Humpback Whale Predation and the Case for Top-Down Control of Local
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RESEARCH FEATURE Humpback Whale Predation and the Case for Top-Down Control of Local Herring Populations in the Gulf of Alaska by Ron Heintz, John Moran, Johanna Vollenweider, Jan Straley, Kevin Boswell and Jeep Rice During late winter in Lynn Canal Steller sea lions closely coordinate foraging dives with a humpback whale to capitalize on minor disruptions to the herring school caused by whales. Photo by John Moran. ontrol over the production of highly constrained. Anecdotal evidence of hump- is a goal of the Auke Bay Laboratories’ Cfecund species such as Pacific her- back whales foraging in locations where (ABL) Nutritional Ecology Lab. Beginning ring (Clupea pallasii) has been attributed herring populations are depressed led to in 2001, we began documenting seasonal historically to bottom-up effects such as the hypothesis that humpback whale pre- changes in the energy content of herring as ocean conditions or food availability. In dation impedes the recovery of depressed winter progresses in the local population in contrast, top-down control exists when herring populations, even when the com- Lynn Canal, a fjord adjacent to the ABL fa- predation limits population production. mercial herring fisheries have been closed cility. Winter is an overlooked time of year Recent observations of humpback whales for decades. critical to the survival and production of (Megaptera novaeangliae) foraging on It is important to understand the effect many marine species, and our location near depressed herring populations suggests of humpback whales on herring because a significant herring biomass facilitates our top-down control of herring may be un- both species are conspicuous elements in ability to understand these processes. In derappreciated. Pacific herring popula- the Gulf of Alaska ecosystem. Herring are 2007, our studies expanded when we began tions are depressed in several locations ubiquitously distributed and play a key role comparing the effects of whales on three in Alaska, and humpback whale popula- in the ecosystem by making the energy different herring populations. Our ap- tions are increasing. A 2004-06 census bound in primary consumers available to proach was to estimate the biomass of her- estimated the North Pacific humpback apex predators. Humpback whales are vora- ring consumed by whales in each location whale population at 18,000 to 20,000 and cious predators. A humpback whale weighs and observe herring behavior in response concluded that the population of whales around 30 metric tons (t) and requires the to whale predation. wintering in Hawaiian waters (one half of equivalent of about 1,100 herring per day the North Pacific population) is doubling to meet its average daily metabolic cost. approximately every 15 years. Humpback However, while it is evident that whales de- Herring and whale studies in the Gulf whales could potentially be a significant pend on herring to some degree, the impact of Alaska source of mortality on herring popula- of this dependence on herring is unknown. tions because they are large homeotherms If we are to evolve towards ecosystem- The location of greatest concern for the that often consume herring, and they dis- based management, we will need to begin impacts of whale predation on herring play a remarkable fidelity to their feeding quantifying the benefits whales gain from stocks has been Prince William Sound. In grounds. If whales repeatedly return to lo- herring and the costs of whale predation 1993 the Prince William Sound herring cations to forage with increasing numbers, to herring. Understanding the relationship stock, which had been fished commercially then production of their prey may become between herring and humpback whales for decades, suddenly collapsed due to an AFSC QUARTERLY REPORT 1 RESEARCH FEATURE outbreak of viral hemorrhagic septicemia. The fishery was closed and has essentially remained so to this day. Whale predation has been cited as one possible explanation for the failed recovery. This hypothesis de- rived from reports of humpback whales foraging in Prince William Sound all win- ter in locations where herring were known to congregate. We began examining this question in detail in Prince William Sound in fall 2007 and included Sitka Sound and Lynn Canal (Fig. 1) as reference sites. We included Sitka Sound because it supports a commercially viable herring population with a total bio- mass currently near 85,000 t. Whales have been observed foraging on herring in Sitka Sound since the early 1980s. We included Lynn Canal because the herring population has failed to support a commercial fishery after being closed to commercial fishing for more than a quarter of a century. Though the cause for the depression of the Lynn Canal population is unknown, humpback whale populations there have been increas- ing. We focused on these three areas in winter, a time when herring congregate in each location. These concentrations of her- ring made it easier for us to observe whale foraging behaviors. While there were reports of whales for- aging in winter in Alaskan waters it was unlikely that whales foraged on herring all winter. Humpback whales make transoce- anic migrations from their feeding grounds in the North Pacific to calving grounds near Figure 1. Location of the Prince William Sound (PWS), Lynn Canal, and Sitka Sound study Hawaii in winter. It is known that whales areas in Southeast Alaska (SEAK). stagger their departure from their feeding grounds, suggesting they also stagger their return. This could create the impression that whales were present throughout the entire winter. Only by identifying individ- ual whales and enumerating them would it be possible to establish the extent of win- ter foraging. Individual identification of humpback whales is done by photograph- ing and cataloging the unique markings on the undersides of their flukes (Fig. 2). We used these individual markings in mark-recapture studies to estimate the number of whales present throughout the winter in each of the three locations. We developed monthly estimates of whale abundance for the winters of 2007-08 and 2008-09. We concurrently identified the types of prey consumed by whales through Figure 2. The unique patterns on the flukes of humpback whales identify individuals allowing fatty acid analysis, direct observation, and the use of mark-recapture models to estimate whale abundance. Photo by John Moran. acoustic observation (Fig. 3). Combining 2 OCTOBER NOVEMBER DECEMBER 2010 RESEARCH FEATURE our observations of whale abundance and responded to the time when herring have the proportion of whales consuming her- their highest energy content. ring with bioenergetic models, we esti- In Sitka Sound, the lack of feeding in the mated the biomass of herring consumed early fall on herring was directly related to by whales over winter in each location. We the absence of herring and abundance of compared these estimates to the estimat- krill. In winter, the number of whales ob- ed total biomass of herring present after served eating herring increased after her- spawning for the three populations to de- ring arrived in Sitka Sound. However, as termine what proportion of the total bio- winter progressed herring began staging for mass the whales were likely consuming. The spawning at the same time whales departed herring population numbers were obtained for their breeding grounds. Therefore, even from age-structured stock assessments for though herring were abundant, there were Sitka and Prince William Sounds and esti- few whales present and consumption rates mates of spawning stock biomass for Lynn were low. Canal. All of these herring estimates were In Lynn Canal we observed the oppo- produced by the Alaska Department of Fish site pattern: whales consumed herring in and Game (ADF&G). October and November when whales were Our previous work with Lynn Canal her- abundant and herring densities were just ring indicated comparisons between whale beginning to increase. However, by mid- consumption and spawning stock biomass December when herring density was very would overestimate the whale impact be- high we could no longer determine what Figure 3. In this echogram recorded at 50 kHz a cause there are significantly more herring whales were eating and their numbers were humpback whale can be seen attacking a small present in winter than are observed dur- in steep decline. herring school near the surface. Depth intervals ing spawning. Consequently, we conducted In contrast to Sitka Sound and Lynn are displayed in meters. In January, when this echogram was recorded, herring schools were monthly acoustic surveys in Lynn Canal Canal, whales in Prince William Sound al- dispersed by whales and scattered throughout during winter to estimate herring abun- most always consumed herring and stayed the water column in Prince William Sound. dance and estimate the potential predation until January. Krill appeared to be relatively rate on a monthly basis. One of the benefits rare in the area. of simultaneously conducting the acoustic The differences in whale abundance and mass. Lynn Canal whales ate 500-700 t of and whale abundance surveys was that we apparent preference for herring led to very herring over winter or almost all of the could examine the effects of whales on her- different estimates of herring consumption local spawning stock. But, recall herring ring by mapping the distribution of whales in each location (Table 2). In Sitka Sound, abundance in Lynn Canal during winter and herring and comparing the abundance whales consumed an estimated 800-1,000 exceeds the spawning stock biomass. In of whales with the depth and location of t of herring in late winter. However the November, whales consumed 1%-2% of herring schools in the water column. estimated biomass of herring the previ- the biomass present. After November con- ous spring was about 100,000 t, so whale sumption dropped to less than 1% because consumption amounted to less than 1% of whales departed and herring continued to Estimates of whale predation on the herring biomass.