FISHERIES OCEANOGRAPHY
17:2, 126–136, 2008
Remotely sensed spawning habitat of Pacific sardine
(Sardinops sagax) and Northern anchovy (Engraulis mordax)
within the California Current
CHRISTIAN S. REISS,1,* DAVID M.
CHECKLEY JR2 AND STEVEN J. BOGRAD3
1
Antarctic Ecosystem Research Division, NOAA Fisheries,
Southwest Fisheries Science Center, 8604 La Jolla Shores Dr.,
La Jolla, CA 92037, USA
2
Integrative Oceanography Division, Scripps Institution of
Oceanography, University of California San Diego, 9500
Gilman Dr., La Jolla, CA 92093-0218, USA
3
Environmental Research Division, NOAA Fisheries, Southwest
Fisheries Science Center, 1352 Lighthouse Ave., Pacific Grove,
CA 93950-2097, USA
ABSTRACT
We use trivariate kernel density estimation to define
spawning habitat of northern anchovy (Engraulis
mordax) and Pacific sardine (Sardinops sagax) in the
California Current using satellite data and in situ egg
samples from the Continuous Underway Fish Egg
Sampler (CUFES) deployed during surveys in April by
the California Cooperative Oceanic Fisheries Inves-
tigations (CalCOFI). Observed egg distributions were
compared with monthly composite satellite sea surface
temperature (SST) and surface chlorophyll a (chl a)
data. Based on the preferred spawning habitat, as
defined in SST and chl a space, the satellite data were
used to predict potential spawning habitat along two
areas of the west coast of North America. Data from
the southern area (21.5 to 39N) were compared to
observations from the CUFES data for the period
1998–2005. Northern anchovy and Pacific sardine
exhibited distinctly different spawning habitat distri-
butions. A significant relationship was found between
satellite-based spawning area and that measured dur-
ing surveys for sardine. CUFES area estimated for
sardine was similar in magnitude to that estimated
from satellite data (60 000 km2). In contrast,
spawning habitat of anchovy averaged between 1000
and 200 000 km2 for the period 1998–2005, for
*Correspondence: e-mail: christian.reiss@noaa.gov
Received 12 December 2006
Revised version accepted 21 August 2007
126 doi:10.1111/j.1365-2419.2008.00469.x
Fish. Oceanogr.
CUFES and satellite estimates, respectively. Interan-
nual variability in the area (km2) and duration
(months) of estimates of suitable habitat varied
between species and between the northern (39 to
50.5N) and southern portions of the California Cur-
rent. Long-term monitoring of habitat variability using
remote sensing data is possible in the southern portion
of the California Current, and could be improved upon
in the northern area with the addition of surveys
better timed to describe relationships between
observed and estimated spawning habitats.
Key words: California Cooperative Oceanic Fisheries
Investigations, California Current, northern anchovy,
Pacific sardine, remotely sensed, Sea-viewing Wide
Field-of-view Sensor, spawning habitat, sea surface
temperature
INTRODUCTION
Annual egg and larval surveys for the estimation of
spawning biomass and recruitment, or for the assess-
ment and delineation of essential fish habitat, are
conducted for many Coastal Pelagic Species (CPS; Lo
et al., 2001) of fish. Recent advances in continuous
in situ monitoring of eggs using the Continuous
Underway Fish Egg Sampler (CUFES; Checkley et al.,
1997, 2000a) provide fine-scale delineations of egg
occurrence that may be used to define spawning hab-
itats within the survey areas. The rather fixed nature
and expense of ship-based point estimate sampling
limits inference about changes in seasonal spawning
peaks and spatial changes in habitat use by CPS and
other co-sampled fish species (Smith, 1990, 1995).
This has important implications for management be-
cause indices of abundance may become biased, and
may simply reflect the availability of fish eggs to the
survey, when spawning shifts geographically with
change in the environment rather than population
productivity. Such shifts are predicted to occur with
global climate change (Perry et al., 2005; Rose, 2005).
Within the California Current (CC), Pacific sar-
dine (Sardinops sagax) and Northern anchovy
Journal compilation  2008 Blackwell Publishing Ltd.
No claim to original US government works

(Engraulis mordax) exhibit long-term, quasi-decadal
variability in abundance even in the absence of fishing
(Soutar and Isaacs, 1974; Baumgartner et al., 1992).
Considerable effort has been expended developing
ecological hypotheses to explain this variability in
abundance, and the changing effect of ocean condi-
tions on biological processes (Parrish et al.,1981;
MacCall, 1990; Jacobson and McCall, 1995). Mac-
Call’s (1990) basin hypothesis predicts that as popu-
lations of CPS stocks increase, they expand the area
occupied to fill other available habitats within the
basin and contract around a more or less fixed center
during periods of low population size. Others have
suggested that CPS populations will not only expand
and contract around a given fixed center but that
changes in suitable habitat may cause population
centers to shift latitudinally (Rodriguez-Sanchez et al.,
2002).
Testing such hypotheses requires observing or
sampling important components of the environment
that may affect productivity of populations across the
totality of potential habitat. Pacific sardine and
northern anchovy generally occupy different areas of
the CC for spawning. Egg distributions of each species,
based on CUFES data (Checkley et al., 2000b; Lynn,
2003) as well as quantitative net tows (Fiedler, 1983),
show distinct spatial segregation. Lluch-Belda et al.
(1991) showed that the thermal preferences of each
species differed, and that the thermal preference of
anchovy was broader than that of sardine.
Several studies within the CC have attempted to
describe the relationship between the distribution of
eggs of anchovy or sardine and the environment in
order to define spawning areas over limited spatial and
temporal periods (Fiedler, 1983; Lynn, 2003). Fiedler
(1983) used satellite-derived sea surface temperature
(SST) and chlorophyll a (chl a) data from the Coastal
Zone Color Scanner (CZCS) to describe the rela-
tionship between anchovy egg distributions, derived
from 4-km spaced vertical net tows, and the ocean
environment of the Southern California Bight (SCB).
Lynn (2003) used a combination of SST, measured
continuously from ship, and zooplankton biomass
inferred from Acoustic Doppler Current Profiler
(ADCP) backscatter, to describe the spawning habitat
of Pacific sardine based on egg distributions from three
CUFES sampling periods. In these cases, strong rela-
tionships between egg abundance and distribution and
the environment were observed.
More recently, the NOAA Fisheries Service
(Macewicz and Griffith, 2006) conducted CUFES
surveys off the Oregon coast (2003–2005) during July
and found Pacific sardine and northern anchovy eggs
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Satellite-derived spawning habitat estimates 127
in waters of similar temperature to the SCB, a char-
acteristic also noted by Emmett et al. (2005). The
presence of Pacific sardine larvae off Vancouver Island
from May through September was documented
between 1992 and 2004, although the contribution of
this spawning to population productivity remains
uncertain (McFarlane et al., 2005). A recent survey of
Pacific sardine off the US West Coast found its eggs
farther offshore and farther north than previously
suggested (Lo et al., 2007). The results of these studies
support the hypothesis that preferred spawning tem-
peratures are similar throughout the California Cur-
rent (CC) ecosystem, although the month of spawning
may vary along the west coast of North America.
These consistent coast-wide thermal preferences allow
us to use satellite and shipborne measurements, either
together or separately, to make inferences about the
spawning habitats of northern anchovy and Pacific
sardine off the US West Coast. As satellite-based time
series become longer, remote sensing of habitats could
provide a mechanism to test hypotheses regarding
environmental changes that may control the quasi-
decadal variability in population size, and could pro-
vide information to develop models in order to explain
long-term variability of populations of CPS in the CC
ecosystem for use in formulating management
decisions.
In this study, we examine relationships between the
egg distributions of Pacific sardine and northern
anchovy to SST and chl a, inferred from remote sens-
ing in the southern California Current. Kernel density
estimates are used to define habitat polygons based on
the distributions of SST, chl a, and the concentration
of eggs within the CalCOFI survey area. We then
investigate the relationship between satellite-based
spawning habitat and empirically derived estimates of
spawning habitat area based on the distribution of eggs
obtained using CUFES. The seasonal patterns and
extent of suitable habitat for each species are then
estimated to provide a baseline for the development of
remote indices of spawning habitat.
MATERIALS AND METHODS
Biological data
Egg distributions of Pacific sardine and northern
anchovy were assessed during the April CalCOFI
cruises from 1998 through 2005. The April cruise is
conducted with the principal goal of assessing the
biomass of spring spawning CPS, principally Pacific
sardine in the Southern California Bight (SCB;
Fig. 1). The general sampling scheme for CalCOFI has
128 C.S. Reiss et al.
Figure 1. Map of the west coast of North America from
Canada to Baja California showing the locations of the fixed
stations occupied during April CalCOFI cruises conducted
between 1997 and 2005.
50
Vancouver
Northern area
45
40
San Francisco
Monterey
Latitude (N)
35 Southern Pt. Conception
California San Diego
Bight
30
Southern
area
25
130 125 120
Longitude (W)
been described elsewhere (Hayward et al., 1999). In
2002 and 2003, two ships were used to sample the
ocean environment concurrently, and in those years
sampling density between San Diego and Pt. Con-
ception was twice that of other years (<40 km
between west to east lines).
Egg abundance (numbers min)1) was determined
using CUFES (Checkley et al., 1997, 2000a) by
pumping water from a fixed depth (3 m), passing it
through a concentrator and into a sample collector
(505 lm), then counting the number of eggs in each
sample collected over a fixed interval (30 min).
Sampling period is increased (15 min) once a high
abundance (>1 egg min)1) of Pacific sardine eggs is
found. At a nominal ship speed of 10 knots
(5 m s)1), these sampling intervals result in a broad
spatial sample of about 5 nmi (9 km) and 19 m3 of
water filtered, assuming a pumping rate of
640 L min)1 (Checkley et al., 1997). Because Pacific
sardine is the target species, sampling and along-
transect spatial resolution of the egg distributions are
finer than those of northern anchovy, a species that is
currently not assessed by NOAA Fisheries.
Journal compilation  2008 Blackwell Publishing Ltd, Fish. Oceanogr., 17:2, 126–136.
No claim to original US government works
In addition to egg abundance, the filtered water is
passed through a SBE-21e thermosalinograph (SBE
Inc.) providing geo-referenced near-surface (3 m)
temperature and salinity, concurrent with the egg
abundance data. Standard processing using SBE Inc.
supplied software was used to calculate salinity from
conductivity.
Spawning area
Spawning area from CUFES sampling was determined
by integrating the area (km2) encompassing all sam-
ples with more than one Pacific sardine or northern
anchovy egg per minute. We did not calculate a
standard error of the spawning area because the error
for the CUFES samples is unknown. We also did not
adjust egg abundance for extrusion that may underes-
timate the abundance of northern anchovy eggs when
a larger mesh netting is used to sample plankton
(Checkley et al., 2000b).
Satellite data
Sea surface chlorophyll. Monthly standard mapped
images of chl a were extracted for the region from
21.5 to 50.5N, and 105 to 130W from the Sea-
viewing Wide Field-of-view Sensor (SeaWiFS) archive
115 110 (oceans.gsfc.nasa.gov⁄SeaWiFS⁄L3SMI⁄). Monthly data
from September 1997 through December 2005
(100 months) were used to develop time series of chl
a along the west coast. These data have a nominal
spatial resolution of about 4 km at the equator, and
about 8 km at 36N, and are extracted as a nominal
9-km equidistant cylindrical grid.
Sea surface temperature. Global standard mapped
images of monthly SST data (ver. 5) were extracted
from the PO.DAAC Pathfinder archive (http://www.
nodc.noaa.gov/sog/pathfinder4 km/) for each month
from September 1997 through December 2005. These
images were extracted as a nominal 4-km equal angle
grid and then mapped onto an equi-rectangular
projection. The reported accuracy of these SST
data is 0.3–0.5C. The final array size for these data
was about three times that of the SeaWiFS data. The
SST data were therefore gridded to the array size of
the chl a data using the nearest neighbor algorithm
in MATLAB (The Mathworks, Inc. Natick, MA,
USA).
Spawning habitat distribution. To develop a habitat
model that included chl a and SST, we pooled the
data from the first half of the available time series of
egg surveys (1998–2001) and extracted chl a and SST
data from the satellite images for each corresponding
CUFES sample for each survey. We searched for data

that were ±4 km from each CUFES position, and
averaged the satellite data within this area. These 4 yr
included both an El Niño (1998) and a La Niña
(1999) event. Preliminary analysis of the relationship
between satellite-based SST and near surface SST
from the thermosalinograph showed a strong
correlation (range of r2 = 0.64–0.83). No large
deviations in the relationships were observed across
both El Niño (1998) and La Niña (1999) periods,
indicating that satellite data adequately reflected the
temperature patterns in the CalCOFI area during the
surveys. No continuous underway chl a data were
available for direct comparison with the satellite-
derived chl a. However, much of the data used to
calibrate the SeaWiFS were collected in the SCB
(Mitchell and Kahru, 1998), so our satellite-based
estimates of chl a should adequately reflect the surface
water chl a concentrations in this area.
We then used a three-dimensional, kernel density
estimator to determine the trivariate (SST, chl a, and
eggs min)1) probability density function (PDF) from
the 4 yr of pooled data using MATLAB and the KDE
toolbox (http://science.ntu.ac.uk/msor/ccb/densest.
html). Kernel density methods are non-parametric
probability density estimators (Scott, 1992) and are
generalizations of standard univariate procedures, like
histograms. The PDF is calculated by fitting a weigh-
ted kernel function for each data point and then
averaging over the data. In this case, a weight, h, is
determined by the desired amount of smoothing that
trades off between bias and variance in the data series
in each of the three dimensions. We used a normal
density kernel and applied the normal reference rule
for determining the optimal window size to smooth
each function. The result is a multivariate PDF
describing the habitat space. We investigated the
sensitivity of the resulting multivariate PDFs to three
different types of kernel functions (normal, triangular
and tri-weight). No major differences were found with
the exception of the smoothness of the multivariate
PDF.
The useful result of the kernel-density estimate in
this analysis is a polygon of the probability of finding
eggs of each species described by generic temperature
and chl a properties where eggs were actually observed.
Lynn (2003) showed that the spawning habitat of
sardine was well defined when contours of the ratio of
CUFES samples positive for eggs versus total CUFES
samples in a bin (temperature, potential food)
exceeded 50%. Thus, we chose to define spawning
habitat (SST and chl a) area as the polygon encom-
passing the upper 50% of the egg abundance of each
species, thereby retaining the most dense contribu-
Journal compilation  2008 Blackwell Publishing Ltd, Fish. Oceanogr., 17:2, 126–136.
No claim to original US government works
Satellite-derived spawning habitat estimates 129
tions of eggs, and potentially maximizing differences in
habitat use between the two species. These boundaries
in SST and chl a space form the boundaries of the
spawning habitat as described from remote sensing.
Because recent data show Pacific sardine and
northern anchovy spawn over a variety of months and
locations along the west coast of North America, but
at similar temperatures (McFarlane et al., 2005; Em-
mett et al., 2005; Richard Charter, SWFSC La Jolla,
CA pers. comm.), we used the boundaries of the 50%
polygon to define spawning habitat for each species
each month by mapping the monthly SST and chl a
data, and summing the calculated spawning area.
Because of latitudinal and seasonal variability, areas
north of 39N (San Francisco) were pooled into a
northern area, while areas between 21.5 and 39N
were pooled into a southern area. Further sub-division
in space did not dramatically affect the results.
Estimates of spawning habitat area for each of the
8 yr (1998–2005) inferred from the kernel-density
procedure in the southern area were compared with
the spawning area estimated by integrating the areas
encompassed by the observed CUFES egg distributions
using all 8 yr of CUFES survey data. Because both
estimates have the same units, we expect the rela-
tionships to be linear, although slopes and intercepts
may vary for a number of reasons including the dif-
fering cruise tracks in different years.
RESULTS
Distribution of sardine and anchovy
The distribution and abundance of Pacific sardine and
northern anchovy eggs off the coast of California
during April varied considerably among years (Fig. 2).
In general, northern anchovy eggs were found much
closer to shore and were concentrated in the SCB,
while the highest abundances of Pacific sardine eggs
were found offshore and north of Point Conception.
Occasionally (1999, 2002), Pacific sardine eggs were
distributed widely across the California Current, while
in other years (1998, 2001, and 2003) they were
concentrated more closely along the coast. Northern
anchovy eggs were generally more variable in spatial
distribution, usually occurring within the SCB
although sometimes (2005) found off Monterey.
Pacific sardine and northern anchovy exhibited
distinctly different interannual patterns in the amount
of spawning area in spring (April) between 1998 and
2005 in the CalCOFI region. Mean spawning area of
Pacific sardine during April was 63 940 km2. The
smallest spawning area (<12 000 km2) occurred in
130 C.S. Reiss et al.

1998 1999
35 35

30 30
Latitude (N)

25 25
–130 –125 –120 –115 –130 –125 –120 –115

2000 2001
35 35

30 30

25 25
–130 –125 –120 –115 –130 –125 –120 –115

2002 2003
35 35

30 30
Latitude (N)

25 25
–130 –125 –120 –115 –130 –125 –120 –115

2004 2005 Figure 2. Distribution and abundance

(egg min)1) of eggs of northern anchovy
35 35
(red bars) and Pacific sardine (black
bars) collected in CUFES samples during
30 30 April CalCOFI larval fish surveys
between 1998 and 2005. Open circles
represent the location of the fixed
25 25 CalCOFI stations. Bars in lower left are
–130 –125 –120 –115 –130 –125 –120 –115 scales (1, 15 and 25 eggs min)1, note
Longitude (W) different scales between species).

1998 and the largest in 1999 (125 398 km2), indicat-
ing that spawning area varied by an order of magnitude
during the transition from an El Niño to a La Niña.
The spawning area of northern anchovy was even
more variable, ranging over two orders of magnitude
during the time period. In 1999, just 535 km2 of the
SCB was used for northern anchovy spawning during
April, while in 2005 the spawning area was greater
than 40 000 km2 (Fig. 2).
sardine and northern anchovy (Fig. 3). The upper 50%
of Pacific sardine eggs collected between 1998 and 2001
were found in a relatively small SST–chl a space
(Fig. 3a–b). Pacific sardine eggs were associated with
lower chl a concentration (<1.2 mg m3) and a tem-
perature range of 12–15C, as measured by satellite. In
contrast, northern anchovy were more widely dispersed
in temperature (12.5–16.5C) and chl a (<0.5 mg m)3–
4.5 mg m)3) space (Fig. 3c–d).

Satellite-based estimates of spawning habitat Annual cycle of spawning area

Trivariate kernel density estimates of the PDF resolved Annual patterns in satellite-based estimates of
different patterns of habitat association between Pacific spawning habitat area varied in the northern and
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 Satellite-derived spawning habitat estimates 131

(a) (b)
10 20

Chlorophyll-a (mg m–3)

8
15

–1
Eggs min
6
10
4
5
2

Figure 3. Trivariate kernel density esti- 0 0

12 14 16 18 10 8 6 4 2 0
mates of Pacific sardine (a,b) and Temperature (C) Chlorophyll-a (mg m–3)
northern anchovy (c,d) encompassing (c) (d)
the 50% occurrence of eggs. Environ- 10 20

Chlorophyll-a (mg m–3)

mental space is defined by chl a
(mg m)3) and temperature (C) derived 8
15

–1
from monthly composites of SST and

Eggs min
6
SeaWiFS chl a during April 1998 10
through 2001. Left panels present planar 4
view of the habitat containing 50% of 5
2
the eggs, while the right panel presents
the elevation view of the abundance of 0 0
12 14 16 18 10 8 6 4 2 0
50% of the eggs as a function of chl a
–3
concentration. Temperature (C) Chlorophyll-a (mg m )

southern areas for both Pacific sardine and northern
anchovy (Fig. 4). Pacific sardine spawning habitat was
temporally uni-modal in the southern area and tem-
porally bi-modal in the northern area. In the southern
area, the most potential Pacific sardine spawning
habitat occurred from December through April,
exceeding 100 000 km2. In the north, satellite-based
estimates predicted year-round availability of Pacific
sardine spawning habitat, with the least amount of
habitat available from July through October. The
average amount of spawning habitat in May exceeded
80 000 km2 and in November the amount of potential
spawning habitat exceeded 100 000 km2. In April,
spawning habitat area for Pacific sardine in the
southern area began to decline and was minimal
between August and October.
Northern anchovy spawning habitat showed uni-
modal seasonal cycles in each area (Fig. 4). In the
southern area, potential northern anchovy spawning
habitat was maximal from February through April,
when it exceeded 300 000 km2. Spawning habitat was
available in the southern area year round. Even when
spawning habitat was minimal, more than 45 000 km2
of potential habitat was available for northern
anchovy. In the northern area, the spawning area was
maximal during summer, between May and Novem-
ber. There was a hint of bi-modality, with one mode in
June and July and a second mode in October. Of

Figure 4. Estimated monthly spawning

areas of Pacific sardine and northern
anchovy, predicted based on a trivariate
kernel density estimate of the satellite-
derived SST and chl a. The northern
area (black) represents areas north of
San Francisco and the southern area
(gray) represents the currently extended
CalCOFI sampling areas. Error bars (±1
SD) represent averaging over the 8 yr of
the study from 1998 to 2005.

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No claim to original US government works
132 C.S. Reiss et al.

particular interest was the complete lack of spawning 1998 and the larger area during the La Niña of 1999.
habitat for northern anchovy between January and In contrast, the time series of satellite- and CUFES-
April in the northern area. based estimates of spawning area were widely disparate
for northern anchovy. The seasonal trends of satellite-
Time series of spawning habitat based spawning habitat were an order of magnitude
When satellite-based spawning habitat was compared greater than that estimated by CUFES (Fig. 4). The
to that derived from the April CUFES measurements temporal patterns over the 8-yr period also varied
in the CalCOFI area, several interesting patterns greatly (Fig. 5b).
emerged (Figs. 5 and 6). Time series of spawning area Spawning habitat area of Pacific sardine inferred
estimated from satellite and CUFES data from April using satellite-based data was similar in magnitude to
were similar for Pacific sardine. Satellite-based mea- the area measured using CUFES (Fig. 6a), while this
sures of spawning habitat also resolved the smaller area was not so for northern anchovy (Fig. 6b). The
that was estimated using CUFES during the El Niño in CUFES and satellite-derived spawning areas for

1.5 1.5
Sardine (00 000s km )

Sardine (00 000s km )

2

1.0 1.0

Satellite spawning area

CUFES spawning area

0.5 0.5

2
0 0
1998 1999 2000 2001 2002 2003 2004 2005 Anchovy (00 000s km )

6 4
Anchovy (0 000s km )
2

Figure 5. Estimates of spawning area of

4 3.5 Pacific sardine (upper panel) and north-
ern anchovy (lower panel) in the
2 3
southern area based on data from CUFES
(solid line) and satellite (dashed line) for
April 1998 to 2005. Data from 1998 to
0 2.5
2

1998 1999 2000 2001 2002 2003 2004 2005 2001 were used to generate the trivariate
Year kernel estimates.

2 5

1.8 4.5

1.6 4
1999
2002
Anchovy (00 000s km )
Sardine (00 000s km2)

1999
2

1.4 3.5 2005

2001
1998
1.2 3 2000
Satellite

Satellite

1 2002 2.5
2004
Figure 6. Correlation between esti-
0.8 2
2000 2003 mated spawning area of sardine (left pa-
1998 nel) and anchovy (right panel) derived
0.6 2005 1.5
from CUFES sampling and the area
0.4 1 estimated from satellite data for the
period 1998 to 2005 in the California
0.2 0.5 Current between the US border and San
0
Francisco. Open circles represent the
0
0 0.5 1 1.5 2 0 1 2 3 4 5 data used to generate the trivariate ker-
CUFES CUFES nel density estimates. Closed circles
2 2
Sardine (00 000s km ) Anchovy (00 000s km ) represent independent data.

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No claim to original US government works
 Satellite-derived spawning habitat estimates 133

Pacific sardine were not significantly correlated time (Fig. 7). Spawning habitat was virtually constant
(r2 = 0.68; p < 0.08), although the high explained on an annual basis in both the southern and northern
variance indicates the lack of significance may be due areas for northern anchovy. However, potential
to low sample size. Predictions of spawning area using spawning habitat was available throughout the year
both CUFES and the satellite were lowest during El between 1999 and 2002 in the south but not the
Niño and highest during La Niña. In contrast, there north.
was little overlap in the spawning habitat of northern
anchovy when CUFES satellite estimates were com-
DISCUSSION
pared. For northern anchovy, the spawning area
based on remote sensing exceeded the CUFES-based Using satellite-based measures of two components of
estimate of spawning habitat by an order of the pelagic habitat (SST and chl a) and non-para-
magnitude. metric kernel density estimation techniques, we were
Monthly time series of satellite-based measures of able to quantify and locate the potential spawning
spawning area of both Pacific sardine and northern habitat of Pacific sardine and northern anchovy off the
anchovy from September 1997 through December west coast of the US over an 8-yr period. This result
2005 showed interesting multi-year patterns that dif- expands on several studies that have documented
fered between species and areas (Fig. 7). For example, relationships between spawning distributions of Pacific
between 1999 and 2002 Pacific sardine spawning sardine and northern anchovy in relation to temper-
habitat in the southern area was greater than in other ature and measures of primary or secondary production
years, and averaged more than 60 000 km2. At the in the CC system (Fiedler, 1983; Lluch-Belda et al.,
same time, there was less predicted spawning habitat 1991; Lynn, 2003) and elsewhere (Richardson et al.,
in the northern area. Pacific sardine spawning habitat 1998; Planque et al., 2007).
was virtually non-existent during the summers of 1999 Given that the time series of co-occurring satellite
to 2002 in the northern area. data and CUFES data are still very short, we used half
The lack of an obvious relationship between the time series to construct the three dimensional
CUFES- and satellite-based estimates of spawning area PDFs to determine the spawning preferences. This was
for northern anchovy is apparent and is discussed later. done by pooling the first 4 yr of data that included
Monthly patterns for this species show the range of both an El Niño and a La Niña. We then predicted the
variability that might be expected between areas over habitat area of each of the four remaining years and
also the first 4 yr. This adds bias to the predictions, but
was necessary given the short time series of the data.
Figure 7. Time series of monthly spawning areas As the time series lengthens and other data are
(·105 km2) of Pacific sardine (a) and northern anchovy (b) included (from other surveys during these years, and
predicted using a trivariate kernel density estimate of the future surveys) it will be easy to determine the bias of
satellite-derived SST and chl a. The northern area (black the current analysis. It is interesting to note that the
line) represents areas north of San Francisco and the 4 yr of independent Pacific sardine data are squarely
southern area (gray line) represents current CalCOFI located within the bounds of the 4 yr of data used to
sampling areas.
construct the PDFs.
(a) The lack of a relationship between satellite- and
2 2
CUFES-based estimates of spawning habitat for
northern anchovy may be due to a number of factors.
Southern area (00 000s km2)

Northern area (00 000s km2)

1 1 Firstly, the peak period of spawning for northern an-

chovy is between late January and April (Hunter and
0
Macewicz, 1980), and there are substantial differences
0
1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 in SST and chl a within the SCB and along the west
(b) coast between these months (Lynn et al., 2003). The
2 4
springtime surveys with CUFES may sample northern
anchovy eggs from late in its spawning season and thus
1 2 provide an estimate of the spawning habitat that is
biased in terms of both quantity and location. Sec-
0
ondly, other features of the local environment may be
0
1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 important in constraining the distribution of northern
Year anchovy that are not captured by the two measures
Journal compilation  2008 Blackwell Publishing Ltd, Fish. Oceanogr., 17:2, 126–136.
No claim to original US government works
134 C.S. Reiss et al.
that we have examined. For example, northern
anchovy are more closely associated with upwelling
areas than Pacific sardine, and so may not use all
potential offshore habitat. As a result, satellite-based
estimates may show habitat offshore that in fact would
not be used by northern anchovy. Thirdly, it has been
hypothesized that northern anchovy are more strongly
associated with regions of high primary production
than Pacific sardine because northern anchovy depend
on local primary production for energy for their serial
egg production during the spawning season (Smith and
Eppley, 1982). Such dependence may further restrict
them to localized areas of predictably high produc-
tivity. Finally, it is believed that the northern anchovy
population is at a lower level relative to historical
periods of the mid-1980s (California Department of
Fish and Game, 2001) and is therefore simply not
occupying as much available habitat as when it is more
abundant. This is consistent with both the overesti-
mation of observed (CUFES-based) spawning habitat
using satellite data and the lack of correlation between
CUFES- and satellite-based estimates for the 8-yr
period. Unfortunately, there has been no population
assessment since 1989, and the CPS fishery has almost
completely focused on the more valuable Pacific sar-
dine population that has increased to more than a
million tons since the early 1990s (Lo et al., 2007),
confounding resolution of these competing
explanations.
An important assumption of this approach is that
mis-matches in the temporal and spatial scales and the
influence of advection do not confound the patterns
we are trying to detect (sensu Taggart and Frank,
1990). Temporally, we have used monthly composites
of both SST and chl a as shorter time periods con-
tained large numbers of missing values owing to clouds
and other quality control issues. As our ultimate goal
was also to derive coast-wide indices, monthly com-
posites minimized the loss of data along the entire
coast. The high correlations between the ship-based
and remotely sensed SST and the consistency among
years indicate that the general environmental condi-
tions in each year (e.g., during El Niño and La Niña)
were those experienced by fish during spawning and
manifest in the temporal composites.
Unlike the near-surface SST from the thermosali-
nograph used for comparison with the satellite-based
SST, there were no high-resolution ship-based mea-
surements to compare with the satellite-derived chl a.
However, comparisons of remotely sensed chl a data
with shipborne sampling have shown high correlations
within the CC (Mitchell and Kahru, 1998). Indeed,
many of the data used to determine algorithms for the
Journal compilation  2008 Blackwell Publishing Ltd, Fish. Oceanogr., 17:2, 126–136.
No claim to original US government works
SeaWiFS project were collected during CalCOFI
cruises (Mitchell and Kahru, 1998) and thus the
satellite-based estimates of chl a we used are probably
representative of in situ chl a in the CC during the
sampling periods. The advent of the MODIS AQUA
satellite and its simultaneous measurement of both
SST and chl a should minimize the mis-match
between these space-based measurements in the
future. As the spatial resolution of both satellite and
CUFES data ranges from 4 to 9 km, spatial biases are
probably not severe. Biases owing to the differential
advection of heat, productivity, or eggs are largely
unknown and are an unquantified error in our analy-
ses. Loss or gain of heat by both conduction and
evaporation are also potential sources of variability
that could impact SST over monthly time scales
(Pickard and Emery, 1990). Fish eggs of CPS hatch
after 3 to 4 days in the water column at the temper-
atures we commonly observe in the California Current
(12–14C; Zweifel and Lasker, 1976). With a
10 cm s)1 net alongshore flow (Marchesiello et al.,
2003), eggs would be displaced <40 km before
hatching, a distance less than between nominal lines
of the CalCOFI survey. Although significant vari-
ability in the along- and cross-shelf flow does occur, it
is unlikely that advection of eggs into or out of areas of
differing temperature or chl a is a greater source of
error than the compositing of monthly satellite-based
SST and chl a measures across the survey area.
Finally, this analysis used objective measures based
on empirical probability distributions to define opti-
mal habitat from the abundance of eggs, SST and chl
a distributions, using a normal kernel density esti-
mator. While more precise estimates of the kernel
density may be possible using some other type of
underlying distribution, such improvements would
not necessarily result in a positive relationship
between measured and calculated spawning area for
northern anchovy.
The positive correlation between the satellite-
based measure of spawning area and the area esti-
mated using CUFES for Pacific sardine is promising
and suggests that satellite-based measurement of
spawning habitat and its temporal variability is pos-
sible. Other studies using CUFES data have shown
how egg distributions were related to temperature and
salinity (Checkley et al., 2000a) and show promise to
explain the distribution of eggs with respect to water
masses. Yet, at present, there is neither a method to
measure salinity remotely nor is there a simple rela-
tionship between temperature and salinity in the CC
from which to infer salinity from temperature. Thus,
at present, remote measures of habitat are limited to

those described by SST and chl a, although other
remotely measured variables such as sea surface
height (Espinosa-Carreon et al., 2004), and derived
variables such as particulate organic carbon (Stramski
et al., 1999) and eddy energy (Ducet and Le Traon,
2001), offer potential.
A number of studies have documented the spatial
separation in the distribution of Pacific sardine and
northern anchovy within the CC (Fiedler et al.,
1986; Checkley et al., 2000a). Our results show
similar patterns, but the use of satellite data assesses
potential habitat independent of the area surveyed
using ships. This is critical, as recent data from the
CPS fishery indicates an offshore movement of sar-
dine beginning in the mid-1990s (Hill et al., 2006),
an area not well sampled by current egg surveys.
More recently (2006 and 2007) sardine spawning has
occurred well offshore and at the most seaward Cal-
COFI stations (Richard Charter; SWFSC La Jolla,
CA, USA, pers. comm.). Recent surveys off central
California suggest continued expansion into less fre-
quently sampled areas with similar temperature and
productivity (Richard Charter; SWFSC La Jolla, CA,
USA., pers. comm.). The use of remote sensing to
assess spawning habitat provides information on the
general patterns of spawning associated with tem-
perature and chl a, which may be due to shifts to less
utilized habitat. Several studies have documented the
results of changing environmental conditions in the
CC on the growth and productivity of both anchovy
and sardine (e.g., Fiedler et al., 1986). There are no
clearly quantified studies modeling the spawning
habitat and the environment, despite the ability to
empirically define spawning habitat with increasing
precision in any given season. This study provides the
framework to develop environmental indices of
habitat area. With respect to northern anchovy,
CUFES sampling in a variety of areas along the coast
could help to resolve why we so drastically over-
estimated habitat and found no relationship between
the satellite- and CUFES-based estimates. Continued
sampling in the northern sector of the California
Current system may further contribute to the tem-
perature and chl a relationship we present here for
Pacific sardine, as spawning may have recently
occurred there (Emmett et al., 2005; McFarlane et al.,
2005). Continued assessment of spawning habitat
from both ships (e.g., CUFES) and space (e.g., ocean
color from satellites) will enable characterization of
spawning habitat occurrence and use by Pacific sar-
dine and northern anchovy over annual scales and,
thus, how this habitat variability is linked to popu-
lation dynamics and demography.
Journal compilation  2008 Blackwell Publishing Ltd, Fish. Oceanogr., 17:2, 126–136.
No claim to original US government works
Satellite-derived spawning habitat estimates 135
ACKNOWLEDGEMENTS
We thank all the researchers, technicians and stu-
dents that make a program as large as the CalCOFI
seagoing program a success. The authors also thank
the SeaWiFS Project (Code 970.2) and the Distrib-
uted Active Archive Center (Code 902) at the
Goddard Space Flight Center, Greenbelt, MD 20771,
for the production and distribution of these data,
respectively. These activities are sponsored by
NASA’s Mission to Planet Earth Program. This
research was funded by the Fisheries and the Envi-
ronment (FATE) Program of the National Marine
Fisheries Service.
REFERENCES
Baumgartner, T., Soutar, A. and Ferriera-Bartrina, V. (1992)
Reconstruction of the history of pacific sardine and northern
anchovy populations over the past two millenia from sedi-
ments of the Santa Barbara Basin, California. CalCOFI
Repts. 33:24–40.
California Department of Fish and Game. (2001). California’s
Living Marine Resources: A Status Report. pp. 585.
Checkley, D.M., Ortner, P.B., Settle, L.R. and Cummings, S.R.
(1997) A continuous, underway fish egg sampler. Fish.
Oceanogr. 6:58–73.
Checkley, D.M., Dotson, R.C. and Griffith, D.A. (2000a)
Continuous, underway sampling of eggs of Pacific sardine
(Sardinops sagax) and northern anchovy (Engraulis mordax) in
spring 1996 and 1997 off southern and central California.
Deep-Sea Res. 47:1139–1155.
Checkley, D.M., Hunter, J.R., Motos, L. and van der Lingen,
C.D. (2000b). Report of a Workshop on the Use of the Con-
tinuous Underway Fish Egg Sampler (CUFES) for Mapping
Spawning Habitats of Pelagic Fish. GLOBEC Report 14, pp. 1–
65.
Ducet, N. and Le Traon, P.Y. (2001) A Comparison of Surface
Eddy Kinetic Energy and Reynolds Stresses in the Gulf
Stream and the Kuroshio Current Systems from Merged
TOPEX⁄Poseidon and ERS-1⁄2 Altimetric Data. J. Geophys.
Res. 106:16603–16622.
Emmett, R.L., Brodeur, R.D., Miller, T.W. et al. (2005) Pacific
sardine (Sardinops sagax) abundance, distribution, and eco-
logical relationships in the Pacific northwest. CalCOFI
Repts. 46:122–143.
Espinosa-Carreon, T.L., Strub, P.T., Beier, E., Ocampo-Torres,
F. and Gaxiola-Castro, G. (2004). Seasonal and interannual
variability of satellite-derived chlorophyll pigment, surface
height, and temperature off Baja California. J. Geophys Res.
C03039, doi:10.1029/2003JC002105.
Fiedler, P.C. (1983) Satellite remote sensing of the habitat of
spawning anchovy in the southern California Bight.
CalCOFI Repts. 26:202–209.
Fiedler, P.C., Methot, R.D. and Hewitt, R.P. (1986) Effects of
California El Niño 1982–1984 on the northern anchovy.
J. Mar. Res. 44:317–338.
Hayward, T.L., Baumgartner, T.R., Checkley, D.M. et al. (1999)
The State of the California Current in 1998–1999: Transi-
tion to Cool-Water Conditions. CalCOFI Repts. 40:29–62.
136 C.S. Reiss et al.
Hill, K.T., Lo, N.C.H., Macewicz, B.J. and Felix-Uraga, R.
(2006) Assessment of the Pacific sardine (Sardinops sagax
caerulea) population for U.S. management in 2007. NOAA
Tech. Mem.396 104.
Hunter, J.R. and Macewicz, B.J. (1980) Sexual maturity, batch
fecundity, spawning frequency and temporal pattern of
spawning for the northern anchovy, engrualis mordax during
the 1979 spawning season. CalCOFI Repts. 21:139–148.
Jacobson, L.D. and McCall, A.D. (1995) Stock recruitment
models for Pacific sardine (Sardinops sagax). Can J. Fish.
Aquat. Sci. 52:566–577.
Lluch-Belda, D., Lluch-Cota, D.B., Hernandez-Vazquea, S. and
Salina-Zavala, C.A. (1991) Sardine and anchovy spawning
as related to temperature and upwelling in the California
Current system. CalCOFI Repts. 32:105–111.
Lo, N.C.H., Hunter, J.R. and Charter, R.L. (2001) Use of a
continuous egg sampler for ichthyoplankton surveys: appli-
cation to the estimation of daily egg production of Pacific
sardine (Sardinops sagax) off California. Fish. Bull. 99:554–
571.
Lo, N.C.H., Macewicz, B.J., Griffith, D.A. and Charter, R.L.
(2007) Spawning biomass of Pacific sardine (Sardinops sagax)
off US and Canada in 2006. NOAA Tech. Mem. SWFSC
401. pp. 32.
Lynn, R.J. (2003) Variability in the spawning habitat of Pacific
sardine (Sardinops sagax) off southern and Central California.
Fish. Oceanogr. 12:541–553.
Lynn, R.J., Bograd, S.J., Chereskin, T.K. and Huyer, A. (2003)
Seasonal renewal of the California Current: the spring
transition off California. J. Geophys. Res. 108:3297–3306.
MacCall, A.D.. (1990). Dynamic geography of marine fish
populations. Seattle, WA: Washington Sea Grant. Wash-
ington University Press, Inc. pp. 153.
Macewicz, B.J. and Griffith, D.A. (2006). Initial results from
four Pacific sardine (Sardinops sagax) surveys off the coasts
of Oregon and Washington. In: Minutes of the 2005 Tri-
national sardine forum. Administrative Report LJ-06-05.
N.C.H. Lo, A.C. Allen & S.Z. Herzka, (eds). La Jolla, CA:
NOAA SWFSC, pp. 80–85.
Marchesiello, P., McWilliams, J.C. and Shchepetkin, A. (2003)
Equilibrium structure and dynamics of the California current
system. J. Phys. Ocean. 33:753–783.
McFarlane, G.A., Schweigert, J., MacDougall, L. and Hrabok,
C. (2005) Distribution and biology of Pacific Sardine (Sar-
dinops sagax) off British Columbia, Canada. CalCOFI Repts.
46:144–160.
Mitchell, B.G. and Kahru, M. (1998) Algorithms for SeaWiFs
standard products developed with the CalCOFI bio-optical
data set. CalCOFI Repts. 39:133–147.
Journal compilation  2008 Blackwell Publishing Ltd, Fish. Oceanogr., 17:2, 126–136.
No claim to original US government works
Parrish, R.H., Nelson, C.S. and Bakun, A. (1981) Transport
mechanisms and reproductive success of fishes in the Cali-
fornia current. Biol. Oceanogr. 1:175–203.
Perry, A.L., Low, P.J., Ellis, J.R. and Reynolds, J.D. (2005)
Climate change and distribution shifts in marine fishes.
Science 308:1912–1915.
Pickard, G.L. and Emery, W.J. (1990) Descriptive Physical
Oceanography: An Introduction Oxford, England: Pergamon
Press, Inc.
Planque, B., Bellier, E. and Lazure, P. (2007) Modelling
potential spawning habitat of sardine (Sardina pilchardus) and
anchovy (Engraulis ecrasicolus) in the Bay of Biscay. Fish.
Oceanogr. 16:16–30.
Richardson, A.J., Mitchell-Innes, B.A., Fowler, J.A. et al.
(1998) The effect of sea temperature and food availability on
the spawning success of Cape anchovy Engraulis capensis in
the southern Benguela. S. Afr. J. Mar. Sci. 19:275–290.
Rodriguez-Sanchez, R., Lluch-Belda, D., Villalobos, H. and
Ortega-Garcia, S. (2002) Dynamic geography of small
pelagic fish populations in the California Current System on
the regime scale (1931–1997). Can. J. Fish. Aquat. Sci.
59:1980–1988.
Rose, G. (2005) On distributional responses of North Atlantic
fish to climate change. ICES J. Mar. Sci. 62:1360–1374.
Scott, D.W. (1992). Multivariate Density Estimation: Theory,
Practice, and Visualization, New York: John Wiley & Sons, Inc.
Smith, P.E. (1990) Monitoring interannual changes in spawning
area of Pacific sardine (Sardinops sagax). CalCOFI Repts.
31:145–151.
Smith, P.E. (1995) Development of the population biology of
the Pacific hake, Merluccius productus. CalCOFI Repts.
36:144–152.
Smith, P.E. and Eppley, R.W. (1982) Primary production and
the anchovy population in the Southern California Bight:
comparison of time series. Limnol and Oceanogr., 27:1–17.
Soutar, A. and Isaacs, J.D. (1974) Abundance of pelagic fish
during the 19th and 20th centuries as recorded in anaerobic
sediment off the Californias. Fish. Bull. 72:257–273.
Stramski, D., Reynolds, R.A., Kahru, M. and Mitchell, B.G.
(1999) Estimation of particulate organic carbon in the ocean
from satellite remote sensing. Science 285:239–242.
Taggart, C.T. and Frank, K.T. (1990) Perspectives on larval fish
ecology and recruitment processes: probing the scales of
relationships. In: Large Marine Ecosystems: Patterns, Processes
and Yields (AAAS Selected Symposium Series). K. Sherman &
L.M. Alexander (eds) Washington, D.C: Am. Assoc.
Advancement of Science, pp. 151–164.
Zweifel, J.R. and Lasker, R. (1976) Prehatch and Posthatch
Growth of Fishes – a General Model. Fish. Bull. 74:609–621.