North American Journal of Fisheries Management 22:1270–1275, 2002

q Copyright by the American Fisheries Society 2002

Reducing Gill-Net Mortality of

Incidentally Caught Coho Salmon
SEANA BUCHANAN AND ANTHONY P. FARRELL
Department of Biological Sciences, Simon Fraser University,
Burnaby, British Columbia V5A 1S6, Canada

JAKE FRASER
Rural Route 1, Site 11B, C-11,
Madeira Park, British Columbia V0N 2H0, Canada

PATRICIA GALLAUGHER
Continuing Studies in Science, Simon Fraser University,
Burnaby, British Columbia V5A 1S6, Canada

RUTH JOY AND RICK ROUTLEDGE*

Department of Statistics and Actuarial Science, Simon Fraser University,
Burnaby, British Columbia V5A 1S6, Canada

Abstract.—The abundance of coho salmon Oncorhyn-
chus kisutch has declined dramatically over much of the
southern part of its range along the Pacific Coast of
North America. This decline has created the need to
reduce fishing mortalities, including bycatch mortalities
in fisheries that are targeting other species. Traditional
gill-net fishing causes an estimated 35–70% mortality
rate on incidentally caught coho salmon. A reduction in
this high mortality rate is necessary if gill nets continue
to be used in fisheries that inadvertently intercept de-
pressed coho salmon stocks while fishing other species.
By using modified gear, short net soak times, careful
handling of fish on removal from the gill net, and a newly
designed recovery box, the short-term mortality rate on
incidentally caught coho salmon can be reduced to as
little as 6%, possibly even lower in some circumstances.
This substantial reduction in mortality on nontargeted
species expands the possible role of gill nets in the de-
velopment of selective fisheries.
The abundance of coho salmon Oncorhynchus
kisutch has declined drastically over much of the
southern part of its range in North America (Weit-
kamp et al. 1995). In 1996, coho salmon on the
central California coast were formally declared as
threatened under the U.S. Endangered Species Act
(National Marine Fisheries Service 1996). By
1998, two more formal declarations had extended
this status to coho salmon on the coasts of northern
California and Oregon (National Marine Fisheries
Service 1997, 1998). In Washington State, coho
salmon outside of the Olympic Peninsula are cur-
* Corresponding author: routledg@stat.sfu.ca
Received April 30, 2001; accepted February 6, 2002
rently candidates for formal listing (National Ma-
rine Fisheries Service 2001a). In British Columbia,
major conservation efforts have been initiated in
response to severely depressed coho salmon stocks
in the Upper Skeena and Thompson rivers (Coho
Response Team 1998).
To preserve these dwindling populations, man-
agement agencies have imposed substantial re-
strictions on commercial fisheries. In many in-
stances, regional coho salmon fisheries have been
eliminated, and incidental mortality of coho salm-
on in commercial fisheries that target other salmon
species has been substantially reduced, primarily
through major restrictions on harvesting times and
areas (see Routledge and Wilson 1999; Fisheries
and Oceans Canada 2001a; National Marine Fish-
eries Service 2001b). Furthermore, in 1998, the
Canadian federal government announced a new
Selective Fishing Policy subsequently revised
(Fisheries and Oceans Canada 2001b). In addition
to avoidance measures (e.g., through restrictions
on time and area fished), the policy calls for re-
ductions in bycatch mortality. For any fishing
method to remain a viable part of the Canadian
commercial salmon fishery, its coho salmon by-
catch mortality cannot be too high. Bycatch mor-
tality is a product of two factors: the incidental
encounter rate and the mortality rate of those fish
that are incidentally encountered. This paper fo-
cuses on the latter factor.
Although gill-net fishing can often be conducted
in such a way as to avoid nontarget species, more
than half of the incidentally caught fish still may

1270
 MANAGEMENT BRIEFS
die. The Canadian Department of Fisheries and
Oceans estimates that 35–70% of all coho salmon
caught in gill-net fisheries die shortly after en-
countering the gear (Coho Response Team 1998);
it has formally applied a 60% mortality rate for
encountered fish in estimating total mortalities for
gill-net-caught coho salmon (Fisheries and Oceans
Canada 1999). This high mortality rate has se-
verely restricted opportunities for gill-net fisher-
ies. It has also prompted efforts to devise tech-
niques to increase the survival of gill-net-caught
coho salmon.
As part of the selective fishing policy, each com-
mercial vessel has been mandated to carry and use
a recovery box for the purpose of reviving inci-
dentally caught coho salmon. However, there are
reasons to question the effectiveness of the recov-
ery box. Although Blewett and Taylor (1999) con-
cluded that the boxes were very effective in in-
creasing survival of nontarget species in some cas-
es, earlier studies into the physiology of salmon
had indicated that holding live fish in tanks in-
creased the fishes’ lactate levels (Parker and Black
1959; Ellis 1964). The controversy led to exper-
iments in 1998 to investigate the performance of
the standard recovery box in reviving salmon cap-
tured with all commercial gear types. Farrell et al.
(2000) found very little evidence of physiological
recovery in commercially caught coho salmon af-
ter 60 min in the standard revival box.
To explore a promising alternative, a study was
undertaken in 1999 to compare the effectiveness
of the standard recovery box with that of a newly
designed Fraser recovery box (Fisheries and
Oceans Canada 2001c). The new design force-ven-
tilates fish while they recover metabolically, mak-
ing recovery possible even if a fish has been as-
phyxiated and cannot ventilate for itself. In an ex-
perimental comparison, the Fraser box reduced the
coho salmon mortality rate substantially below the
50% level for the standard box in that experiment
(Farrell et al. 2001). In addition, the Fraser recov-
ery box proved to be more beneficial than the stan-
dard revival box from a physiological perspective
because it led to decreased muscle lactate levels
and increased muscle phosphocreatine and gly-
cogen after only 1 h (Farrell et al. 2001). When
the use of the new recovery box was coupled with
modified gear, net soak times of less than 60 min,
and careful fish handling, coho salmon mortality
was less than 2%.
Nevertheless, to better represent the effective-
ness of the Fraser box and other tools in the context
of a commercial fishing situation, we undertook
the current study in 2000. Several trained fishers
1271
in multiple locations used the above-described
modifications to evaluate coho salmon mortality
in conditions that better represented the commer-
cial sockeye salmon O. nerka fishery.
Methods and Study Sites
The primary goal of the experiment was to test
the effectiveness of the methods that had been de-
veloped in the earlier experiments, when used by
newly trained crews in a setting that more closely
mimicked a commercial fishery. We assessed the
mortality rate of coho salmon that were caught by
a gill net in an approximately 40-min set, handled
carefully, and placed in a Fraser recovery box if
in need of resuscitation. Furthermore, being de-
signed to assess the mortality rate in a wide variety
of conditions and locations, the experiment re-
tained an exploratory flavor.
The experiment was conducted at three sites.
The first site was located in the Skeena Estuary in
northern British Columbia, the other two on the
inner South Coast of southern British Columbia in
Johnstone Strait and Knight Inlet. The first two
sites were in major migration routes that contained
mixed stocks of coho salmon. The Knight Inlet
site was believed to contain mostly coho salmon
returning to spawn in the inlet. Coho salmon
caught in the Skeena Estuary and in Knight Inlet
showed early signs of the epidermal thickening
that is commonly associated with sexual matura-
tion in Pacific salmon (Idler et al. 1961; McBride
and van Overbeeke 1971). The observations from
Johnstone Strait, where coho salmon did not dis-
play these signs, provided an opportunity to probe
for differences in mortalities between sites that
might be related to different levels of spawning
maturation.
Fishing procedures.—Fishers attempted to
achieve soak times (the time from when the first
cork hit the water as the net was being set out until
the last cork was removed as the net was being
hauled and cleared of fish) of 40 min, but spring
tides, strong river currents, and floating debris in
the Skeena Estuary imposed considerable vari-
ability in soak times (mean 5 45 min, SD 5 17
min); soak times were less variable in Johnstone
Strait (mean 5 41 min, SD 5 10 min) and Knight
Inlet (mean 5 42 min, SD 5 6 min). In addition,
because Knight Inlet was unlikely to contain se-
riously depleted coho salmon stocks like those
originating in either the Upper Skeena or Thomp-
son watersheds, this site permitted the testing of
longer soak times, which one would expect to pro-
duce higher mortality rates (Farrell et al. 2000).
To this end, additional sets were made at this site
1272 BUCHANAN ET AL.

TABLE 1.—A summary of the experimental methods employed at each study site. The letters indicate individual
vessels, with vessel A being present at all three sites. Net abbreviations are as follows: AK4, four-strand Alaska twist
tooth tangle net; Multi, 16-gauge river tangle net; AK6, six-strand Alaska twist outside or ocean net. Intended soak
times are shown; actual values were variable.

Area Boats Nets Dates Soak times

Skeena Estuary A, B, C AK4, Multi Jul 26–Aug 7 40 min
Johnstone Strait A, D, E, F AK6 Aug 27–Sep 2 40 min
Knight Inlet A, E AK6 Sep 9–Sep 18 40 and 140 min

with a target soak time of 140 min (mean 5 142
min, SD 5 7 min). Logistical constraints forced
most of the shorter soak times to be taken earlier
in the experimentand thus required investigating
the potential for confounding between soak times
and date in the statistical analysis.
Six vessels were used in the experiment. In
keeping with the goal of mimicking natural fishing
conditions, three different net designs were de-
ployed. Shorter lengths were chosen to promote
the achievement of short soak times at times of
higher abundance. In addition, net designs were
selected with the aim of reducing bleeding and
suffocation. In the Skeena Estuary, the following
two net types were used: a 0.15 mm by four-strand
Alaska Twist tooth tangle net, with an 80- or 100-
fathom shackle length, a 60-mesh depth, a 10-cm
web, a 2.2:1 hang ratio, and a weed line; and a
35-strand, 16-gauge multistrand river tangle net,
with a 100-fathom shackle length, a 60-mesh
depth, an 11-cm web, a 3.0:1 hang ratio, and a
weed line.
In the last two sites, only one net type was used:
a 0.2 mm by six-strand Alaska Twist net, with a
200-fathom shackle length, a 90-mesh depth, a 12-
cm web, a 2.0:1 hang ratio, and a weed line.
Crews and onboard observers on each vessel
generally remained on the same vessel for the du-
ration of the experiment at each site. Hence, be-
tween-boat variation was driven by a composite of
several factors, including net design (only in the
Skeena Estuary), crews, and observers. Crews and
observers were all trained to handle coho salmon
TABLE 2.—Number of coho salmon brought on board
the gill-net vessels and the number of those that subse-
quently died on board, by site and, for Knight Inlet, by
set length. For Knight Inlet, the short sets were all less
than 60 min, the long sets more than 130 min.
Number
Site Set length caught
Skeena Estuary Short 102
Johnstone Strait Short 83
Knight Inlet Short 101
Long 84
as carefully as possible to minimize abrasion, gill
damage, and air exposure. Where appropriate, the
net was cut to ease fish removal. Key aspects of the
experimental design are summarized in Table 1.
All coho salmon that were not in an advanced
stage of rigor mortis were promptly placed in on-
board Fraser recovery boxes and held there until
they were upright and ventilating normally. They
were then transferred to recovery pens as soon as
possible. A few fish escaped during transfer to net-
pens, and some vigorous fish were released di-
rectly from recovery boxes if space was needed
for less vigorous fish. Status of fish in the net-pens
was monitored approximately 24 and 48 h after
transferal.
After 48 h in the net-pen, the coho salmon were
either released or tested for swimming perfor-
mance as an indirect measure of the physiological
state of the fish when released. As many fish as
possible at each site were tested for maximum sus-
tainable swimming speed in a swim tunnel. Equip-
ment and procedures were as described in Farrell
et al. (2001). Body measurements were taken after
the swim test was completed.
Statistical analysis.—Onboard mortality rates
(the fraction of coho salmon removed from nets
that died on board the vessel) were analyzed by
logistic regression analysis (Agresti 1990:84–91)
with respect to the effect of soak time, between-
site and between-boat variation, and potential con-
founding of soak time and date. This modification
and extension of the standard chi-square test for
frequency data is the most appropriate way to test
frequency-count data for the effects of several fac-
tors. Although it depends on a specific form for
the dependence on a quantitative factor such as
soak time, given that soak times primarily clus-
tered tightly around only two values (40 and 140
min), the assumption is no more critical than the
Number standard assumption of linearity in a regression
dying analysis in which the x-observations similarly
11 clustered around only two values. Where appro-
6 priate, observations were categorized into set-
6 length and date intervals and were analyzed by
53
log-linear model analyses (Agresti 1990:143–150)
 MANAGEMENT BRIEFS
to confirm the lack of importance of this assump-
tion.
Net-pen mortalities were compared between the
1999 and 2000 studies by means of a simple chi-
square test, confirmed with a log-linear model
analysis (Agresti 1990, pp. 143–150) to account
for confounding with fish condition.
Results and Discussion
The experiment provided highly significant ev-
idence of a substantial reduction in mortality as-
sociated with shorter soak times (P 5 0.0001).
Also between-site (P 5 0.57) and between-boat (P
5 0.46) variations were not statistically signifi-
cant. Hence, there is no evidence that any of the
factors (e.g., net configuration) associated with be-
tween-site or between-boat variation was influ-
ential.
We analyzed the data for Knight Inlet separately
to assess the potential confounding between soak
time and date. We found significant evidence of
an increasing trend in mortality with date (P 5
0.002 for the logistic regression and P 5 0.011 for
the log-linear analysis on the grouped data), with
further evidence that the trend was more prevalent
for shorter sets (P 5 0.024 for a quadratic inter-
action term in the logistic regression and P 5 0.002
for the log-linear analysis). The reason for this
trend is unclear and presents a mild source of po-
tential confounding with soak time.
Nonetheless, the soak time effect was still high-
ly significant (P 5 0.008) and still pointed to a
substantial reduction in mortality with a shortened
soak time. Because the mean mortality for 40- ver-
sus 140-min sets showed signs of varying sepa-
rately with time, straightforward comparisons can
be made only on specific dates. On the midrange
date for the experimental fishing in Knight Inlet,
estimates of mortality for 40- versus 140-min sets
were 2.5% versus 60.4%.
The combined observations from all sites (with
no date effect included in the model) demonstrate
the remarkable decrease in estimated mean mor-
tality with shorter soak times (Figure 1). For 40-
min sets, the estimated mortality was 6.7% (95%
confidence limits, 4.3–10.2%). This supports ob-
servations from the previous study, in which the
estimated mean mortality for 40-min sets was
5.7% (95% confidence limits, 2.7–8.7%) and con-
trasts sharply with the 95% confidence limits of
52.0–72.0% for 140-min sets obtained from the
current study (these limits for the longer soak time
apply of course only to Knight Inlet, the only site
where the longer soak times were possible).
We also note the lack of data in the range from
1273
50 to 130 min. The shape of the curve remains
untested in this region, and mortality estimates ob-
tained from the logistic curve are not reliable for
soak times in this range.
These confidence limits reflect uncertainty in the
mean mortality, averaged across all sites, boats,
and dates. The large P-values for site-to-site dif-
ferences (0.57) and boat-to-boat differences (0.46)
show that the amount of variation among these
estimates is no more than one would expect to be
generated by chance alone. Nonetheless, it does
not follow that the actual differences are small.
Estimates of site-specific mortality rates ranged
from 4.9% to 8.0%, and estimates of boat-specific
mortality rates ranged from 4.6% to 13.2%. The
real extent of between-site and between-boat dif-
ferences can only be assessed through more ex-
tensive experimentation.
The experiment also revealed that the Fraser re-
covery box successfully resuscitated asphyxiated
fish—but only those captured during short soak
times. Of 52 fish brought on board in an apparently
lifeless condition from short sets, 26 (50%) were
resuscitated in the recovery box. In contrast, after
sets of around 140 min, only 2 of 57 (4%) were
resuscitated.
Net-pen mortalities were higher (9.7%) in this
study than in the previous year, where the mortality
rate was 2.1% (Farrell et al. 2001) and were in-
fluenced by fish condition on arrival (Table 3).
The causes of these higher mortality rates are
unknown. The differences cannot be attributed to
chance alone (P 5 0.039 for the chi-square test;
P 5 0.026 for the log-linear model analysis that
adjusts for differences in fish condition on removal
from the gill net). Unfavorable conditions in the
net-pens in the present experiment could have been
a contributing factor. In the Skeena River, strong
tidal currents may have pinned weaker fish against
the side of the pen. In Johnstone Strait, heavy algal
growth on the pens may have interfered with water
exchange and created hypoxia. Notably, in Knight
Inlet where neither of these net-pen difficulties was
observed (but also where conditions were most
similar to those in the previous experiment), none
of the 36 fish placed in net-pens after having been
caught in short sets died. In fact, the 2 fish (of 62
fish placed in net-pens) that did die were taken in
long sets.
The swim tests provided further evidence of re-
covery. Of 44 fish placed in the swim tunnel, all
but 4 swam, and these attained a swimming speed
of at least 1.44 body lengths per second, a velocity
comparable to the speeds reported in Farrell et al.
(2001) for physiologically recovered fish. One of
1274 BUCHANAN ET AL.

FIGURE 1.—Estimated average mortality rate as a function of soak time. The curve is the estimate produced by
the logistic regression analysis. The circles are the observed mortality rates for soak times within 10 min of the
value on the horizontal axis (e.g., 40 6 10 min). Solid circles represent data from the current study, open circles
data from the 1999 study for comparison. (The open circle corresponding to the 40-min soak time was moved
slightly to the left to improve its visibility, and only data from the present study were used in fitting the curve.)
Vertical lines represent approximate 95% confidence intervalss (intended to portray the accuracy of the logistic
regression fit to the observed mortality rates, these limits were calculated directly from the corresponding observed
mortality rates). There were too few sets in the 70–130-min range to provide direct, informative estimates of
mortality rates. The shape of the curve remains untested in this region.

the four fish that did not swim had lost its pectoral Nonetheless, four important caveats must be
fins; the other three appeared to have been sub- recognized. First, effects on long-term survival
jected to considerable stress during the transfer and eventual spawning success are unknown. This
process from the net-pen to the swim tunnel. appears directly testable only through comprehen-
sive tagging experiments. Second, encounter rates
Conclusions and Management must be kept low. In particular, this experiment
Recommendations did not assess the impacts of multiple encounters.
The experiment provides strong evidence that Third, these major reductions were achieved only
48-h coho salmon mortality from encounters with with educated crews, trained in the use of short
gill nets can be reduced substantially, as evidenced soak times and careful fish handling. Thorough
by the surviving fish having recovered their swim- training of commercial fishers is required transfer
ming ability. This success indicates the possibility of technology and knowledge to be adequate.
of an expanded role for gill nets in selective fish- Fourth, economic incentives to process target spe-
eries. cies may well interfere with the motivation and
ability to handle incidentally caught coho salmon.
Some combination of changes to economic incen-
TABLE 3.—Number of fish that died in net-pens, by con-
dition when removed from the gill nets. tives, education, monitoring, and enforcement will
be needed to achieve these results.
Number dying Finally, the experiment pointed to a notable in-
Fish condition Number captured in net-pens crease in mortality over time at one site. The re-
Vigorous 34 0 sults were also consistent with considerable be-
Lethargic 89 6 tween-site variability. These observations under-
Asphyxiated 22 8
score the potential value of more extensive ob-
 MANAGEMENT BRIEFS
servations on the dependence of mortality rates on
time and area. Vulnerability of coho salmon to
encounters with gill nets may vary with such fac-
tors as fish maturity and water temperature. Such
dependencies, if better understood, could be used
strategically to reduce bycatch impacts.
Acknowledgments
This research was funded by the Natural Sci-
ences and Engineering Research Council of Can-
ada, the Canadian Department of Fisheries and
Oceans, and commercial fishermen. Gordon Currie
and Brigid Payne provided liaison support with
the Department of Fisheries and Oceans. Fishing
was conducted by Ken and Jordan Lawson, Clar-
ence and Pearl Innes, Murray Tanner, Adrian Bel-
veal, and Ian MacKay. Rebekah Leakey, Mitch
Canlas, Ron Laplante, Susan Jirik, and Darlaene
Eccleston served as observers. Technical and lo-
gistical support were provided by Danielle Pike,
Brooke Fraser, Michael and Maureen Berry,
George Jolliffe, Butch and Kathy Duthie, the com-
munities of Dodge Cove and Alert Bay, staff of
North Pacific Cannery Historic Site, Alert Bay Ma-
rine Laboratory, Nimpkish hatchery, Stolt Sea
Farms Inc., and Ocean Fisheries.
References
Agresti, A. 1990. Categorical data analysis. Wiley-In-
terscience, New York.
Blewett, E., and T. Taylor. 1999. Selective fisheries. Re-
view and evaluation. Edwin Blewett and Associates,
Inc. and Timothy Taylor Consulting, Inc. January
1999 report to Fisheries and Oceans Canada, Van-
couver.
Coho Response Team. 1998. Coho salmon final report.
Fisheries and Oceans Canada, Pacific Region, Van-
couver.
Ellis, R. J. 1964. The effect of confinement on blood
lactate levels in chinook and coho salmon. Oregon
Fish Commission Research Briefs 10:28–33.
Farrell, A. P., P. Gallaugher, C. Clarke, N. DeLury, H.
Kreiberg, W. Parkhouse, and R. Routledge. 2000.
Physiological status of coho salmon (Oncorhynchus
kisutch) captured in commercial nonretention fish-
eries. Canadian Journal of Fisheries and Aquatic
Sciences 57:1668–1678.
Farrell, A. P., P. E. Gallaugher, J. Fraser, D. Pike, P.
Bowering, A. K. M. Hadwin, W. Parkhouse, and R.
Routledge. 2001. Successful recovery of the phys-
iological status of coho salmon on board a com-
mercial gillnet vessel by means of a newly designed
revival box. Canadian Journal of Fisheries and
Aquatic Sciences 58:1932–1946.
Fisheries and Oceans Canada. 1999. Fishing salmon se-
lectively in British Columbia. Report of the third
selective fisheries multi-stakeholder workshop.
1275
Fisheries and Oceans Canada and Fisheries Renewal
British Columbia, Vancouver.
Fisheries and Oceans Canada. 2001a. Pacific region in-
tegrated fisheries management plan, salmon. Fish-
eries and Oceans Canada. Available: www.pac.
dfo-mpo.gc.ca/ops/fm/mplans/mplans.htm#salmon.
(September 2002).
Fisheries and Oceans Canada. 2001b. A policy for se-
lective fishing in Canada’s Pacific fisheries. Fish-
eries and Oceans Canada, Vancouver.
Fisheries and Oceans Canada. 2001c. Coho revival
tanks construction plans. Fisheries and Oceans Can-
ada. Available: www.pac.dfo-mpo.gc.ca/ops/fm/
Salmon/index.htm (September 2002).
Idler, D. R., I. I. Bitners, and P. J. Schmidt. 1961. 11-
ketotestosterone: an androgen for sockeye salmon.
Canadian Journal of Biochemistry and Physiology
39:1737–1742.
McBride, J. R., and A. P. van Overbeeke. 1971. Effects
of androgens, estrogens, and cortisol in the skin,
stomach, liver, pancreas, and kidney in gonadec-
tomized adult sockeye salmon (Oncorhynchus ner-
ka). Journal of the Fisheries Research Board of Can-
ada 28:485–490.
National Marine Fisheries Service. 1996. Endangered
and threatened species; threatened status for central
California coast coho salmon evolutionarily signif-
icant unit (ESU). Federal Register 61(31 October
1996):56138–56149.
National Marine Fisheries Service. 1997. Endangered
and threatened species; threatened status for north-
ern California/southern Oregon coast evolutionarily
significant unit (ESU) of coho salmon. Federal Reg-
ister 62(6 May 1997):24588–24609.
National Marine Fisheries Service. 1998. Endangered
and threatened species; threatened status for the
Oregon coast evolutionarily significant unit of coho
salmon. Federal Register 63(10 August 1998):
42587–42591.
National Marine Fisheries Service. 2001a. Endangered
Species Act status of West Coast salmon and steel-
head. National Marine Fisheries Service. Available:
www.nwr.noaa.gov/1salmon/salmesa/pubs/1pgr.
pdf. (September 2002).
National Marine Fisheries Service. 2001b. Fisheries off
West Coast states and in the Western Pacific: West
Coast salmon fisheries; 2001 management mea-
sures. Federal Register 66(8 May 2002):23185–
13295.
Parker, R. R., and E. C. Black. 1959. Muscular fatigue
and mortality in troll-caught chinook salmon (On-
corhynchus tshawytscha). Journal of the Fisheries
Research Board of Canada 16:95–106.
Routledge, R. D., and K. Wilson. 1999. Coast-wide
coho. Pacific Fisheries Resource Conservation
Council, 1998–1999 Annual Report, Background
Paper 1999/1d, Vancouver.
Weitkamp, L. A., T. C. Wainwright, G. J. Bryant, G. B.
Milner, D. J. Teel, R. G. Kope, and R. S. Waples.
1995. Status review of coho salmon from Wash-
ington, Oregon, and California. NOAA Technical
Memorandum NMFS-NWFSC-24. Available: www.
nwfsc.noaa.gov/pubs/tm/tm24/coho.pdf. (September
2002).