Please feel
free to contact me if you would like a reprint of any of these papers.
J.L. Miksis-Olds, P.J. Stabeno, J.M. Napp, A.I. Pinchuk, J.A. Nystuen,
J.D. Warren, S.L. Denes. 2013.
Ecosystem response to a temporary sea ice retreat in the Bering Sea: Winter 2009. Progress in Oceanography. 111: 38-51. doi: 10.1016/j.pocean.2012.10.010. [
.pdf]
Adding acoustic systems onto ocean moorings and observatories provides
additional data to more fully document ecosystem responses to
environmental perturbations. A passive acoustic recorder and
three-frequency echosounder system were integrated into a biophysical
mooring on the central eastern Bering Sea continental shelf. An
unexpected, transient, mid-winter retreat of the seasonal sea ice was
observed over the mooring for a 2-week period in March 2009.
Interpretation of the passive acoustic data provided information about
sea ice conditions and included the detection and identification of
vocalizing marine mammals, while the acoustic backscatter provided
information on relative zooplankton and fish abundance before, during,
and after the retreat. Hydrographic data confirmed the acoustic signal
was associated with changing surface ice conditions, and the combined
information from the biophysical mooring sensors revealed changes in
winter trophic level dynamics during the retreat, which would have
otherwise been undetected by traditional ship-based observations.
Changes in the acoustic environment, zooplankton dynamics, and acoustic
detection of marine mammals were observed amidst a physically stable
and uniform water column with no indication of a phytoplankton bloom.
These data demonstrate the value of acoustic technologies to monitor
changing ecosystems dynamics in remote and hazardous locations.
J.D. Warren. 2012.
Counting critters in the sea using active acoustics. Acoustics Today. Volume 8. Issue 3. 25-34.
doi: 10.1121/1.4753914 [
.pdf]
An invited paper for the lay-language magazine for the Acoustical
Society of America as part of a special issue on Remote Sensing of
Animals.
Joy N. Smith, P.H. Ressler,
J.D. Warren. 2013.
A distorted wave Born approximation target strength model for Bering Sea euphausiids. ICES Journal of Marine Science. 70(1): 204-214. doi: 10.1093/icesjms/fss140 [
.pdf]
Acoustic surveys monitor euphausiid populations in the Bering Sea
because of their importance as prey for walleye pollock and other
organisms. Various scattering models exist to convert acoustic
backscatter data to estimates of euphausiid numerical density or
biomass, but a target strength (TS) model specific to Bering Sea
euphausiids has not been available. This study parameterized a
distorted wave Born approximation (DWBA) scattering model using
physical (length and body shape) and material (density contrast, g, and
sound speed contrast, h) properties measured from live euphausiids. All
model parameters (length, shape, material properties, orientation) were
evaluated for their effect on predicted TS. A polynomial function was
used to describe animal shape and produced smaller TS estimates
compared to a taper function, as is traditionally used in DWBA
scattering models of euphausiids. Animal length was positively
correlated with TS, but variations in other parameters (including
material properties and orientation) also produced large changes in TS.
Large differences in TS between estimates calculated using measured
versus literature material property values caused large variations in
acoustic estimates of euphausiid numerical densities (animals m23)
which emphasizes the importance of collecting site-specific g and h
measurements when possible.
P.H. Ressler, A. De Robertis,
J.D. Warren, Joy N. Smith, S. Kotwicki. 2012.
Developing an acoustic survey of euphausiids to understand trophic interactions in the Bering Sea ecosystem. Deep Sea Research II. 65-70: 184-195. doi: 10.1016/j.dsr2.2012.02.015 [
.pdf]
Euphausiids (principally Thysanoessa spp.) are a key group of organisms
in the Bering Sea ecosystem, linking production at lower trophic levels
to top predators and important commercial fish stocks such as
walleye pollock (Theragra chalcogramma). Here, we combine
multifrequency acoustic survey methods, physics-based models of
euphausiid backscatter, and net sampling to provide a means of
monitoring the status and trends of euphausiid standing stock biomass
on the Bering Sea shelf. Observations made using this approach during
six summers (2004 and 2006–2010) indicate that standing stocks of
euphausiids and pollock were inversely correlated over time as well as
in space across the continental shelf. First-order calculations show
that when pollock abundance was at its peak during these years, the
pollock stock could have consumed 10–87% of the euphausiid standing
stock between May and September. We hypothesize that predation by
pollock is a significant top-down control on euphausiid standing stock
in this system.
S.E. Parks,
J.D. Warren, K. Stamieszkin, C.A. Mayo, D. Wiley. 2012.
Dangerous dining: surface foraging of North Atlantic right whales increases risk of vessel collisions. Biology Letters. 8(1): 57-60. doi: 10.1098/rsbl.2011.0578 [
.pdf]
North Atlantic right whales are critically endangered and, despite
international protection from whaling, significant numbers die from
collisions with ships. Large groups of right whales migrate to the
coastal waters of New England during the late winter and early spring
to feed in an area with large numbers of vessels. North Atlantic right
whales have the largest per capita record of vessel strikes of any
large whale population in the world. Right whale feeding behaviour in
Cape Cod Bay (CCB) probably contributes to risk of collisions with
ships. In this study, feeding right whales tagged with archival suction
cup tags spent the majority of their time just below the water’s
surface where they cannot be seen but are shallow enough to be
vulnerable to ship strike. Habitat surveys show that large patches of
right whale prey are common in the upper 5 m of the water column in CCB
during spring. These results indicate that the typical spring-time
foraging ecology of right whales may contribute to their high level of
mortality from vessel collisions. The results of this study suggest
that remote acoustic detection of prey aggregations may be a useful
supplement to the management and conservation of right whales.
J.N. Smith, P.H. Ressler, and J.D. Warren. 2010.
Material properties of euphausiids and other zooplankton from the Bering Sea. Journal of the Acoustical Society of America. 128(5): 2664-2680. doi: 10.1121/1.3488673 [
.pdf]
Acoustic assessment of Bering Sea euphausiids and their predators can
provide useful data for ecosystem studies if the acoustic scattering
characteristics of these animals are known. The amount of acoustic
energy that is scattered by different marine zooplankton taxa is
strongly affected by the contrast of the animal’s density g and sound
speed h with the surrounding seawater. Density and sound speed contrast
were measured in the Bering Sea during the summer of 2008 for several
different zooplankton and nekton taxa including: euphausiids
(Thysanoessa inermis, Thysanoessa raschii, and Thysanoessa spinifera),
copepods, amphipods, chaetognaths, gastropods, fish larvae, jellyfish,
and squid. Density contrast values varied between different taxa as
well as between individual animals within the same species. Sound speed
contrast was measured for monospecific groups of animals and
differences were found among taxa. The range, mean, and standard
deviation of g and h for all euphausiid species were: g = 1.001– 1.041;
1.018; 0.009 and h = 0.990– 1.017; 1.006; 0.008. Changes in the
relationship between euphausiid material properties and animal length,
seawater temperature, seawater density, and geographic location were
also evaluated. Results suggest that environmental conditions at
different sample locations led to significant differences in animal
density and material properties.
J.D. Warren and D.A. Demer.
2010.
Abundance and distribution of Antarctic krill
(Euphausia superba) nearshore of Cape Shirreff, Livingston Island,
Antarctica, during six austral summers between 2000 and 2007. Canadian Journal of Fisheries and Aquatic
Sciences. 67(7): 1159-1170. doi:10.1139/F10-042 [
.pdf]
Abundance
and distribution of Antarctic krill (Euphausia superba) in the
nearshore waters north of Livingston Island, Antarctica, were
characterized from six small-boat surveys conducted in late January or
early February from 2000 to 2007. The first three surveys (2000, 2002,
2004) were conducted using a 120 kHz split-beam echosounder to measure
water column acoustic backscatter. The last three surveys (2005–2007)
were conducted using 38 kHz and 200 kHz single-beam echosounders. A
portion of the acoustic backscatter was attributed to Antarctic krill
based on the results of net tows, underwater video observations, and a
multiple-frequency acoustic classification algorithm. The annual mean
krill biomass density in the survey area ranged from 11 to 84 g·m–2.
Results are compared with the western Scotia Sea area of the US
Antarctic Marine Living Resources (AMLR) program’s acoustic surveys of
krill biomass density for the same years. Nearshore krill biomass
densities were significantly larger (t test, p < 0.05), more stable,
and the coefficients of variation were smaller than the much larger
AMLR surveys. Increased competition between seals, penguins, and humans
for the nearshore krill resource, especially during the austral summer
months, could impact the recruitment success of these land-based krill
predators. Implications of nearshore krill biomass on small-scale
management units are discussed.
M.J. Cox, J.D. Warren, D.A.
Demer, G.R. Cutter, and A.S. Brierley. 2010.
Three-dimensional
observations of swarms of Antarctic krill (Euphausia superba) made using a
multi-beam echosounder. Deep Sea Research II.
doi:10.1016/j.dsr2.2009.10.003 [
.pdf]
Antarctic krill (Euphausia superba) aggregate in dense swarms. Previous
investigations of krill swarms
have used conventional single- or split-beam echosounders that, with
post-processing, provide a two-
dimensional (2-D) view of the water column, leaving the third dimension
to be inferred. We used a
multi-beam echosounder system (SM20, 200kHz, Kongsberg Mesotech Ltd,
Canada) from an inflatable
boat (length=5.5m) to sample water-column backscatter, particularly
krill swarms, directly in 2-D and,
with post-processing, to provide a three dimensional (3-D) view of
entire krill swarms. The study took
place over six days(2–8 February 2006) in the vicinity of Livingston
Island, South Shetland Islands,
Antarctica (62.41S, 60.71W). An automatic 3-D aggregation detection
algorithm resolved 1006 krill
swarms from the survey data. Principal component analyses indicated
that swarm morphology metrics
such as length, surface area and volume accounted for the largest
between swarm variance, followed by
echo energy, and finally swarm geographic location. Swarms did not form
basic cylindrical or spherical
shapes, but had quite consistent surface area to volume ratios of 3.3
m. Swarms were spatially
segregated, with larger sizes (mean north-south length = 276 m, at
least double that of two other swarm
classifications), found to the northwest of the survey area.The
apparent clustering of swarm types
suggests that krill biomass surveys and ecosystem investigations may
require stratified survey design,
in response to varying 3-D swarm morphology, variation that may be
driven in turn by environmental
characteristics such as bathymetry.
K.A. Forman and J.D. Warren.
2009.
Variability
in the density and sound-speed of coastal zooplankton and nekton.
ICES Journal of Marine Science. doi:
10.1093/icesjms/fsp217 [
.pdf]
Acoustic sampling techniques provide an advantage over traditional
net-sampling by increasing scientific ability to survey a large area in
a relatively short period, as well as providing higher-resolution data
in the vertical and horizontal dimensions. To convert acoustic data
into measures of biological organisms, physics-based scattering models
are often used. Such models use several parameters to predict the
amount of sound scattered by a fluid-like or weakly scattering animal.
Two important input parameters are the density (g) and sound-speed (h)
contrasts of the animal and the surrounding seawater. The density and
sound-speed contrasts were measured for coastal zooplankton and nekton
species including shrimps (Palaemonetes pugio and Crangon
septemspinosa), fish (Fundulus majalis and Fundulus heteroclitus), and
polychaetes (Nereis succinea and Glycera americana) along with multiple
physiological and environmental variables. Factors such as animal size,
feeding status, fecundity, gender, and maturity caused variations in g.
The variations in g observed for these animals could lead to large
differences (or uncertainties) in abundance estimates based on acoustic
scattering models and field-collected backscatter data. It may be
important to use a range of values for g and h in the acoustic
scattering models used to convert acoustic data into estimates of the
abundance of marine organisms.
M.J. Cox, D.A. Demer, J.D.
Warren, G.R. Cutter, and A.S. Brierley. 2009.
Multibeam
echosounder observations reveal interactions between Antarctic krill
and air-breathing predators .
Marine Ecology Progress Series 378: 199-209. doi: 10.3354/meps07795 [
.pdf]
A multibeam echosounder (MBE) was deployed on an inflatable boat
(length = 5.5 m) to observe swarms of Antarctic krill
Euphausia superba
in the nearshore environment off Livingston Island, South Shetland
Islands, Antarctica. Visual observations of air-breathing predators,
including penguins and fur seals, were made from the boat at the same
time. MBEs extend the 2-dimensional acoustic observations that can be
made with conventional vertical echosounders to 3 dimensions, enabling
direct observation of the surface areas and volumes of entire krill
swarms. Krill swarms exhibited a wide range of various size metrics
(e.g. height, length and width) but only a narrow range of
surface-area-to-volume ratios or ‘roughnesses’, suggesting that krill
adopt a consistent group behavior to maintain swarm shape. The
variation in R was investigated using generalized additive models
(GAMs). GAMs indicated that the presence of air-breathing predators
influenced swarm shape (R decreased as the range to predators
decreased, and the swarms became more spherical), as did swarm
nearest-neighbor distance (R decreased with increasing distance) and
swarm position in the water column (R decreased in the upper 70% of the
water column). Therefore, swarm shape appears to be influenced by a
combination of behavioral responses to predator presence and
environmental variables. MBEs have the potential to contribute much to
studies of krill, and can provide data to improve our understanding of
the behavior of krill
in situ.
J. D.
Warren, J.A. Santora, and D.A. Demer. 2009.
Submesoscale
distribution of Antarctic krill and its avian and pinniped predators
before and after a near gale.
Marine
Biology 156: 479-491.
doi:
10.1007/s00227-008-1102-0 [
.pdf]
We conducted two ship-based surveys of the nearshore ecosystem north of
Livingston Island, Antarctica during 2–10 February 2005. Between the
two surveys, a low-pressure system (963 mbar) passed through
the area
providing the opportunity to measure ecosystem parameters before and
after a near gale. A ship-based multiple-frequency acoustic-backscatter
survey was used to assess the distribution and relative abundance of
Antarctic krill (
Euphausia superba).
Net tows, hydrographic profiles, and meteorological data were collected
to measure biological and physical processes that might affect the
krill population. During the survey, the distribution and behavior of
several krill predators [chinstrap penguins (
Pygoscelis
antarctica), cape petrels (
Daption capense),
and Antarctic fur seals (
Arctocephalus gazella)]
were measured from the vessel by visual observations. The survey
encompassed an area of roughly 2,500 km
2,
containing two submarine canyons with one to the west and one to the
east of Cape Shirreff, which had different abundances of krill and
predators. Several aspects of the nearshore ecosystem changed after the
near gale including: hydrography of the upper 100 m of the
water
column, phytoplankton biomass, the abundance and distribution of krill,
and the distribution of some krill predators. Differences in these
parameters were also measured between the two canyons. These changes in
the physical and biological environment during the survey period are
quantified and show that the ecosystem exhibited significant changes
over relatively short spatial (tens of kilometers) and time
(tens of hours) scales.
To convert measurements of backscattered acoustic energy to estimates
of abundance and taxonomic information about the zooplankton community,
all of the scattering processes in the water column need to be
identified and their scattering contributions quantified. Zooplankton
populations in the eastern edge of Wilkinson Basin in the Gulf of Maine
in the Northwest Atlantic were surveyed in October 1997. Net tow
samples at different depths, temperature and salinity profiles, and
multiple frequency acoustic backscatter measurements from the upper
200 m of the water column were collected. Zooplankton samples
were
identified, enumerated, and measured. Temperature and salinity profiles
were used to estimate the amount of turbulent microstructure in the
water column. These data sets were used with theoretical acoustic
scattering models to calculate the contributions of both biological and
physical scatterers to the overall measured scattering level. The
output of these predictions shows that the dominant source of acoustic
backscatter varies with depth and acoustic frequency in this region. By
quantifying the contributions from multiple scattering sources,
acoustic backscatter becomes a better measure of net-collected
zooplankton biomass.
The
density and sound speed of two coastal,
gelatinous zooplankton,
Mnemiopsis leidyi
(a ctenophore) and
Cyanea capillata (lion's mane
jellyfish), were
measured. These parameters are
important inputs to acoustic scattering models.
Two
different methods were used to measure the density of
individual
animals: one used a balance and graduated cylinder to
determine
the mass and displacement volume of the animal, the
other
varied the density of the solution the animal was
immersed
in. When the same animal was measured using both
methods,
density values were within 1% of each other. A
travel-time
difference method was used to measure the sound speed
within
the animals. The densities of both zooplankton slightly decreased
as the animals increased in length, mass, and
volume. The
ratio of animal density and sound
speed to the surrounding
seawater (
g
and
h, respectively) are reported for both animals.
For
Mnemiopsis leidyi
ranging in length from 1 to 5 cm,
the
mean value (±standard deviation) of
g
and
h were
1.009
(±0.004) and 1.007 (±0.001). For
Cyanea
capillata ranging in
bell diameter
from 2 to 11 cm, the mean value (±standard
deviation) of
g and single
value of
h were 1.009
(±0.004)
and 1.0004. ©2007
Acoustical Society of America
The acoustic
backscatter intensity signal from a high-frequency
(600 kHz) Acoustic Doppler Current Profiler (ADCP) was used to
categorize four different types of bottom habitat (sand, mud, sparse
and dense vegetation) in a shallow-water estuary (Shinnecock Bay, NY,
USA). A diver survey of the bay measured sediment and bottom vegetation
characteristics at 85 sites within the bay. These data were used to
groundtruth the acoustic data. Acoustic data were collected at four
sites with known bottom types and used to develop an algorithm that
could categorize the bottom type. The slope of the echo intensity
profile close to the bottom was used to determine the bottom type and
the relative numerical density (sparse or dense) of Submerged Aquatic
Vegetation (SAV). In areas where eelgrass (Zostera marina)
was
the dominant SAV species, the intensity profile data were analyzed to
measure the height of the vegetation canopy. An acoustic survey which
categorized the bottom type of the bay was conducted from a small
vessel. The percentage of sampled sites categorized as each bottom
habitat type from the acoustic survey was similar to those obtained by
the diver survey. These methods may provide a means to rapidly survey
estuarine habitats and measure spatial and temporal variations in SAV
populations, as well as changes in the height of the eelgrass canopy.