George M. Watters

Variability in krill biomass links harvesting and climate warming to penguin population changes in Antarctica

The West Antarctic Peninsula (WAP) and adjacent Scotia Sea support abundant wildlife populations, many of which were nearly extirpated by humans. This region is also among the fastest-warming areas on the planet, with 5–6 °C increases in mean winter air temperatures and associated decreases in winter sea-ice cover. These biological and physical perturbations have affected the ecosystem profoundly. One hypothesis guiding ecological interpretations of changes in top predator populations in this region, the “sea-ice hypothesis,” proposes that reductions in winter sea ice have led directly to declines in “ice-loving” species by decreasing their winter habitat, while populations of “ice-avoiding” species have increased. However, 30 y of field studies and recent surveys of penguins throughout the WAP and Scotia Sea demonstrate this mechanism is not controlling penguin populations; populations of both ice-loving Adélie and ice-avoiding chinstrap penguins have declined significantly. We argue in favor of an alternative, more robust hypothesis that attributes both increases and decreases in penguin populations to changes in the abundance of their main prey, Antarctic krill. Unlike many other predators in this region, Adélie and chinstrap penguins were never directly harvested by man; thus, their population trajectories track the impacts of biological and environmental changes in this ecosystem. Linking trends in penguin abundance with trends in krill biomass explains why populations of Adélie and chinstrap penguins increased after competitors (fur seals, baleen whales, and some fishes) were nearly extirpated in the 19th to mid-20th centuries and currently are decreasing in response to climate change.

Visualizing the Food-Web Effects of Fishing for Tunas in the Pacific Ocean

We use food-web models to develop visualizations to compare and evaluate the interactions of tuna fisheries with their supporting food webs in the eastern tropical Pacific (ETP) and the central north Pacific (CNP) Oceans. In the ETP and CNP models, individual fisheries use slightly different food webs that are defined by the assemblage of targeted tuna species. Distinct energy pathways are required to support different tuna species and, consequently, the specific fisheries that target different tuna assemblages. These simulations suggest that catches of tunas, sharks, and billfishes have lowered the biomass of the upper trophic levels in both systems, whereas increases in intermediate and lower trophic level animals have accompanied the decline of top predators. Trade- offs between fishing and predation mortality rates that occur when multiple fisheries interact with their respective food webs may lead to smaller changes in biomass than if only the effect of a single fishery is considered. Historical simulations and hypothetical management scenarios further demonstrate that the effects of longline and purse seine fisheries have been strongest in upper trophic levels, but that lower trophic levels may respond more strongly to purse-seine fisheries. The apex predator guild has responded most strongly to longlining. Simulations of alternative management strategies that attempt to rebuild shark and billfish populations in each ecosystem reveal that (1) changes in longlining more effectively recover top predator populations than do changes in purse seining and (2) restrictions on both shallow-set longline gear and shark finning may do more to recover top predators than do simple reductions in fishing effort.

Range expansion of the Humboldt squid was not caused by tuna fishing

Zeidberg and Robison (1) infer that the range expansion of Dosidicus gigas resulted from a top-down cascade caused by fishing for tunas and billfishes in the eastern equatorial Pacific (EEP). We offer an alternative perspective supported on three grounds. First, the thermal tolerance and plastic life history of D. gigas …

Complex wasp-waist regulation of pelagic ecosystems in the Pacific Ocean

Abstract‘Wasp-waist’ control of marine ecosystems is driven by a combination of top-down and bottom-up forcing by a few abundant short-lived species occupying intermediate trophic levels that form a narrow ‘waist’ through which energy flow from low to high trophic levels is controlled. It has been assumed that wasp-waist control occurs primarily in highly productive and species-poor systems (e.g. upwelling regions). Two large, species-rich, pelagic ecosystems in the relatively oligotrophic eastern and western Pacific Ocean also show wasp-waist-like structure, in that short-lived and fast-growing cephalopods and fishes at intermediate trophic levels comprise the vast majority of the biomass. Possible forcing dynamics of these systems were examined using ecosystem models by altering the biomass of phytoplankton (bottom-up forcing), large pelagic predators (top-down forcing), and intermediate ‘wasp-waist’ functional groups independently and observing how these changes propagated throughout the ecosystem. The largest effects were seen when altering the biomass of mid trophic-level epipelagic and mesopelagic fishes, where dramatic trophic cascades occurred both upward and downward in the system. We conclude that the high productivity and standing biomass of animals at intermediate trophic levels has a strong top-down influence on the abundance of primary producers. Furthermore, their importance as prey for large predators results in bottom-up controls on populations at higher trophic levels. We show that these tropical pelagic ecosystems possess a complex structure whereby several waist groups and alternate trophic pathways from primary producers to apex predators can cause unpredictable effects when the biomasses of particular functional groups are altered. Such models highlight the possible structuring mechanisms in pelagic systems, which have implications for fisheries that exploit these wasp-waist groups, such as squid fisheries, as well as for fisheries of top predators such as tunas and billfishes that prey upon wasp-waist species.

Identifying Risk: Concurrent Overlap of the Antarctic Krill Fishery with Krill-Dependent Predators in the Scotia Sea

Mitigating direct and indirect interactions between marine predators and fisheries is a motivating factor for ecosystem-based fisheries management (EBFM), especially where predators and fisheries compete for a shared resource. One difficulty in advancing EBFM is parameterizing clear functional responses of predators to indices of prey availability. Alternative characterizations of fishery-predator interactions may therefore benefit the implementation of EBFM. Telemetry data identify foraging areas used by predators and, therefore, represent critical information to mitigate potential competition between predators and fisheries. We analyzed six years (2009–2014) of telemetry data collected at Cape Shirreff, Livingston Island and Admiralty Bay, King George Island, Antarctica, on three species of Pygoscelid penguins and female Antarctic fur seals. In this region, all four species are primarily dependent on Antarctic krill. The tracking data demonstrate local movements near breeding colonies during the austral summer and dispersal from breeding colonies during the winter. We then assessed overlap between predators and the Antarctic krill fishery on a suite of spatiotemporal scales to examine how different data aggregations affect the extent and location of overlap. Concurrent overlap was observed on all spatiotemporal scales considered throughout the Antarctic Peninsula and South Orkney Islands region, including near tagging locations and in distant areas where recent fishing activity has concentrated. Overlap occurred at depths where mean krill densities were relatively high. Our results demonstrate that direct overlap of krill-dependent predators with the krill fishery on small spatiotemporal scales is relatively common throughout the Antarctic Peninsula region. As the krill fishery continues to develop and efforts to implement ecosystem-based management mature, indices of overlap may provide a useful metric for indicating where the risks of fishing are highest. A precautionary approach to allocating krill catches in space would be to avoid large increases in catch where overlap on small spatiotemporal scales is common.

Impacts of rising sea temperature on krill increase risks for predators in the Scotia Sea

Climate change is a threat to marine ecosystems and the services they provide, and reducing fishing pressure is one option for mitigating the overall consequences for marine biota. We used a minimally realistic ecosystem model to examine how projected effects of ocean warming on the growth of Antarctic krill, Euphausia superba, might affect populations of krill and dependent predators (whales, penguins, seals, and fish) in the Scotia Sea. We also investigated the potential to mitigate depletion risk for predators by curtailing krill fishing at different points in the 21st century. The projected effects of ocean warming on krill biomass were strongest in the northern Scotia Sea, with a ≥40% decline in the mass of individual krill. Projections also suggest a 25% chance that krill biomass will fall below an established depletion threshold (75% of its unimpacted level), with consequent risks for some predator populations, especially penguins. Average penguin abundance declined by up to 30% of its unimpacted level, with up to a 50% chance of falling below the depletion threshold. Simulated krill fishing at currently permitted harvest rates further increased risks for depletion, and stopping fishing offset the increased risks associated with ocean warming in our model to some extent. These results varied by location and species group. Risk reductions at smaller spatial scales also differed from those at the regional level, which suggests that some predator populations may be more vulnerable than others to future changes in krill biomass. However, impacts on predators did not always map directly to those for krill. Our findings indicate the importance of identifying vulnerable marine populations and targeting protection measures at appropriate spatial scales, and the potential for spatially-structured management to avoid aggravating risks associated with rising ocean temperatures. This may help balance tradeoffs among marine ecosystem services in an uncertain future.