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Diet energy density estimated from isotopes in predator hair associated with survival, habitat, and population dynamics

Sea ice loss is fundamentally altering the Arctic marine environment. Yet there is a paucity of data on the adaptability of food webs to ecosystem change, including predator-prey interactions. Polar bears ( Ursus maritimus ) are an important subsistence resource for Indigenous people and an apex predator that relies entirely on the under-ice food web to meet their energy needs. Here, we assessed whether polar bears maintained dietary energy density by prey switching in response to spatial-temporal variation in prey availability. We compared the macronutrient composition of diets inferred from stable carbon and nitrogen isotopes in polar bear guard hair (primarily representing summer/fall diet) during periods when bears had low and high survival 2004-2016, between bears that summered on land versus pack ice, and between bears occupying different regions of the Alaskan and Canadian Beaufort Sea. Polar bears consumed diets with lower energy density during periods of low survival suggesting that concurrent increased dietary proportions of beluga whales ( Delphinapterus leucas ) did not offset reduced proportions of ringed seals ( Pusa hispida ). Diets with the lowest energy density and proportions from ringed seal blubber were consumed by bears in the western Beaufort Sea (Alaska) during a period when polar bear abundance declined. Intake required to meet energy requirements of an average free-ranging adult female polar bear was 2.1 kg/day on diets consumed during years with high survival but rose to 3.0 kg/day when survival was low. Although bears that summered onshore in the Alaskan Beaufort Sea had higher fat diets than bears that summered on the pack ice, access to the remains of subsistence-harvested bowhead whales ( Balaena mysticetus ) contributed little to improving diet energy density. Because most bears in this region remain with the sea ice year-round, prey-switching and consumption of whale carcasses onshore appear insufficient to augment diets when availability of their primary prey, ringed seals, is reduced. Our results show that a strong predator-prey relationship between polar bears and ringed seals continues in the Beaufort Sea. The method of estimating dietary blubber using predator hair, demonstrated here, provides a new metric to monitor predator-prey relationships that affect individual health and population demographics.

Ecological Applications↗

Modelling predation by transient leopard seals for an ecosystem-based management of Southern Ocean fisheries

Correctly quantifying the impacts of rare apex marine predators is essential to ecosystem-based approaches to fisheries management, where harvesting must be sustainable for targeted species and their dependent predators. This requires modelling the uncertainty in such processes as predator life history, seasonal abundance and movement, size-based predation, energetic requirements, and prey vulnerability. We combined these uncertainties to evaluate the predatory impact of transient leopard seals on a community of mesopredators (seals and penguins) and their prey at South Georgia, and assess the implications for an ecosystem-based management. The mesopredators are highly dependent on Antarctic krill and icefish, which are targeted by regional fisheries. We used a state-space formulation to combine (1) a mark-recapture open-population model and individual identification data to assess seasonally variable leopard seal arrival and departure dates, numbers, and residency times; (2) a size-based bioenergetic model; and (3) a size-based prey choice model from a diet analysis. Our models indicated that prey choice and consumption reflected seasonal changes in leopard seal population size and structure, size-selective predation and prey vulnerability. A population of 104 (90-125) leopard seals, of which 64% were juveniles, consumed less than 2% of the Antarctic fur seal pup production of the area (50% of total ingested energy, IE), but ca. 12-16% of the local gentoo penguin population (20% IE). Antarctic krill (28% IE) were the only observed food of leopard seal pups and supplemented the diet of older individuals. Direct impacts on krill and fish were negligible, but the "escapement" due to leopard seal predation on fur seal pups and penguins could be significant for the mackerel icefish fishery at South Georgia. These results suggest that: (1) rare apex predators like leopard seals may control, and may depend on, populations of mesopredators dependent on prey species targeted by fisheries; and (2) predatory impacts and community control may vary throughout the predator's geographic range, and differ across ecosystems and management areas, depending on the seasonal abundance of the prey and the predator's dispersal movements. This understanding is important to integrate the predator needs as natural mortality of its prey in models to set prey catch limits for fisheries. Reliable estimates of the variability of these needs are essential for a precautionary interpretation in the context of an ecosystem-based management. ?? 2009 Elsevier B.V.

Ecological Modelling↗

Numerical simulation of the effects of groundwater withdrawal and injection of high-salinity water on salinity and groundwater discharge, Kaloko-Honokōhau National Historical Park, Hawaiʻi

Kaloko-Honokōhau National Historical Park (KAHO) is located on the west coast of the island of Hawaiʻi and contains water resources exposed in fishponds, anchialine pools, and marine waters that are cultural resources and that provide habitat for threatened, endangered, and other culturally important native species. KAHO’s water resources are sustained by and dependent on groundwater discharge. In 1978, the year of KAHO authorization, the lands immediately surrounding KAHO were undeveloped and zoned for conservation purposes; at present, most surrounding lands are either developed or zoned for industrial, commercial, or residential use. Urbanization of the North Kona District has increased the need for additional drinking and nonpotable (irrigation) water. Because KAHO’s water resources may be affected by existing and proposed groundwater withdrawals and injections of high-salinity water in the surrounding area, the U.S. Geological Survey, in cooperation with the National Park Service, undertook this study to refine the understanding of how groundwater withdrawals and injection of high-salinity water may affect KAHO’s water resources. Changes in KAHO water resources, in terms of changes in salinity and groundwater discharge, were modeled for selected scenarios of groundwater withdrawal and high-salinity water injection in the aquifer. The numerical model was developed using the model code SUTRA, which accounts for density-dependent flow and salinity transport, and included the coastal-confined groundwater system beneath the coastal freshwater-lens system. Model results indicate that withdrawal of additional groundwater from the coastal freshwater-lens system will affect the salinity of KAHO’s anchialine pools, which provide habitat for the endangered orangeblack Hawaiian damselfly ( Megalagrion xanthomelas ). The magnitude of the effect is dependent on the amount and location of the withdrawal. Greater withdrawal rates cause greater increases in salinity in KAHO, other factors being equal. For a given withdrawal rate, the greatest increase in salinity in KAHO is associated with wells withdrawing groundwater in an area inland of KAHO and the least increase in salinity is associated with wells near the coast. Model results also indicate that withdrawal of additional groundwater from the coastal freshwater-lens system will affect the groundwater discharge, in terms of the freshwater component (water with zero salinity) of the discharge, through KAHO. Greater withdrawal rates cause greater reductions in freshwater discharge through KAHO. For a given withdrawal rate, the greatest reduction in freshwater discharge through KAHO is associated with wells near the north boundary of KAHO and the least reduction is associated with wells near the coast to the north and south of KAHO. Injection of high-salinity water that is denser than ocean water can affect the salinity of damselfly habitat in KAHO, with the magnitude of the effect dependent on the location of the injection. Model results indicate that salinity may either increase or decrease in the anchialine pools that provide damselfly habitat in KAHO, depending on the site of injection. Injection inland of KAHO and at sites immediately north and south of KAHO causes a simulated decrease in salinity at the damselfly habitat, whereas injection farther north and south of KAHO causes an increase in salinity. Injection of high-salinity water also causes a reduction in freshwater discharge through KAHO, with the greatest reduction associated with distant injection wells to the north and south of KAHO and the least reduction associated with wells located near and immediately inland from KAHO. The numerical groundwater models developed for this study have a number of limitations. Lack of understanding of the subsurface geology constrains the ability to accurately model the groundwater-flow system. The models developed for this study are nonunique, cannot account for local-scale heterogeneities in the aquifer, and contain uncertainties related to recharge, boundary conditions, assigned parameter values in the model, and representations of the different hydrogeological features. Confidence in model results can be improved by addressing these and other limitations. In spite of these limitations, the three-dimensional numerical model developed for this study provides a useful conceptual understanding of the potential effects of additional withdrawals and injections on groundwater resources in KAHO. Further evaluation of the ecologic effects of the simulated changes in groundwater quality and quantity in KAHO is needed but is beyond the scope of this study.

Hawaii↗

Sea otter studies in Glacier Bay National Park and Preserve: annual report 2002

Since 1995, the number of sea otters in Glacier Bay proper has increased from around 5 to more than 1200. Sea otter distribution is mostly limited to the Lower Bay, south of Sandy Cove, and is not continuous within that area. Concentrations occur in the vicinity of Sita Reef and Boulder Island and between Pt. Carolus and Rush Pt. on the west side of the Bay, although there have been occasional sightings north of Sandy Cove (Figure 1). Large portions of the Bay remain unoccupied by sea otters, but recolonization is occurring rapidly. Most prey recovered by sea otters in Glacier Bay are ecologically, commercially, or socially important species. In 2002 sea otter diet consisted of 35% clam, 26% mussel, 3% crab, 3.0% snail, 2% starfish, 11% urchins, 2% other, and 20% unidentified. Dominant clam species include the butter clam, Saxidomus gigantea , the Greenland cockle, Serripes groenlandicus , and the littleneck clam, Protothaca staminea . Urchins are primarily green urchins, Strongylocentrotus droebachiensis , and the mussel is Modiolus modiolus . Crabs observed in 2002 include the Dungeness, Cancer magister , the kelp crab Pugettia gracilis , and the helmet crab, Telmessus cherigonus . Although we characterize diet at broad geographic scales, we have previously found diet to vary between sites separated by as little as several hundred meters. Dietary variation among and within sites can reflect differences in prey availability as well as individual specialization. We estimated species composition, density, biomass, and sizes of subtidal clams, urchins, and mussels at 13 sites in Glacier Bay and 5 sites in nearby Port Althorp, where sea otters have been present for at least 20 years. All sites were selected based on the presence of abundant clam siphons and the absence of sea otters (Glacier Bay) or abundant shell litter and the presence of sea otters (Port Althorp). Glacier Bay sites were selected to achieve a broad geographic sample of dense subtidal clam beds within Glacier Bay prior to occupation and foraging by sea otters. Port Althorp sites were chosen to achieve a representative sample of subtidal clam beds already under prolonged foraging pressure by sea otters. There was no direct evidence of otter foraging at any of our Glacier Bay sampling sites. In Glacier Bay, we sampled 15,338 bivalves (average of 1,180/site) representing 14 species of clam, 2 species of mussel, and a single scallop and we sampled 6,917 urchins (average of 513/site). In Port Althorp, we sampled 1,034 bivalves (average of 207/site) representing 14 species of clam. We found only 5 urchins, all S. droebachiensis . Mean densities and biomass of all subtidal clams were significantly greater in Glacier Bay (59.2 and 99/0.25m 2 compared to Port Althorp (10.3 and 5.8/0.25m 2 (p<0.002 for both). Our contrasts of subtidal clam populations between Glacier Bay and Port Althorp suggest that clam densities will likely decline by about a factor of six and that clam biomass estimates will decline by more than a factor of ten. Numerically dominant species of clams, P. staminea , S. gigantea , Macoma sp. and Mya sp. were all significantly greater in density and biomass in Glacier Bay, while C. nutalli density was low but significantly higher in Port Althorp. Subtidal clam species diversity was significantly greater in Port Althorp compared to Glacier Bay, although this may simply reflect habitat differences. Sea urchin densities were high in Glacier Bay, while in Port Althorp urchins were virtually absent. Sea otters are now well established in limited areas of the lower portions of Glacier Bay. It is likely that distribution and numbers of sea otters will continue to increase in Glacier Bay in the near future. Glacier Bay supports large and diverse populations of clams that are largely unexploited by sea otters at present. It is predictable that the density and sizes of clam populations will decline in response to otter predation. This will result in fewer opportunities for human harvest, but will also trigger ecosystem level changes, as prey for other predators, such as octopus, sea stars, fishes, birds and mammals are modified. Sea otters will also modify benthic habitats through excavation of sediments required to extract burrowing infauna such as clams. Effects of sediment disturbance by foraging sea otters are not understood. Glacier Bay also supports large populations of other preferred sea otter prey, such as king ( Paralithodes sp. ), tanner ( Chionoecetes sp. ) and dungeness ( Cancer magister ) crabs and green sea urchins ( S. droebachiensis ). As the colonization of Park waters by sea otters continues, it is also likely that dramatic changes will occur in the species composition, abundance, and size class distribution of many components of the nearshore marine ecosystem. Many of the changes will occur as a direct result of predation by sea otters. Others will result from indirect or cascading effects of sea otter foraging, such as increased kelp production and modified prey availability for other nearshore predators. Without recognizing and quantifying the extent of change initiated by the colonization of Glacier Bay by sea otters, management of nearshore resources will be severely constrained for many decades.

Alaska↗

Near-field receiving-water monitoring of trace metals and a benthic community near the Palo Alto Regional Water Quality Control Plant in south San Francisco Bay, California—2020

Trace-metal concentrations in sediment and in the clam Limecola petalum (World Register of Marine Species, 2020; formerly reported as Macoma balthica and M. petalum ), clam reproductive activity, and benthic macroinvertebrate community structure were investigated in a mudflat 1 kilometer (km) south of the discharge of the Palo Alto Regional Water Quality Control Plant (PARWQCP) in south San Francisco Bay, California. This report includes the data collected by the U.S. Geological Survey (USGS) for January 2020–December 2020 (Cain and others, 2022). These data append to long-term datasets extending back to 1974. A major focus of the report is an integrated description of the 2020 data within the context of the longer, multidecadal dataset. This dataset supports the City of Palo Alto’s Near-Field Receiving- Water Monitoring Program, initiated in 1994. Silver and copper contamination substantially decreased at the site in the 1980s following the implementation by PARWQCP of advanced wastewater-treatment and source-control measures. Since the 1990s, concentrations of these elements in surface sediments have continued to decrease, although more slowly. For example, from 1994 to 2020, the minimum annual mean silver concentration—0.20 milligram per kilogram (mg/kg)—was observed in multiple years. In 2020, silver concentrations ranged from 0.18 to 0.28 mg/kg. These concentrations are 2 to 3 times higher than the regional background concentration. Presently (2020), sediment-copper concentrations appear to be near the regional background level. Over the same period (1994–2020), sedimentary iron and zinc exhibited modest decreases. Sedimentary aluminum, chromium, mercury, nickel, and selenium have not exhibited any trend. Since 1994, silver and copper concentrations in L. petalum have varied seasonally, apparently in response to a combination of site-specific metal exposures and cyclic growth and reproduction, as reported previously. Seasonal patterns for other elements, including chromium, mercury, nickel, selenium, and zinc, generally were similar in timing and magnitude as those for silver and copper. Downward trends in the silver and zinc concentrations in L. petalum during 1994–2020 were evident and appeared to be related to the general physiological condition of the clam, indicated by a condition index. Biological effects of elevated silver and copper contamination at the Palo Alto site have been interpreted from data collected during and after the recession of these contaminants. Concentrations of both elements in the soft tissues of L. petalum decreased with sedimentary copper and silver. This pattern was associated with changes in the reproductive activity of L. petalum , as well as the structure of the benthic invertebrate community. Reproductive activity of L. petalum increased as metal concentrations in L. petalum decreased (Hornberger and others, 2000), and presently is stable with almost all animals initiating reproduction in the fall and spawning the following spring. Analyses of the benthic community structure indicate that the infaunal invertebrate community has shifted from one dominated by several opportunistic species when silver and copper exposures were highest to one in which the species abundance is more evenly distributed, a pattern that indicates a more stable community that is subjected to fewer stressors. Importantly, this long-term change is unrelated to other metals and other measured environmental factors, including salinity and sediment composition. In addition, two of the opportunistic species ( Ampelisca abdita and Streblospio benedicti ) that brood their young and live on the surface of the sediment in tubes have shown a continual decrease in dominance coincident with the decrease in metals. Both species had short-lived rebounds in abundance in 2008, 2009, and 2010 and showed signs of increasing abundance in 2020. Heteromastus filiformis (a subsurface polychaete worm that lives in the sediment, consumes sediment and organic particles residing in the sediment, and reproduces by laying its eggs on or in the sediment) showed a concurrent increase in dominance and, in the last several years before 2008, showed a stable population. H. filiformis abundance increased slightly from 2011 to 2012 and returned to pre-2011 numbers in 2020. The reproductive mode of most species that were present in 2020 was indicative of species that were capable of movement either as pelagic larvae or as mobile adults. Although oviparous species were lower in number in this group, the authors hypothesize that these species will return slowly as more species move back into the area. The use of functional ecology was highlighted in the 2020 benthic community data, which showed that the animals that have now returned to the mudflat are those that can respond successfully to a physical, nontoxic disturbance. Today, community data show a mix of species that consume the sediment, or filter feed, those that have pelagic larvae that must survive landing on the sediment, and those that brood their young. The long-term recovery observed after the 1970s can be ascribed to the decrease in sediment pollutants.

California↗

Nitrogen excess in North American ecosystems: Predisposing factors, ecosystem responses, and management strategies

Most forests in North America remain nitrogen limited, although recent studies have identified forested areas that exhibit symptoms of N excess, analogous to overfertilization of arable land. Nitrogen excess in watersheds is detrimental because of disruptions in plant/soil nutrient relations, increased soil acidification and aluminum mobility, increased emissions of nitrogenous greenhouse gases from soil, reduced methane consumption in soil, decreased water quality, toxic effects on freshwater biota, and eutrophication of coastal marine waters. Elevated nitrate ( NO 3 − ) loss to groundwater or surface waters is the primary symptom of N excess. Additional symptoms include increasing N concentrations and higher N:nutrient ratios in foliage (i.e., N:Mg, N:P), foliar accumulation of amino acids or NO 3 − , and low soil C:N ratios. Recent nitrogen-fertilization studies in New England and Europe provide preliminary evidence that some forests receiving chronic N inputs may decline in productivity and experience greater mortality. Long-term fertilization at Mount Ascutney, Vermont, suggests that declining and slow N-cycling coniferous stands may be replaced by fast-growing and fast N-cycling deciduous forests. Symptoms of N saturation are particularly severe in high-elevation, nonaggrading spruce–fir ecosystems in the Appalachian Mountains and in eastern hardwood watersheds at the Fernow Experimental Forest near Parsons, West Virginia. In the Los Angeles Air Basin, mixed conifer forests and chaparral watersheds with high smog exposure are N saturated and exhibit the highest streamwater NO 3 − concentrations for wildlands in North America. High-elevation alpine watersheds in the Colorado Front Range and a deciduous forest in Ontario, Canada, are N saturated, although N deposition is moderate (∼8 kg·ha −1 ·yr −1 ). In contrast, the Harvard Forest hardwood stand in Massachusetts has absorbed >900 kg N/ha during 8 yr of N amendment studies without significant NO 3 − leaching, illustrating that ecosystems vary widely in the capacity to retain N inputs. Overly mature forests with high N deposition, high soil N stores, and low soil C:N ratios are prone to N saturation and NO 3 − leaching. Additional characteristics favoring low N retention capacity include a short growing season (reduced plant N demand) and reduced contact time between drainage water and soil (i.e., porous coarse-textured soils, exposed bedrock or talus). Temporal patterns of hydrologic fluxes interact with biotic uptake and internal cycling patterns in determining ecosystem N retention. Soils are the largest storage pool for N inputs, although vegetation uptake is also important. Recent studies indicate that nitrification may be widespread in undisturbed ecosystems, and that microbial assimilation of NO 3 − may be a significant N retention mechanism, contrary to previous assumptions. Further studies are needed to elucidate the sites, forms, and mechanisms of N retention and incorporation into soil organic matter, and to test potential management options for mitigating N losses from forests. Implementation of intensive management practices in N-saturated ecosystems may only be feasible in high-priority areas and on a limited scale. Reduction of N emissions would be a preferable solution, although major reductions in the near future are unlikely in many areas due to economic, energy-use, policy, and demographic considerations.

Ecological Applications↗