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A fundamental goal in ecology is to understand
the factors that determine the distribution and
abundance of organisms. While a vast literature
exists relating factors important in determining
the distribution and movement patterns of
terrestrial and near-shore predators to spatial and
temporal patterns in the distribution of their food
resources, similar studies for pelagic marine
species are rare. This largely stems from the
expense and difficulty of simultaneously measuring
the distribution and abundance of predators, prey,
and related environmental factors in the open ocean
over the larger temporal and spatial scales they
forage. As a result, most studies correlate
distribution and abundance patterns of oceanic
predators to direct or indirect indices of prey
abundance and environmental parameters over small
spatial scales and short time periods. This is
particularly true for studies of the foraging
ecology of large predators.
The blue whale (Balaenoptera musculus) is
the largest organism that has ever existed. This
combined with its mammalian metabolic rate, leads
to the highest average daily total energy
requirements of any species. Blue whales feed
almost exclusively on euphausiids, at rates of up
to 2 metric tons per day (Rice 1978). They satisfy
this high energy demand by feeding in patchily
distributed seasonal but ephemeral aggregations
where euphausiids are found at high densities
(Tomilin 1967, Yochem and Leatherwood 1985,
Schoenherr 1991, Tershy 1992, Croll et al. 1998,
Fiedler et al. 1998). Thus, blue whale foraging
appears to only occur in regions of exceptionally
high productivity. Traditionally, it has been
speculated that blue whale distribution and
movement patterns consist of a seasonal migration
from high latitudes where foraging takes place to
low latitudes where they mate and give birth (e.g.
Mackintosh 1965; Lockyer 1981). However, data from
the Pacific indicate that feeding also takes place
at low latitude, "upwelling-modified" waters
(Reilly and Thayer 1990, Fiedler et al. 1998, Croll
et al. 1998), and data from both the Pacific and
Indian Oceans indicate that some individuals remain
at low latitudes year-round (Yochem and Leatherwood
1985).
Detailed information on the movement patterns of
blue whales off California is not available, but
they are believed to migrate annually between
foraging areas in Central/Southern California
(May-September), the west coast of Baja California
(September-December) and the Gulf of California
(January-April) (Calambokidis 1995). A number of
studies have found that blue whales off California
seasonally forage upon dense euphausiid schools in
highly productive coastal upwelling regions
(Schoenherr 1991, Croll et al. 1998, Forney and
Barlow 1998, Fiedler et al. 1998). Croll et al.
(1998) proposed that: 1) the distribution of blue
whales in the coastal California current region is
defined by their attraction to areas of predictably
high prey density; 2) the preferred prey of these
whales are several species of euphausiids
(Euphausia pacifica , Thysanoessa spinifera, and
Nyctiphanes simplex) that are abundant in the
California Current region; 3) blue whale foraging
efforts are concentrated on dense aggregations
found at discrete depths in the water column; 4)
these localized areas of high euphausiid densities
are predictable and sustained by upwelling regions
of seasonally high primary production; 5)
topographic breaks in the continental shelf located
downstream from these regions work in concert with
euphausiid behavior to collect and maintain large
concentrations of euphausiid swarms; and 6) despite
seasonal and inter-annual variability, these
processes are sufficiently consistent that the
distribution of Balaenoptera whales can be
predicted. In this study, we test these predictions
for California blue whales feeding in a
well-defined foraging area (Monterey Bay,
California) over several years.
Since at least 1986 blue whales have been
reported seasonally foraging in Monterey Bay on
dense aggregations of euphausiids (T.
spinirfera, E. pacifica) (Schoenherr 1991,
Calimbokidis 1995). Due to their high prey demands,
we hypothesized that blue whales in Monterey Bay
forage on dense aggregations of euphausiids that
result from the processes we proposed above. To
test this hypothesis, we used a combination of
concurrent ship- and mooring-based oceanographic
sampling, hydroacoustic sampling, net sampling,
opportunistic cetacean sighting records, visual
surveys, and time-depth recorder deployments to
describe the foraging ecology of whales within the
seasonal upwelling context of Monterey Bay between
1992-1996. Specifically, we: 1) define prey patches
and whale foraging behavior within prey patches, 2)
determine spatial and temporal patterns in the
distribution and abundance of whale prey patches,
and 3) examine physical and biological factors
important in creating whale foraging patches.
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