Vol 23 Bailey Etal

Download as pdf or txt
Download as pdf or txt
You are on page 1of 18

BAILEY ET AL.

: PACIFIC WHITING LIFE HISTORY AND FISHERY


CalCOFl Rep., Vol. XXIII, 1982

THE LIFE HISTORY AND FISHERY


OF PACIFIC WHITING, MERLUCClUS PRODUCTUS
KEVIN M. BAILEY,' ROBERT C. FRANCIS PAYSON R. STNENS
Northwest and Alaska Fisheries Center Scripps Institution of Oceanography
National Marine Fisheries Service University of California, San Diego
Seattle, Washington 98112 La Jolla, California 92093

ABSTRACT coastal stock of Pacific whiting. Since the publication


The Pacific whiting is one of the most abundant and of a similar synopsis in 1970 (U.S. Fish and Wildlife
important fishes of the California Current region. This Service 1970), a great deal of new information has
report synthesizes available data, published and un- become available. Most of this material is unpublished
published, on the life history and population dynamics and thus is generally unavailable to scientists, mana-
of whiting. Aspects of the life history described are gers, and fishermen. Further goals of this synopsis are
distribution, spawning, early life history, feeding, and to present new information, particularly concerning
growth. Information on the population dynamics of the migration of whiting, and to suggest areas of
the stock is summarized with attention to stock abun- needed research.
dance, recruitment variability, and mortality. A syn-
thesis of the fishery, its development and manage- THE CALIFORNIA CURRENT SYSTEM-THE
ment, is presented. HABITAT OF PACIFIC WHITING
Pacific whiting ranges from the Gulf of Alaska to
RESUMEN the Gulf of California (Hart 1973); however, it is most
Merluccius productus es una de las especies mas abundant within the region of the California Current
abundantes e importantes en la regi6n de la Corriente de system. The California Current system is the eastern
California. En este trabajo se recopilan 10s datos pu- boundary current system of the North Pacific Ocean. It
blicados e inkditos sobre el ciclo de vida y dinimica de extends from the coastal divergence of the westwind
poblaciones de M. productus. Los aspectos del ciclo drift at 45"N in winter (and 50" in summer) southward
de vida que se discuten incluyen; distribucih, puesta, to about 23"N, where California Current water mixes
fases larvales y juveniles, alimentacibn y crecimiento. with equatorial water and bends westward to form the
La informacih sobre la dinimica de la poblacih se North Equatorial Current. The California Current I

resume en relaci6n con la abundancia de las existen- system is composed of (1) an equatorward surface
cias, variaciones en el reclutamiento y la mortalidad. flow-the California Current; (2) a seasonally occur-
Se presenta ademis una shtesis de las pesquerias, su ring poleward surface current identified as the David-
desarrollo y administracibn. son Current north of Pt. Conception, and as the
California Countercurrent in southern California; and
INTRODUCTION (3) a poleward subsurface flow-the California Un-
Commercially and ecologically the Pacific whiting dercurrent. Numerous gyres, including the Southern
(also called Pacific hake), Merluccius productus, is California Eddy, are semipermanent features of the
one of the most important fish species on the west California Current system. The individual currents are
coast of North America. It supports a large commer- briefly discussed below; a more detailed review can be
cial fishery that has been dominated by foreign na- found in Hickey (1979).
tions. In recent years, however, a U.S. fishery has The flow of the California Current is driven by
developed through ventures with foreign nations. Be- winds and is slow, broad, and shallow. Water of the
sides being an important resource to man, whiting is California Current is subarctic in physical and chemi-
an important trophic link in the California Current cal properties at high latitudes, characterized by low
ecosystem. As a large predator, whiting interacts with salinity and temperature. As the water flows south-
other fish and shellfish populations, notably the com- ward, it becomes more intermediate in nature through
mercially important stocks of Pacific herring, Clupea mixing with the high-salinity and high-temperature
harengus pallasi; northern anchovy, Engraulis mor- water of the North Pacific Current and the Central
dux; and shrimp. Whiting is also important as prey in Pacific water mass. Eventually, California Current
the diets of marine mammals itnd large fishes. water becomes semitropical off southern Mexico after
The objective of this synopsis is to synthesize avail- mixing with equatorial water.
able information on the biology and fishery of the The Davidson Current is the surface poleward flow
'Current address: College of Fisheries, University of Washungton, Sea&, WA 98195
north of Pt. Conception that develops in winter. The
[Manuscrip received January 1 1 , 1982.1 Davidson Current appears in October off Vancouver

81
BAILEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

Island and develops later farther south. It exists off the spawn in winter off the coasts of southern California
Oregon-Washington coast from October until Feb- and Baja California. In spring and summer, large fish
ruary and off the California coast from November until migrate northwards as far as central Vancouver Island,
January. and juveniles remain off the Californias. The migra-
The California Undercurrent is a northward flow of tion of whiting is outlined in Figure 1 and is described
high-salinity , high-temperature water occurring sea- in detail below.
ward of the continental shelf and below the main pyc-
Spawning
nocline. Wooster and Jones (1970) reported that the
Spawning schools of Pacific whiting have been dif-
undercurrent was found 75 km offshore of Cab0 Col-
ficult to locate. Nelson and Larkins (1970), Tillman
nett, Baja California, and was centered at 200-500-m (1968), Bureau of Commercial Fisheries 1964l, Erich
depth off the Oregon-Washington coast. A poleward
et al. (1980), and StepanenkoZ report spawning
undercurrent develops over the continental shelf in late schools off southern California in midwater at depths
summer and early fall (Hickey 1979).
of 130-500 m and over bottom depths corresponding
LIFE HISTORY to those of the continental slope. (Spawned at these
depths, eggs float upwards to the base of the mixed
Stocks and Distribution layer.) Ermakov (1974) also reports spawning over
Stocks. At least four distinct stocks of Pacific whit- the continental slope. However, Erich et al. (1980)
ing may exist. These include (1) a coastal stock rang- report a spawning school some 400 km seaward in the
ing from Canada to Baja California, (2) a Puget Sound southern part of the Southern California Eddy, and
stock, (3) a Strait of Georgia stock, and (4) a dwarf Stepanenko3 reports a spawning school about 300 km
stock found off Baja California. Two of the stocks, offshore in central California.
h g e t Sound and the coastal stock, have been iden- The distribution of eggs and small larvae (2-3 mm)
tified as genetically distinct spawning stocks (Utter indicates that whiting spawn from Cape Mendocino to
and Hodgins 1971). southern Baja California. Almost all eggs and larvae
The separate identities of the dwarf and coastal are located over water depths corresponding to depths
stocks are at present controversial. Ahlstrom and of the continental slope, except in the Southern
Counts (1955) examined larvae found off Baja California Eddy, where eggs and larvae are often
California and were not able to distinguish two sepa- found over very deep water and far out to sea (400
rate stocks, thus supporting a concept of one spawning km). Bailey (1981a) postulated that whiting spawn in
stock. However, MacGregor (1971) and Vrooman and the California Undercurrent, which usually occurs
Paloma (1977) discussed several differences in adult over the continetal slope at depths of 200-400 m, but
dwarf whiting found off Baja California compared spreads seaward some 200-400 km in the Southern
with the adult coastal whiting found farther north. California Eddy and some other locations where ed-
Dwarf whiting grow slower from age one onwards, dies occur. Large concentrations of eggs and larvae
mature earlier, and have several different morphomet- are found overlying areas of northward geostrophic
ric and meristic characteristics compared with the flow at 200-m depth (Figure 2).
coastal whiting. Vrooman and Paloma ( 1977) believed Variation may exist in the latitudinal distribution of
that these differences indicate separate stocks. How- spawning. The location of the apparent northern front
ever, the differences may not be genetic, and are not of spawning is correlated to the sea surface tempera-
inconsistent with changes caused by environmental ture (Table 1). Assuming that temperatues at the sea
effects in the different habitats. surface are correlated to those at the depth of spawn-
The remainder of this report deals with the coastal ing, this indicates that in warm years when subtropical
stock, which is the most abundant and commercially water is farther north, spawning occurs at higher
important. latitudes. Alternatively, larvae may be transported by
Distribution. As indicated previously, Pacific a northward flow.
whiting are found within the coastal region of the IBureau of Commercial Fisheries. 1964. Cruise report: exploratory cruise No. 64.
California Current system. Normally whiting are not Unpubl. manuscr. Northwest and Alaska Fish. Cent., Natl. Mar. Fish. Serv., NOAA,
caught seaward of the continental slope, although 2725 Montlake Blvd. E . , Seattle, WA 98112.
2Stepanenko, M.A. 1978. m e patterns of the abundance of the California anchovy
there are occasional reports of whiting eggs and larvae and Pacific hake, and estimation of their biomass, 1976-1977. Unpubl. manuscr.
(as well as of juveniles and adults) far seaward of the Pacific Scientific Research Institute of Marine Fisheries and Oceanography (TINRO),
Vladivostok, USSR.
slope (Frey 1971). The latitudinal distribution of 3Stepanenko, M.A. 1980. Reproductive condition and assessment of the spawning
whiting varies seasonally. In autumn adult whiting stocks of Pacific hake, California anchovy, horse mackerel, and some other fish
species in the California Current zone in 1979. Unpubl. manuscr. Pacific Scientific
make an annual migration from the summertime Research Institute of Marine Fisheries and Oceanography (TINRO), Vladivostok,
feeding grounds off the Pacific Northwest coast to USSR.

82
BAILEY ET AL.:PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

w JY
PACIFIC WHITING h
51 30N
MIGRATION BEHAVIOR CANADA

I ‘
Cape Flattery \
.-::..
By JulyIAugust WASHINGTON
4 1 30N
moving to outer Destruction Island
continental shelf

Heceta Bank

OREGON
43 30N

- -----_ _

39 30N

CALIFORNIA
WINTER
Migrating offshore over slope
in California current (south)

4
\ 35 30N

31 30N

Ea Spawning

Feeding

Main Schooling Area


27 30N

138 O O U 128 oou I 18 oou

Figure 1. Migratory patterns of Pacific whiting.

83
BADLEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

130. 125' 1200 115' 110.


I I
I I I 1 I I I 1 I I 4 1
I I I 1 1
I
'AL'OFI

JANUARY
1950-65 MEAN
CAPE
GEOSTROPHIC FLOW AT 200m DEPTH
ME NDOCINO ( TOPOGRAPHY OF THE 2 0 0 OEClOAR SURFACE,
IN D Y N A M I C M E T E R S , R E L A T I V E TO T H E 500-
O E C I B A R SURFACE 1
40 0''

42

*
\

I
SAN
FRANCISCO

I
I
I
I
I

44
\, .:*
46 Y
\ A42 \
\ \

4? ..t3

\
\ \
- 4s , %. . \

50.
-a
\ \ \ a
\
\ \
I
I
I
Y I I
r.50 / I
/ I
... '
-
,40
/ /
0
/

. /
f
I..

-
.
,& /e*-

/ A - \

,/.
-
I
/ \

I O> 1000 Eggs and larvae/lOm


- \
\
- t-
!
- 46

-+CU(lltNT C4RCCTION *OICATLO W bRROWS -+---


-
ACPWO~IIATE zoo i f w n MTTOY cnmroun
'COG
1
mea
1 I I 1 I 1 1 I I I I I I 1
I 1 I 1 1
130-__ 125. 120" I 15' I IO"

MEAN 200/500 db
JANUARY
Figure 2. Large catches of Pacific whiting eggs and larvae (all size classes) in January surveys, 1950-79, plotted on a chart showing geostropic flow at 200-m depth
(from Wyllie 1967).

84
BAILEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

TABLE 1 180-
The Northward Extent of 2-3-mm Standard Length Whiting u)
z
Larvae' during January Surveys, Compared to the Average 160-
January 50-Meter Temperature in the Los Angeles Bight U
I-
u)
Temperature Distance cI 140-
Year "C Rank Line Rank wa
1963 12.0 9 87 8.5 30 120-
1964 13.2 4 76 4
1965 12.4 7 80 6 ?
1966 13.5 3 70 2.5 y 100-
5
1968 12.8 5 70 2.5 a
U
1969 13.6 2 80 6 -I 80.
1972 12.2 8 87 8.5 u.
0
1975 12.6 6 80 6
1978 13.9 1 63 1
*Measured by the northernmost CalCOFI line where larvae occurred in
numbers greater than 100 larvae/lO mz.
Smaller line numbers are farther north. Temperature and northward extent
of larvae are significantly correlated using a Spearman rank correlation
statistic (P<.Ol).
b
Larvae of all size classes occur in significant num- FEE IWRCH RPRIL
I

M Y .&
bers in the water from December to May (Stauffer and MONTH
Smith 1977), but some 80% of eggs and small larvae Figure 3. The average monthly number of small whiting larvae per occupied
are found in two months, January and February (Fig- station in the California Cooperative Oceanic Fisheries Investigationssurvey
region, lines €0-120, Over the years 1963-79. Small larvae are 2-3 mmSL.
ure 3), which indicates a sharp peak in spawning.
Most Soviet reports also indicate that January and Early Life History
February are the primary spawning months, but some- Egg stage. Ahlstrom and Counts (1955) described
times heavy spawning is reported in March. Spawning the eggs of Pacific whiting. They are smooth spheres,
is generally completed by late March; in several con- have a single oil droplet, and are 1.14-1.26 mm in
secutive years Ermakov (1974) observed schools of diameter (after accounting for 7% shrinkage caused by
postspawning whiting off northern California by early preservation in Formalin). Egg hatching is tempera-
March. ture dependent (Bailey, in press; Zweifel and Lasker
Pacific whiting females mature and spawn at 3 to 4 1976). Whiting eggs may be expected to hatch in
years of age and at lengths of 34-40 cm (Best 1963; 100-120 hr at temperatures found at their habitat depth
MacGregor 1966, 1971; Ermakov 1974). MacGregor on the spawning grounds, where temperatures range
(1971) found some males maturing at 28 cm. Spawn- from 11" to 14°C.
ing whiting do not appear to migrate vertically, and In a laboratory setting, predators capable of eating
bilayered schools have been observed on sonar traces. whiting eggs are numerous and include, among others,
(R. McNeely, Northwest and Alaska Fisheries Center, medusae, ctenophores, and amphipods (Bailey and
Seattle, WA 98112, pers. comm.; J. Mason, Pacific Yen, in press). Whiting eggs may be somewhat re-
Biological Station, Nanaimo, B.C., pers. comm.). sistant to tactile and small invertebrate predators be-
Several modes of eggs appear in whiting ovaries cause they are motionless and have a very hard cuticle.
(MacGregor 1966; Ermakov et al.4). MacGregor Fish predation may also be heavy: Ermakov and Khar-
suggests that only one mode develops, because of the chenkd report finding the stomachs of threadfin bass,
poor condition of the females, but he did not examine Anthias gordensis, full of whiting eggs off Baja
the ovaries histologically, Foucher and Beamish California. Northern anchovy could also be feeding on
(1977) reported that only one mode of eggs develops whiting eggs, for they consume considerable numbers
in the Strait of Georgia whiting stock. Ovaries average of their own eggs (Hunter and Kimbrell 1980) and are
about 8% of the body weight of spawning females. believed to feed at depths where whiting eggs occur
Ripe ovaries contain 80-600 advanced-mode eggs per (Holliday and Larsen 1979).
gram of ovary wet weight (MacGregor 1966). An Larval stage. Ahlstrom and Counts (1955) de-
equation relating total fecundity to length of the fe- scribed the larvae of Pacific whiting. They are distin-
male is E = 0.00142*L3 (MacGregor 1966). guished by a pigment band around the tail, pigment
4Emakov, Y,,V. Snytko, L. Kodolov, I. Serobaba, L. Borets, and N. Fadeev. (Date
5Ermakov, Y., and A.M. Kharchenko. 1976. Biological characteristics of Pacific
unknown). Biological characteristics and the condition of stocks of Pacific hake,
hake and the estimation of its abundance in 1975. Unpubl. manuscr. Pacific Scientific
rockfish, blackcod, and pollock in 1972. Unpubl. manuscr. Pacific Scientific Re-
Research Institute of Marine Fisheries and Oceanography (TlNRO), Vladivostok,
search Institute of Marine Fisheries and Oceanography (TINRO), Vladivostok,
USSR.
USSR.

85
BAILEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

spots on the dorsal crown of the head, sturdy bodies, TABLE 2


and 51-54 myomeres (Figure 4). Preserved yolk-sac Percent Biomass of Juvenile Pacific Whiting by
larvae are 2.5-3.0 mm standard length. Shrinkage of Depth Interval and by Age
larvae due to handling is highly variable, from 1 0 4 % Depth Age (YO
depending on the preservative and on the time from (fathoms) 0 1 2 ~~
3
death to preservation (Bailey, in press; Theilacker 0-99 95.4 35.8 14.6 30.7
1981). 100-199 4.4 63.4 59.6 41.8
150-199 - 0.2 - 0.8 -25.8 27.5
Time to absorption of the yolk is temperature de-
100.0 100.0 100.0 100.0
pendent (Bailey, in press). At ambient temperatures,
From the summer 1977 Northwest and Alaska Fisheries Center bottom-
absorption of the yolk may take 120-200 hours. A trawl survey.
mouth develops before the yolk is fully depleted, and
yolk-sac larvae are observed to feed (Sumida and
(Ahlstrom 1959). If a strong pycnocline does not
Moser 1980). Larvae take 150-250 hours to starve
exist, eggs and larvae may be distributed through the
after yolk depletion (Bailey, in press).
mixed layer. Some evidence exists that larger larvae
Daily growth of whiting larvae has been described
may be distributed deeper than small larvae (Bailey, in
by counting growth increments on their otoliths
press).
(Bailey, in press). Growth in length appears to be slow
Juvenile stage. Not much is known about juvenile
and constant for the first 20 days, after which it rapidly
whiting. Juveniles 1-3 years of age are found primarily
accelerates (Figure 5). off central and southern California (Figure 6). Most
In the laboratory, predators on yolk-sac larvae are
0-1 year-olds occur inshore of the 200-fathom (fm)
more varied than those on eggs and include
isobath, and older fish are distributed somewhat
euphausiids, medusae, ctenophores, amphipods, and
farther offshore than younger fish (Table 2). The food
carnivorous copepods. Invertebrate predation on
of juvenile whiting is mainly copepods and euphau-
whiting larvae is stage- or size-specific; larger larvae
siids (P. Livingston “The Feeding Biology of Pacific
are not as vulnerable to predators as yolk-sac larvae
Whiting,” in review).
(Bailey and Yen, in press).
The diet of larval whiting is composed mostly of Adult Life History
copepod eggs, calanoid copepod nauplii, copepo- Migratory behavior. Tagging of Merluccidae has
dites,and copepod adults (Sumida and Moser 1980). not proved feasible (Jones 1974); thus the migrations
Whiting larvae have relatively large mouths and feed of Pacific whiting are inferred from survey and
on a broad size range of prey from 50-400 p m in fisheries data.
width. Pacific whiting become scarce in survey catches
Competitors of whiting larvae in the ichthyo- (Table 3) and in the fishery (Table 4) from autumn
plankton sharing the same temporal and spatial dis- until early spring (see also Jow 1973; Best 1963; Alton
tributions are California smoothtongue, Bathylagus 1972), and whiting eggs and larvae are most abundant
stilbius, and snubnose blacksmelt, Bathylagus in winter off California. These observations have led
wesethi. Overlap in the vertical and horizontal dis- to a hypothesis that adult whiting leave the coastal
tribution also occurs with Vinciguerria lucetia; waters in autumn to migrate from the shelf and south-
rockfish, Sebastes spp. ; and jack mackerel, Trachurus ward for spawning in winter, and then return north-
symmetricus. Numerous carnivorous invertebrates are ward in early spring (Alverson and Larkins 1969).
also competitors. This migratory pattern has been verified from more
The vertical distribution of whiting eggs and larvae. recent data (Ermakov 1974; Dark et al. 1980).
Eggs are released at 130-500 m in spawning, and most Speeds of migration may be estimated from the se-
rise upwards to a depth of neutral buoyancy, usually at quential appearance of fish up the coast after spawn-
40-60-m depth, near the base of the mixed layer ing. Postspawning accumulations of whiting normally

TABLE 3
Average Trawl Catches (Pound per Hour) of Pacific Whiting by Month and Year

Year Jan. Mar. May July Aug. Sept. Nov. Dec .


1961 - - - 2,403 - 9,784 - 97
1962 - 45 71 - 11,131 - 425 -
1963 100 - 175 - 4,315 - 450 -
- - - ~ ~ ~ - -
Mean 100 45 123 2,403 7,723 9,784 438 97
From Alton 1972.

86
BAILEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

Figure 4. Stages of Pacific whiting larvae (from Ahlstrom and Counts 1955).

87
BAILEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

TABLE 4 kGE 2 AGE I

The Proportion of Soviet Catches in the INPFC

Month
Vancouver-Columbia Area by Month, 1973-76

1973 1974 1975 1976 X


n
1 - - - - -
2 - - - - -
3
4
5
6
-
.032
.008
,106
,012
,073
,085
.049
-
-
,026
,163
-
-
,122
,207
,003
,026
,060
.131
AGE' r i AGE L

7 ,305 ,212 .406 ,202 ,281


8 .296 .116 ,167 ,261 ,210
9 ,158 ,453 ,136 .188 .234
10 ,094 - ,086 .017 ,049
11 - - .016 - .004
12 - - - .002 -

have appeared around San Francisco (38"N) in early


March (Ermakov 1974; Erich et al. 1980) and have
been later observed off southern Oregon (42"N) in the AREA
third week of April for five consecutive years from Figure 6. The distribution of biomass by International North Pacific Fisheries
1966 to 1971 (Ermakov 1974). A population traveling Commission area for each age dass in the 1977 Northwest and Alaska
Fisheries Center trawl survey: Area 1, Conception; Area 2, Monterey; Area
on this schedule would move, on the average, about 3, Eureka; Area 4, Columbia; Area 5, Vancouver (from Bailey and Ainley, in
10 k d d . By May, concentrations appear off Van- press).
couver Island. These estimated mean population
1975) indices of wind stress tends to support this
speeds compare favorably to speeds obtained from hypothesis. Adult whiting generally begin to disappear
direct observation of individual schools. Ermakov
from the Pacific Northwest in autumn when the wind
(1974) concluded from direct observation of a lead direction shifts and the Davidson Current appears.
school that the northward migration is at speeds of
Adult whiting also make seasonal inshore-offshore
5-1 1 k d d .
migrations. Ermakov (1974) reports that in spring and
Ermakov (1974) hypothesized that the timing of the early summer whiting schools concentrate over the
spawning migration was linked to the seasonal appear-
continental slope. By mid-June, a large portion of the
ance of the Davidson Current off the Oregon-
stock moves inshore to depths less than 100 m. Later,
Washington coast. Analysis of the movement of the
in early August, whiting move offshore, and by mid-
Soviet fishing fleet in relation to Bakun's (1973, October they begin to migrate southward for spawn-
8 Or
1
ing. These observations of bathymetric migrations are
supported by data in Alton (1972) showing that the
40.0-
w
average depth of catches in bottom trawls decreased in
early summer and increased in autumn (Figure 7).
These movements are similar to the dynamics of the
0 5.0 10.0 15.0 20.0
30.0- INCRWENTS California Undercurrent, which is located over the
continental slope in spring and spreads over the shelf
in early summer (Huyer et al. 1975; Huyer and Smith
1976). Further research on the migration of whiting in
relation to ocean currents is needed and would be of
value to stock assessment and management efforts.
Adult whiting also migrate on a diurnal schedule.
10.0-
2 Fish are dispersed from near surface to 20-m depth at
night (10 p.m. to 3 a.m.). They descend quickly at
dawn and form schools. At night they rise to the sur-
01 face again in 30-40 min (Nelson and Larkins 1970;
0 25 .O 50 .O 75.0 100.0 125.0 150.0
INCREMENTS Ermakov 1974). These diurnal migrations have been
Figure 5. The growth of larvae caught off southern California determined from
compared to the migrations of their primary prey,
otolith increments. A Gompertz curve was fitted to the data, Y = 1.72 euphausiids, as a causal mechanism (Alton and Nelson
'exp[3.15 '(/ - exp [-0.02624'Xl)1.Insef?: dailygrowthforthefirst 20days
was better fitted with a straight line (Y = 2.75 + 0.16x) (from Bailey,
1970). As noted previously, spawning whiting do not
in press). appear to migrate vertically.

88
BAILEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFl Rep., Vol. XXIII, 1982

TABLE 5

jool Parameters of the von Bertalanffy Growth Equation

Source and sex


classification Ira k 1"

Dark (1975)
Male 56.29 0.39 0.20
Female 61.23 0.30 0.01
Male, female combined 60.85 0.30 0.03
Francis (1982)
cl Male, female combined 55.40 0.26 - 1.61
cl

0' 1 Individual males may reach 66 cm, and some females


1 2 3 4 5 6 7 8 9 1 0 1 1 1 2
may reach 80 cm in length. Growth in length was
tlONTH analytically described by the von Bertalanffy growth
Figure 7. The average depth of Pacific whiting bottom-trawlcatches by month, equation:
plotted from data in Alton (1972).
lt = 1, (/-e - k(t-10))
Schooling. Pacific whiting form schools in daytime where
near the bottom. Schools are sometimes shaped in It = body length at age t
bands composed of distinct clusters (T. Dark, Nor- and 1 00, k , and to are parameters of the curve.
thwest and Alaska Fisheries Center, Seattle, WA Table 5 gives values of these parameters estimated
98112, pers. comm.) whose long axes are often for Pacific whiting. Francis (1982) found that between
parallel to isobaths (Nelson and Larkins 1970). ages 3 and 7 growth in length is not uniform through-
Soviets reported that schools may be from less than out the main feeding season (April-October) and that it
0.5 km up to 20 km in length and 0.25 to 3.2 km in appears to reach a maximum during midsummer
width. Above the shelf, schools are, in general, within (June-August).
20 meters of the bottom and are 6-12 m thick. Often The length-weight relation empirically fits the fol-
the underside of a school is 2 m off bottom. Quite a bit lowing equation:
of variability in school size, depth, and structure is
observed (M. Nelson, Northwest and Alaska Fisheries W=aP
Center, Seattle, WA 98112, pers. comm.), and school where
characteristics are more variable and less oriented over W = weight in grams, and
the continental slope than over the shelf (Nelson and 1 = length in centimeters.
Larkins 1970). Table 6 gives estimates of a and b for Pacific whiting.
Ermakov (1974) concluded that schools are formed By age 3, males have grown to between 50 and 60% of
of similar-sized fish. He reports densities of 15-19 their total weight at age 1 1, and females to between 40
fish/1000 m3 in daytime and less than 1 fish/1000 m3 and 50% of their total weight at age 11. Males attain
at night. Spawning schools of whiting form dense an average weight of between 900 and 1000 g by age
aggregations in the pelagic layer, ranging in depth 11 and females between 1100 and 1200 g. Francis
from 100 to 500 m (Stepanenko6; Ermakov 1974; Nel- (1982) found that growth in weight is markedly sea-
son and Larkins 1970). Stepanenko7 reported one sonal. During the winter spawning season (November-
school of spawning whiting that was 4.2 miles long March), adults between ages 4 and 11 lose a minimum
and had a biomass of 81 thousand MT. of between 5 and 10% of their total body weight, and
Age and growth. Age compositions of commercial during the feeding season (April-October) adults be-
catches are determined from annual growth patterns tween ages 4 and l l gain a minimum of between l l
observed from otoliths. The primary source of data on and 30% of their initial body weight. Francis (1982)
Pacific whiting age and growth comes from the
analysis of commercial age compositions (Dark 1975; TABLE 6
Francis 1982). Growth in length is rapid during the Parameters of the Weight-Length Equation w = a P
first 3 years, then it slows and approaches an asymp- Source and sex
tote in the oldest ages (10-13 yr). At about 4 years of classification U b
age, females grow noticeably faster, and by age 11 Dark ( 1975)
may average 32 cm larger than males (Dark 1975). Male ,034682 2.55618
Female ,020444 2.69509
%e footnote 2 on p g e 82.
Francis ( 1982)
Male, female combined .001815 2.13343
'See footnote 3 on page 82.

89
BAILEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFl Rep., Vol. XXIII, 1982

TABLE 7
The Percent Occurrence of Food Types in the Diet of Pacific Whiting Determined by Soviet Scientists.

Washington-Oregon California
Month Month
Food type 5 6 9 10 3 4 6 5 11
Euphausiids 81.6 99.6 40.2 1.2 83.3 83.1 99.8 84.0 93.5
Shrimp 4.9 0.4 1.3 17.3 - - - - 15.2
Squid - - - 1.3 - - - - -
Fish 14.3 -
12.4 37.7 5.6 0.9 1.4 1.4 10.9
*Ermakov, Y., and A.M. Kharchenko. 1976. Biological characteristics of Pacific hake and the estimation of its abundance in 1975. Unpubl. manuscr. Pacific
Scientific Research Institute of Marine Fisheries and Oceanography (TINRO), Vladivostok, USSR.

TABLE 8
The Percent by Weight of Food Types in the Diet of Pacific
Whiting Determined by Polish Scientists* in Summer 1979

Region
Food type Eureka Columbia Vancouver
Euphausiids 94.2 94.0 85.6
Juv. rockfish 1 .o 1.6 -
Pacific hemng - - 5.9
Juv. Pacific herring - - 6.6
Osmerids - 0.4 -
Pacific whiting 0.5 - -
Sablefish - 2.0 0.1
Flatfish - 0.4 -
L
30
I
40
,
50 GO
I 1
70
1
80
1
90 100 I10
I
120 Squid 0.7 - -
Age l y r i Shrimp - 1.6 -
Figure 8. The annual growth in weight of Pacific whiting (from Dark 1975, solid Other fish 3.2 - 1.7
lines) compared to seasonal estimates of growth by age dass (from Francis, Other 0.4 - 0.1
unpubl. manuscr., dotted lines).
*Jackowski, E. 1980. Biological characteristics of Pacific whiting from
Polish surveys of the west coast of the U.S.A. and Canada in 1979.
also found that to accurately represent the seasonal Unpubl. manuscr., presented at :he U.S.-Poland bilateral meetings, 1980.
dynamics of growth a separate weight-length equation
was needed for each age. Figure 8 gives a comparison in frequency of occurrence in the diet in autumn com-
of the weight-age relationships arrived at by Dark pared to summer (Table 7). Shrimp and fish some-
(1975) for 1964-69 and Francis (1982) for 1976-80. times occur frequently. Jackowskiy found that in
Feeding. The feeding behavior of Pacific whiting summer off Vancouver Island, Pacific herring were
has been studied by several investigators, but a com- important in the diet of whiting (Table 8); and in
prehensive seasonal and geographic examination of northern California waters where adult and juvenile
feeding is lacking. distributions overlap, cannibalism is often observed
Adult whiting probably do not feed on the spawning (T. Dark, pers. comm.).
grounds (Tillman 1968) but begin to feed “raven- Livingston (“The Feeding Biology of Pacific
ously” during the postspawning migration north (Er- Whiting, in review) found that Pacific herring were

makov 1974). In summer, whiting are observed to an important component in the diet of whiting off
feed at night towards the surface (Alton and Nelson Oregon-Washington in 1980, composing almost 70%
1970); however, if patches of prey are abundant near of the diet (by weight) of whiting greater than 55 cm,
bottom, whiting may remain there at night to feed and 50% of the diet of whiting less than 55 cm. Alton
(Ermakov 1974). and Nelson (1970) found that in the spring and sum-
There are apparent geographic, seasonal, annual, mer of 1965 and 1966, euphausiids, mostly
and size-specific differences in feeding behavior. The Thysanoessa spinifera, composed 57% of the biomass
most frequently occurring prey items in the summer of whiting stomach contents. Fish, mostly deepsea
diet are euphausiids and Pacific sand lance, Ammo- smelts or Osmeridae spp., composed another 34% of
dytes hexapterus, off Vancouver Island (Outram and stomach contents.
Haegele 1972) and euphausiids and shrimps from Gotshall’s ( 1969a) study demonstrated considerable
California to Washington (Alton and Nelson 1970; seasonality in the diet of whiting off northern Califor-
Gotshall 1969a). Ermakov and Kharchenko8 found nia. Crustaceans, which are the major food in spring
that off Washington and Oregon euphausiids decrease
’Jackowski, E. 1980. Biological characteristics of Pacific whiting from Polish sur-
veys of the west coast of the U.S.A. and Canada in 1979. Unpubl. manuscr., pre- ’
8See footnote 5 on page 8 5 . sented at the U.S.-Poland bilateral meetings. 1980.

90
BAILEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

40 ,
t M l S C INVERTEBRATES
I
r0
t-
‘01
1964
v)
80 \ 1965

- a
CRAB
40
5 0 1966
Y -

FISH
/A-
#I’
I
cn
IO- 20- 30- 40- 50- 6 0 - 70-

19 29 39 49 59 69 89
801
EUPHAUSIIDS LENGTH
a 4G
Figure 11. The number of ocean shrimp per stomach of Pacific whiting off
- ~
1
Vancouver Island (data from Gotshall 1969b).

shrimp is a significant factor in shrimp abundance has


provoked some controversy. Catches of shrimp off the
RONTH
Oregon-Washington coast have increased significantly
Figure 9. The percentage (by volume) of different food types in the diet of
Pacific whiting by month (data from Gotshall 1969a).
since the late 1960s, and this increase appears corre-
lated to the harvest of whiting (Figure 12). It has been
and summer, decline in the diet in winter, and are hypothesized that removing large whiting by fishing
replaced by fish as the dominant food (Figure 9). In has reduced predation pressure on the shrimp popula-
spring and summer an average of 50-60% of the tion. This same trend has occurred off the California
stomach contents of whiting was ocean shrimp; how- coast. However, other factors, such as increasing
ever, these results should be viewed conservatively fishing effort or normal changes in abundance, cannot
because the sample size was small. be ruled out as responsible for the increase in shrimp
Larger whiting more frequently eat fish and less catches, and the question of a whiting-shrimp interac-
frequently eat euphausiids compared to smaller whit- tion deserves more rigorous examination. Francis
ing (Figure 10). Larger whiting also appear to con- (1982) briefly addresses this issue.
sistently eat more shrimp than smaller whiting (Fig-
ure 11).
The question of whether whiting’s feeding on ocean I
1 g 60 1
.60

>
0
.40
8
3
01 0
w
cc
U 113
.20

r
c

0
42-51 52-61 62-71

LENGTH (CM)
Figure 10. Frequency of occurrence of prey types in different Pacific whiting Figure 12. Shrimp catches off the Oregon-Washington coast and catches Of
size classes off Vancouver Island (data from Outram and Haegele 1972). Pacific whiting.

91
BAILEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

TABLE 9
Distribution of Pacific Whiting Biomass by INPFC Area from Three Northwest and Alaska Fisheries Center Surveys

Monterey
and
Year Source Vancouver Columbia Eureka Conception Total
Metric tons
1975* Midwater 3,791 165,941 25,596 42,020 237,348
Percentage 2 70 11 18
Bonom 667 189,630 7,222 10,107 207,626
Percentage - 91 3 5
Total 4,458 355,571 32,818 52,127 444,974
Percentage 1 80 7 12
1977 Midwater 343,821 316,440 360,944 108,087 1,129,292
Percentage 30 28 32 10
Bottom 6,560 32,917 9,501 20,662 69,640
Percentage 9 47 14 26
Total 350,381 349,357 370,445 128,749 1,198,932
Percentage 29 29 31 11
1980 Midwater 322,335 260,476 182,783 578,841 1,344,435
Percentage 24 19 14 43
Bottom 16,678 16,938 13,579 127,647 174,832
Percentage 10 10 8 73
Total 339,013 277,404 196,362 706,488 1,519,267
Percentage 22 18 13 47
*1975 areas do not correspond to 1977 and 1980 survey areas

Competitors. Competitors with Pacific whiting for lected from California sea lion scats at the Farallon
food resources are numerous. Among competitive Islands for 4 yrs, 1974-78, and describe the seasonal I
fishes are other Gadidae; flatfish, Pleuronectidae; and annual dynamics of sea lion feeding on Pacific
soles, Bothidae; smelts, Osmeridae; Cottidae; Pacific whiting. Sea lions fed most heavily on whiting,
herring; albacore, Thunnus alalunga; Hexagram- primarily juveniles, in spring and summer and may
midae; lingcod, Ophiodon elongatus; Myctophidae; consume about 185 thousand tons each year.
rockfish, Scorpaenidae; sablefish, Anoplopoma fim-
bria; and Salmonidae. Numerous marine mammals POPULATION DYNAMICS
may also be competitors, including the rough-toothed
Size of Stocks
dolphin, Steno bredanensis; gray whale, Eschrichtius
The abundance of the Pacific whiting stock has been
robustus; minke whale, Balaenoptera acutorostrata;
assessed from trawl-hydroacoustic surveys. In 1980
Bryde's whale, B . edeni; sei whale, B . borealis; fin
the abundance of whiting from central California to
whale, B. physa1us;blue whale, B . musculus; hump-
southern Vancouver Island over the continental shelf
back whale, Megaptera novaeangliae; and right
was estimated by scientists of the Northwest and
whale, Balaena glacialis (Fiscus 1979).
Alaska Fisheries Center to be 1.52 million metric tons
Predators. Predators of Pacific whiting reported in
the literature include the great white shark, Carcharo- (MT). Most of this biomass was composed of
juveniles off the coast of central California. From
don carcharias; soupfin shark, Galeorhinus zyop-
similar surveys in 1975 and 1977, the stock biomass
terus; Pacific electric ray, Tetranarce californica;
was estimated at 0.44 million and 1.20 million MT
bonito, Sarda chiliensis; albacore; bluefin tuna, Thun-
(Table 9). Based on earlier surveys by the U . S .
nus thynnus; rockfish; sablefish; lingcod; dogfish,
Bureau of Commercial Fisheries, Alverson (cited in
Squalus acanthias; and arrowtooth flounder,
Atheresthes stomias (Best 1963; Nelson and Larkins
Tillman 1968) calculated about 0.68 million MT of
whiting. Estimates of whiting abundance based on
1970; Pinkas et al. 1971). Marine mammals that feed
hydroacoustic methods (Kramer and Smith 1970; Dark
on whiting include the California sea lion, Zalophus
et al. 1980), however, have limitations. Critical prob-
californianus; northern elephant seal, Mirounga an-
lems include (1) the difficulty in calibrating target
gustirostris; Pacific whiteside dolphin, Lagenorhyn-
chus obliquidens; killer whale, Oreinus orca; Dall strength, (2) the failure to identify species by acoustic
porpoise, Phocoenoides dalli; sperm whale, Physeter signals, and (3) the detection of whiting near the
macrocephalus; northern sea lion, Eumetopias bottom (Thorne'O). Regardless of these problems,
jubatus; and northern fur seal, Callorhinus ursinus
"'Thome, R.E. Assessment of population abundance by echo integration. SCOR,
(Fiscus 1979). Working Group No. 52. Symposium on assessment of micronekton, April 27-30,
Bailey and Ainley (1982) analyzed otoliths col- 1980.

92
BAILEY ET AL.:PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

whiting offer one of the more optimum circumstances 1966 1968


100
for hydroacoustic assessment compared with many
other species. 1961

Soviet scientists have determined that the average 1961

biomass of whiting from 1967-73 was 1.36 million 50


MT (Efimov", Ermakov and Kharchenko12, Volog-
din"). They estimated 1.40 and 1.86 million MT of
whiting in 1974 and 1975, respectively. The Soviets 0
conducted two surveys in 1979 with resulting esti- .-
c
0

mates of 1.20 and 2.88 million MT. E 100 1977 1978


Estimates of spawning biomass of whiting deter-
mined from egg and larval surveys are considerably
higher than those stated above. Ahlstrom (1968) cal- 1970
culated that the spawning biomass of whiting was l . 8 1973 1970

to 3.6 million MT. StepanenkoI4 estimated that the


spawning biomass of whiting was 2.4 million MT in
0
1977 and 2.65 million MT in 1979.
Estimates of spawning biomass from egg and larval 1 3 5 7 9 11 1 3 5 7 9 1 1
Age ( y r l
surveys are extremely crude in the case of whiting
Figure 13. Age compositions of Pacific whiting caught off Oregon and
because: the size composition and fecundity of the Washington from research surveys (1966 and 1968) and from U.S. observer
spawners is relatively unknown; the stage duration of fishery data (1977 and 1978) (from Bailey 1981a.b).
eggs and larvae was not used for these approxima-
tions; fecundity schedules of adults are based on very supply, predation, and larval transport. Although can-
little data; and it is unknown whether whiting are mul- nibalism is sometimes observed, it is probably fairly
tiple spawners. Estimates are further confounded by low because the majority of adult whiting spend a
the extreme patchiness of eggs and larvae. In spite of limited time in the spawning area (1-3 mo). However,
these problems, ichthyoplankton surveys are useful for Sumida and Moser (1980) found that some large lar-
assessing the relative abundance of the stock, and the vae eat smaller larvae, and there are a few reports of
California Cooperative Oceanic Fisheries Investiga- adult predation on juveniles.
tions (CalCOFI) ichthyoplankton surveys conducted Bailey (in press) found that the food requirement of
since 1950 have been useful in monitoring changes in whiting larvae is low because of relatively low growth
the spawning potential of the population. The spawn- and metabolic rates. The large mouth size of larvae
ing stock appears to have decreased in the late 1960s enables first-feeding larvae to ingest a wide spectrum
and early 1970s compared with earlier years, but has of food particles, including juvenile and adult
recently increased to previous levels (Stauffer and copepods (Sumida and Moser 1980). Bailey (in press)
Smith 1977). calculated that a first-feeding whiting larva can satisfy
growth and metabolic requirements by ingesting 3 1
Recruitment copepod nauplii, 6 small calanoid adult copepods, or
Because of the spatial distribution of age classes, 0.6 Culunus copepodites per day. By comparison, a
recruitment of the exploited stock occurs at 3-6 years first-feeding northern anchovy larva, with its small
of age depending on the location of fishing. Inter- mouth, must capture at least 200 Gymnodinium cells
annual variations in recruitment are great, as exempli- per day to satisfy metabolic (excluding growth) re-
fied by the dominance of the age composition of the quirements alone (Hunter 1977). It was concluded that
stock by strong year classes for several consecutive starvation from first-feeding failure is probably not as
years (Figure 13). variable for whiting larvae as it appears to be for
The factors most often considered to affect repro- northern anchovy and that whiting may not be as de-
ductive success of marine fishes are cannibalism, food pendent on finding patches of prey as are northern
anchovy (Lasker 1975).
"Efimov, Y . N . 1974. The size of stocks and status of fishery of Pacific hake. Predation on eggs and larvae is a difficult problem
Unpubl. manuscr. Pacific Scientific Research Institute of Marine Fisheries and
Oceanography (TINRO), Vladivostok, USSR. to assess and is poorly understood. A wide variety of
I2See footnote 5 on p g e 85. invertebrate organisms are capable of feeding on
"Vologdin, V. 1980. Results of the hydroacoustic surveys with trawlings off the whiting eggs and larvae. Yolk-sac stages are most
Pacific coast of North America in 1979. Unpubl. manuscr. Pacific Scientific Research
Institute of Marine Fisheries and Oceanography (TINRO), Vladivostok, USSR. vulnerable to predation by invertebrates (Bailey and
I4See footnotes 2 and 3 on p g e 82. Yen, in press). Predation by invertebrates may be

93
BAILEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

Y = 34.56 + 1.66X ( R = .70; P = ,001) of larvae offshore is expected to be detrimental to


+ survival. In fact, Ekman transport during the spawning
months is negatively correlated to year-class strength
(Figure 15). Further work must be done on the sur-
+ vival of larvae swept offshore to test this hypothesis.
Temperature may also influence recruitment, possi-
bly by the predation-temperature interactions noted
above and the previously described influence of temp-
erature on the location of spawning. The average
winter temperature and Ekman transport in a multiple
regression model account for 68% of the observed
variation in an index of year-class strength (Bailey
1981 a,b). There is no apparent relationship between
spawning biomass of the population and recruitment.

+ Mortality
20 - Estimates of annual instantaneous natural mortality
i
t + rates range widely. These estimates, as well as several
estimates for fishing and total mortality rates are pre-
sented in Table 10. A cohort analysis performed by
Francis (1982) on the 1973-80 catch-by-age data gives
estimates of age-specific fishery mortality (catcha-
bility) as well as recruitment of the exploited stock at
age 3.
important in cold years when development is slow
through the stages most vulnerable to predation. TABLE 10
Oceanographic conditions appear to play a major Estimates of Annual Instantaneous Mortality Rates
role in the recruitment of Pacific whiting (Bailey 198 1 of Pacific Whiting
a,b). The offshore distance of larvae is apparently Investigators Males Females Both Sexes
positively correlated to indices of wind-driven Ekman M : natural mortality
transport (Figure 14). Although there is a good deal of Tillman (1968) 0.72 0.62 x=0.67
variability in this relationship, it is statistically sig- Nelson and Larkins (1970) OS6
Efimov (1974)' 0.3.5
nificant and indicates that larvae may be transported PFMC2 0.30-0.60
offshore in years of high npwelling. Since the juvenile Jackowski (1980)3 0.30
nursery is inshore over the continental shelf, advection Ehrich, et al. (1980) 0.56
Low (1978)4 0.50
Francis (in prep.) 0.19-0.86
2.5 Y = 1,449 - 0,020X (R = -0.74;P < 0.001) (variable
age-
specific
x
a 1961 natural
;2.0 mortality)
-
z F: fishing mortality
Efimov ( 1974)' 0.30
co Ehrich, et al. (1980) 0.67
cn
'= Z: total mortality
1.5 Efimov ( 1974)l 0.65
a Ehrich et al. (1980) 1.23
'Efimov Y . N . 1974. The size of stocks and status of fishery of Pacific
hake. Unpubl. manuscr. Pacific Scientific Research Institute of Marine
I A Fisheries and Oceanography (TINRO), Vladivostok, USSR.
*Pacific Fishery Management Council. 1980. Pacific coast groundfish
plan. Draft report. Pacific Fishery Management Council, 526 S.W. Mill
St., Portland, OR 97201,
0.5 L--' 3Jackowski, E. 1980. Biological characteristics of Pacific whiting from
-30 -20 -10 0 30 20 30 Polish surveys of the west coast of the U.S.A. and Canada in 1979.
Unpubl. manuscr., presented at the U.S.-Poland bilateral meetings, 1980.
UPWELLING INDEX 4Low, L.L. 1978. Hake natural mortality and yield potential. Unpubl.
Figure 15. Log of the year-class index against the January upwelling index manuscr. Northwest and Alaska Fish. Cent. Natl. Mar. Fish. Serv.
(from Bailey 1981a,b). NOAA, 272.5 Montlake Blvd. E., Seattle, WA 98112.

94
BAILLY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOl-1 Rep., Vol. XXIII, 1982

Dynamics of the Population 1879 (Jow 1973). This fishery has been rather insig-
Over the past 200 years the Pacific whiting popula- nificant, with catches in the range of 200-500 MT/yr.
tion has experienced some major changes in abun- However, in recent years the domestic fishery has be-
dance (Soutar and Isaacs 1974).Based on an analysis come important: U.S.-foreign joint-venture fishing
of fish scales deposited in sediments, at the turn of the caught 9 thousand MT and 28 thousand MT in 1979
last century the population was almost an order of and 1980, respectively.
magnitude larger than recent abundance levels. These Historical effort statistics for the fishery, excluding
changes in abundance have been correlated to changes Canadian waters, were calculated from weekly aerial
in abundance of the northern anchovy (Soutar and surveillance data from the NMFS Enforcement Divi-
Isaacs 1974) and are inversely correlated to offshore sion (supplied by Bill Dickenson, NMFS, Northwest
Ekman transport (Bailey 1981a,b). Regional Office, Seattle, WA). Effort for two classes
Several mathematical models have been constructed of vessels-large Soviet BMRT stem trawlers and
that simulate changes in the whiting population. These smaller Soviet SRT side trawlers (see “Technical As-
include models by Francis (1982), Francis et al. pects ” below)-were calculated in vessel-days on the
(1982), Bernard (Oregon State University, Newport, fishing grounds. Effort by the SRT trawlers was
OR), Stevens and Goodman (Scripps Institution of greatest in 1966 and declined steadily (Table 12).
Oceanography, La Jolla, CA), Tillman (1968), and Effort by the BMRT trawlers was greatest in 1975
Riffenburgh (1969). and 1976.
To obtain a rough estimate of overall catch per unit
THE PACIFIC WHITING FISHERY of effort (CPUE) for the foreign fishery in U.S. wat-
ers, SRT effort was converted to effective BMRT ef-
Historical Catches and Effort
fort by assuming a relative fishing power of
Pacific whiting has been the target of a large foreign
P ~ R T= 0.3 1 from the ratio of average horsepower of
fishery off the west coast of the United States and
SRT vessels to BMRT vessels (1 150 HP).
Canada (Table 11). A Soviet fishery for whiting began
Catchhtandard BMRT day indicates that the highest
in 1966 with a catch of 137 thousand MT. From
rates occured in 1967 and from 1977 to 1980. Since
1973-76 Poland, West Germany, East Germany, and
the latter period coincides conspicuously with passage
Bulgaria joined fishing operations for Pacific whiting.
of the Magnuson Fishery Conservation Management
Reported catches peaked in 1976 at 237 thousand MT.
Act of 1976 (MFCMA) and the onset of intense ob-
The average annual all-nation reported catch from server coverage, these statistics indicate that actual
1966 to 1980 was 162 thousand MT. (These catches
catches were possibly underreported from 1968 to
were compiled from data at the Northwest and Alaska
1976.
Fisheries Center.)
A small domestic fishery for whiting, used in the
manufacture of pet food, has existed since at least TABLE 12
Historcial Effort Statistics for the U.S.S.R.-Poland Foreign
Pacific Whiting Fishery (Solely) in U.S. Waters
TABLE 11 ~ ~ ~~

Annual All-Nation Catches of Pacific Whiting (x lo3 MT) in US. Standard


and Canadian* Waters. BMRT
BMRT RST days Catch CPLE
Domestic/ Year vessel days vessel days Pqr=O 31 (1000 MT) (MTIBMRTday)
Year U.S.S.R. Poland ioint venture Other Total 1966 2,670 14,490 7,128 137.0 19.2
1966 137.0 - - - 137.0 1967 2,730 10,350 5,915 195.1 33.0
1967 206.1 - - - 206.1 1968 5,677 2,079 6,317 68.0 10.8
1968 103.8 - - - 103.8 1969 5,607 1,589 6,096 109.0 17.9
1969 161.8 - - 0.12 161.9 1970 7,847 658 8,049 200.8 24.9
1970 226.2 - - 2.3 228.5 1971 7,245 65 1 7,445 146.7 19.7
1971 151.8 - - 1.4 153.2 1972 5,131 518 5,290 111.3 21 .0
1972 150.8 - - 0.4 151.2 1973 5,904 - 5,904 141.1 23.9
1973 143.8 2.0 - 5.1 150.8 1974 7,717 - 7,717 201.1 26.1
1974 173.7 44.3 - 8.4 226.5 1975 10,401 - 10,401 196.9 18.9
1975 155.4 57.2 - 5.1 217.7 1976 6,917 - 6,917 177.8 25.7
1976 158.0 25.7 - 53.0 236.8 1977 4,076 - 4,076 127.2 31.2
1977 111.0 19.5 - 1.9 132.4 1978 2,779 - 2,779 96.9 34.9
1978 70.9 27.3 2.7 3.4 104.2 1979 4,452 - 4,452 114.9 25.8
I979 96.8 22.3 13.1 3.6 135.9 1980 1,553 - 1,553 44.0 28.3
1980 0.1 49.0 40.8 0.8 90.7 Assumptions - Ps~=0.31
*Zyblut, E. 1981. Dept. of Fisheries and Oceans, Govt. of Canada, Van- PBMRT(Poland)=1.OO
couver, B.C. Personal communication. CPUE = catch per unit effort

95
BAILEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

Technical Aspects TABLE 13


As the major country fishing for whiting, the Soviet Catch (Percentage) by Depth Strata of Pacific Whiting Taken by
Foreign Trawlers in 1979
Union has improved its whiting fleet considerably. In
1966 the fleet was mainly medium-sized side trawlers Depth (m) 0-99 100-199 200-299 300-399 400-499 >500
(SRTs) of about 500 gross tons. The proportion of Percentaee 15.7 47.7 26.2 6.9 2.3 1.2
large stern trawlers with freezing capacity (BMRTs) From: French, R., R. Nelson Jr., and J . Wall. 1980. Observations of
has gradually increased to replace the side trawlers. foreign and joint venture fishing fleets off the coast of Washington, Ore-
The typical BMRT is 3170 gross tons, has a crew of gon, and California, 1979. Unpubl. rep. Northwest and Alaska Fish.
Cent., Natl. Mar.Fish. Sew., NOAA, 2725 Montlake Blvd. E., Seattle,
22-26, and uses a midwater trawl with a headrope WA 981 12.
length of 97 m. The daily production capacity is 30-50
MT of frozen fish and 20-35 MT of meal and oil. under review. A conservative estimate of maximum
Support vessels in the fishery include factory ships, sustained yield in the plan is 175.5 thousand MT. The
refrigerated transports, oil tankers, personnel carriers, FMP specifies geographic and seasonal restrictions,
tugs, and patrols. mesh size, incidental catch levels, and an optimal
The Soviet fishery is a well-coordinated expedition, yield (in the form of quotas). Under the MFCMA only
and acoustics are used to guide the net over fish con- the Soviets and Poles have been granted licenses as the
centrations. Prior to 1976, about 100 BMRTs would major foreign interests that may fish for Pacific whit-
typically participate in the fishery (Pruter 1976), but ing. Recently. U. s. fishermen have become involved
lately the fleet has been reduced to about 39 large stern in the whiting fishery through joint ventures in which
trawlers. In early years of the fishery most whiting U. S . trawlers harvest whiting for delivery to foreign
were filleted, and small fish were reduced to meal.Re- processing vessels.
cently the average size of hake has become considera- Francis et al. (1 982) present a management analysis
bly smaller, and an increasing proportion of the catch of the Pacific whiting fishery in which a policy al-
is frozen whole. gorithm is developed that aims to use strong year clas-
The foreign fishery is closely tied to the migratory ses in a practical and efficient manner while protecting
movements of the whiting population. Historically, the stock when it is in poor condition and environ-
the fishery began in waters off Oregon in April and mental conditions do not appear conducive to imme-
moved northward as schools made their way up the diate improvement.
coast in summer. This was documented from aerial
sightings of the Soviet fishery (Figure 16). In au- Effects of the Fishery on the Population
Commercial fisheries may affect the abundance and
tumn, as fishing activity halted, whiting began to
recruitment of marine fish populations in several
migrate offshore and southward for spawning. More
ways. Besides reducing the total spawning biomass of
recently, fishing has been restricted by treaty to the
the population, removing a stock’s largest and oldest
period from June until October. Based on aerial sur-
fish also (1) lowers the quality of the spawning pro-
veillance records and Ermakov’s (1974) analysis of
duct if offspring from smaller fish are less fit (Hempel
the fishery, rich fishing grounds appear to be as-
1979); (2) reduces the number of age classes that con-
sociated with prominent geographical sites such as
tribute to spawning; thus the maintenance of healthy

-
banks, sharp “curves” in the continental slope, and
levels of spawning stock depends on successful re-
canyons. Especially productive grounds are found
near the Heceta Banks, Yaquina Head, Cape Flattery,
Cape Blanco, and Destruction Island. Most fish are
caught in depths of 100-199 m (Table 13).
Ln 1 50

Management
Prior to implementation of the MFCMA in 1977,
the foreign fishery was managed by bilateral agree-
ment. Since 1977 management has been directed by a
Preliminary Management Plan (PMP)IS for groundfish
prepared by the Department of Commerce. Sub-
sequently, the Pacific Fishery Management Council 0
has prepared a fisheries management plan (FMP) for 4 5 6 7 8 9 1 0 1 1 1 2
groundfish, including whiting, which is currently MONTH
‘sPacific Fishery Management Council. 1980. Pacific coast groundfish plan. Draft Figure 16. The average monthly number of foreign vessels fishing Pacific
report. Pacific Fishery Management Council, 526 S.W. Mill St., Portland, OR whiting sighted in aerial surveys, 1967-72 (squares) and the average per-
97201. centage of the catch in 3-m0 periods for the same years.

96
BAILEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

cruitment from fewer age classes (Smith 1978); and A 1951-66 Y = 0,23X - 4.80 (R = .63; P < ,01)
(3) changes the distribution of spawning if the popula- 1967-75 Y = 0.1OX - 1.42 (R = ,84; P < ,O1)
tion stratifies by size or age on the spawning ground.
The spawning potential of the whiting population
loa
has had no discernible effect on recruitment from -
c
1960-75 (Bailey 1981a,b), partly because of the m
C

overwhelming effects of environmental factors. For .


example, the strong 1961 year class arose from an
-._a
(0
80

extremely low spawning stock, but under favorable E


.
environmental conditions it became a very strong year 60
class. Similar situations gave rise to the strong 1970 3
E
and 1973 year classes. g 40
-
3
The long life of Merluccius spp., as well as of other 0,
gadids, is probably an adaptation to stabilize the stock C -
4
from the effects of extreme recruitment variability,
t
L

and reduction in the number of spawning age classes 1


d
by heavy fishing must be a destabilizing influence. In O L
a population with fewer age classes, the probability of 21 22 23 24 25
a stock collapse would increase if recruitment failure Sea surface temperature ("C)
occurred in a succession of years. This type of interac- Figure 17. Regressions of the percentage of Pacific Whiting larvae off swthern
tion appears to have influenced the recruitment of California compared to Baja Californiaagainst the mean January-March sea
surface temperature off Baja Californiafor the pre- and postfishing periods,
other stocks. In an analysis of the population dynamics 1951-66 and 1967-75.
of the Pacific sardine, Sardinops sagax, Murphy
(1968) concluded that after the number of spawning R. Lasker reviewed earlier versions of the manuscript
age classes was reduced by fishing, several years of and made helpful comments. We also appreciate the
recruitment failure caused catastrophic population de- encouragement of W. Wooster and R. Marasco.
clines.
Since the mid-l960s, a change in the spawning LITERATURE CITED
grounds of Pacific whiting has occurred. Larvae have Ahlstrom, E.H. 1959. The vertical distribution of fish eggs and larvae off
California and Baja California, Fish. Bull., U.S. 60: 107- 146.
become much less abundant off Baja California and
-. 1968. An evaluation of the fishery resources available to Califor-
more abundant off central California compared with nia fishermen. In D. Gilbert (ed.), The future of the fishing industry of
earlier years (Bailey 1980). In addition, the deposition the United States. Univ. Wash. Press, Seattle, p. 65-80.
of scales from young whiting markedly declined off Ahlstrom, E.H., and R.C. Counts. 1955. Eggs and larvae of the Pacific
Baja California from 1965-69 compared with earlier hake, Merluccius productus. Fish. Bull., U.S. 56:295-329.
Alton, M.S. 1972. Characteristics of the demersal fish faunas inhabiting
prefishing periods (Soutar and Isaacs 1974). Smith the outer continental shelf and slope off the northern Oregon coast. In
(1975) first suggested that this change was related to A.T. Pruter and D.L. Alverson (eds.), The Columbia River estuary and
the beginning of an intensive fishery for adults in adjacent ocean waters. Univ. Wash. Press, Seattle, p. 583-636.
1966. He suggested that large adults spawn farther Alton, M.S., and M.O. Nelson. 1970. Food of the Pacific hake, Merluc-
ciusproductus, in Washington and northern Oregon coastal waters. U.S.
south and that harvesting this component of the popu- Fish Wildl. Serv., Circ. 332:35-42.
lation has caused the spawning decrease in the south- Alverson, D.L., and H.A. Larkins. 1969. Status of the knowledge of the
ern end of the range. Further analysis supports an in- Pacific hake resource. Calif. Coop. Oceanic Fish. Invest. Rep.
teraction between the spawning distribution and the 13:24-3 1.
Bailey, K.M. 1980. Recent changes in the distribution of hake larvae:
fishery (Bailey 1980, 1981a,b). Although the spawn- causes and consequences. Calif. Coop. Oceanic Fish. Invest. Rep.
ing location of whiting is related to temperature, the 21: 167-171.
recent change in the distribution of larvae is inde- - . 1981a. An analysis of the spawning, early life history and re-
pendent of-temperature changes. An analysis of cruitment of the Pacific hake, Merluccius producrus. Ph.D. dissertation,
Univ. Wash., Seattle.
covariance indicated significantly different slopes and -. 1981b. Larval transport and the recruitment of Pacific hake,
intercepts for pre- and post-spawning periods (Fig- Merluccius productus. Mar. Ecol. 6:1-9.
ure 17). -. (in press). The early life history of Pacific hake, Merluccius
productus. Fish. Bull., U.S.
ACKNOWLEDGMENTS Bailey, K.M., and P. Ainley. 1982. The dynamics of California sea lion
predation on Pacific hake. Fisheries Research 1:163- 176.
Numerous scientists at the and Bailey, K.M., and J . Yen (in press). Predation by a carnivorous marine
and Southwest Fisheries Centers freely made data copepod, Euchuera elongaru Esterly, on eggs and larvae of the Pacific
available. T. Laevastu, R. Schwartzlose, T. Dark, and hake, Merluccius productus. J. Plankton Res.

97
BAILEY ET AL.: PACIFIC WHITING LIFE HISTORY AND FISHERY
CalCOFI Rep., Vol. XXIII, 1982

Bakun, A. 1973. Coastal upwelling indices, west coast of North America, Gamer, D., and P. Smith. 1970. Seasonal and geographic characteristics
1946-7 I . U.S. Dep. Commer., NOAA Tech. Rep. NMFS SSRF-67 1. of fishery resources. California Current region-111. Pacific hake.
-. 1975. Daily and weekly upwelling indices, west coast of North Commer. Fish. Rev. 32(7):41-44.
America, 1967-73. U.S. Dep. Commer., NOAA Tech. Rept. SSRF- Lasker, R. 1975. Field criteria for survival of anchovy larvae: the relation
693. between inshore chlorophyll maximum layers and successful first feed-
Best, E.A. 1963. Contribution to the biology of the Pacific hake, Merluc- ing. Fish. Bull., U.S. 73:453-462.
cius productus. Calif. Coop. Oceanic Fish. Invest. Rep. 9:51-56. MacGregor, J.S. 1966. Fecundity of the Pacific hake, Merluccius produc-
Dark, T.A. 1975. Age and growth of Pacific hake, Merlucciusproductus. tus. Calif. Fish Game. 52:111-116.
Fish. Bull., U S . 73:336-355. -. 1971. Additional data on the spawning of the hake. Fish. Bull,
Dark, T.A., M.O. Nelson, J. Traynor, and E. Nunnallee. 1980. The U.S. 69x581-585.
distribution, abundance, and biological characteristics of Pacific whit- Murphy, (3.1. 1968. Pattern in life history and the environment. Am. Nat.
ing, Merluccius productus, in the California-British Columbia region 102~391-403.
during July-September 1977. Mar. Fish. Rev. 42(3-4):17-33. Nelson, M.O., and H.A. Larkins. 1970. Distribution and biology of
Erich, S., F. Mombeck, and G . Speiser. 1980. Investigations on the Pacific hake: a synopsis. U.S. Fish Wildl. Serv., Circ. 332:43-52.
Pacific hake stock (Merluccius productus) in the Northeast Pacific.
Outram, D.N., and C. Haegele. 1972. Food of Pacific hake (Merluccius
Arch. Fishereiwiss 30:17-38.
productus) on an offshore bank southwest of Vancouver Island, British
Ermakov, Y.K. 1974. The biology and fishery of Pacific hake, Merluccius Columbia. J. Fish. Res. Bd. Can. 291792-1795.
productus. Ph.D. dissertation, Pac. Sci. Inst. Mar. Fish. Oceanogr.
(TINRO), Vladivostok, USSR (in Russian). Pinkas, L., M. Oliphant, and I. Iverson. 1971. Food habits of albacore,
bluefin tuna, and bonito in California waters. Calif. Fish and Game.
Fiscus, C. 1979. Interactions of marine mammals and Pacific hake. Mar. Fish. Bull. 152.
Fish. Rev. 41(10):1-9.
Pruter, A.T. 1976. Soviet fisheries for bottomfish and hemng off the
Foucher, R.P., and R.J. Beamish. 1977. A review of oocyte development Pacific and Bering Sea coasts of the United States. Mar. Fish. Rev.
in fishes with special reference to Pacific hake (Merluccius productus) 38( 12):1-14.
Can. Fish. Mar. Serv. Tech. Rep. No. 755.
Francis, R.C. 1982. On the population and trophic dynamics of Pacific Riffenburgh, R.H. 1969. A stochastic model of interpopulation dynamics
whiting. U.S. Dep. Commer., Natl. Oceanic Atmos. Admin., Natl. in marine ecology. J. Fish. Res. Bd. Can. 26:2843-2880.
Mar. Fish. Ser., Northwest and Alaska Fish. Cent., Seattle, Wash., Smith, P. 1975. Pacific hake larval distribution and abundance, 1951-
Processed Rep. 82-07, 68 p. 1975. Southwest Fish. Cent. Admin. Rep. LJ-75-83. Southwest Fish.
Francis, R.C., G.L. Swartzman, W.M. Getz, R. Haar, and K. Rose. 1982. Cent., Natl. Mar. Fish. Serv., NOAA, P.O. Box 271, La Jolla, CA
A management analysis of the Pacific whiting fishery. U.S. Dep. Com- 92038.
mer., Natl. Oceanic Atmos. Admin., Natl. Mar. Fish. Ser., Northwest -. 1978. Biological effects of ocean variability: time and space
and Alaska Fish. Cent., Seattle, Wash., Processed Rep. 82-06, 48 p. scales of biological response. Rapp. P-v. Rtun. Cons. Int. Explor. Mer
Frey, H.W. 1971. Pacific hake. In H.W. Frey (ed.), California’s living 173:117-127.
marine resources and their utilization, p. 71-74. Calif. State Dep. Fish Soutar, A,, and J.D. Isaacs. 1974. Abundance of pelagic fish during the
Game. 148 p. 19th and 20th centuries as recorded in anaerobic sediments off Califor-
Gotshall, D.W. 1969a. Stomach contents of Pacific hake and arrowtooth nia. Fish. Bull., U.S. 72:257-274.
flounder from northern California. Calif. Fish. Game 55:75-82. Stauffer, G.D., and P.E. Smith. 1977. Indices of Pacific hake from 1951-
. 1969b. The use of predator food habits in estimating relative 1976. Southwest Fish. Cent. Admin. Reo. LJ-77, Southwest Fish.
abundance of the ocean shrimp, Pandulus jordani. Food Agric. Organ. Cent., Nat’l. Mar. Fish. Serv., NOAA, P.O. Box 271, La Jolla, CA
U.N., Fish. Rep. 57:667-685. 92038.
Hart, J.L. 1973. Pacific fishes of Canada. Fish. Res. Bd. Can., Bull. 180, Sumida, B.Y., and H.G. Moser. 1980. Food and feeding of Pacific hake
740 p. larvae, Merluccius productus, off southern California and Baja Califor-
Hempel, G. 1979. Early life history of marine fish: the egg stage. Univ. nia. Calif. Coop. Oceanic Fish. Invest. Rep. 21:161-166.
Wash. Press, Seattle. Theilacker, G.H. 1981. Changes in body measurements of larval northern
Hickey, a.M. 1979. The California Current system-hypotheses and anchovy, Engraulis mordax, and other fishes due to handling and pre-
facts. Prog. in Oceanogr. 8:191-279. servation. Fish Bull., U.S. 78:685-692.
Holliday, D.V., and H.L. Larsen. 1979. Thickness and depth distribution Tillman, M.F. 1968. Tentative recommendations for management of the
of some epipelagic fish schools off southern California. Fish. Bull., coastal fishery for Pacific hake, Merluccius productus, based on a
U.S. 771489-494. simulation study of the effects of fishing upon a virgin population. M.S.
Hunter, J. 1977. Behavior and survival of northern anchovy, Engraulis thesis, Univ. Wash. Seattle.
mordax larvae. Calif. Coop. Oceanic Fish. Invest. Rep. 19:138-146. U.S. Fish and Wildlife Service. 1970. Pacific hake. U.S. Fish. Wildl.
Hunter, J., and C. Kimbrell. 1980. Egg cannibalism in the northern an- Serv., Circ. 332, 152 p.
chovy, Engraulis mordax, Fish Bull., U.S. 78:811-816. Utter, F.M., and H.O. Hodgins. 1971. Biochemical polymorphisms in the
Huyer, A . , R.D. Pillsbury, and R.L. Smith. 1975. Seasonal variation of Pacific hake (Merluccius productus). Rapp. P-v. Cons. RCun. Int.
the alongshore velocity field over the continental shelf off Oregon. Lim- Explor. Mer 161:87-89.
nol. Oceanogr. 20:90-95.
Vrooman, A.M., and P.A. Paloma. 1977. Dwarf hake off the coast of Baja
Huyer, A., and R.L. Smith. 1976. Observations of a poleward undercur-
California. Calif. Coop. Oceanic Fish. Invest. Rep. 19:67-72.
rent over the continental slope off Oregon, May-June 1975. EOS
57:263. Wooster, W.S., and J.H. Jones. 1970. California Undercurrent off north-
em Baja California. J. Mar. Res. 28:235-250.
Jones, B.W. 1974. World resources of hakes of the genus Merluccius. In
F.R. Harden Jones (ed.), Sea fisheries research. John Wiley & Sons, Wyllie, J.G. 1967. Geostrophic flow of the California Current at the sur-
New York, p. 139-166. face and at 200 meters. Calif. Coop. Oceanic Fish. Invest. Atlas 4.
Jow, T. 1973. Pacific hake length frequencies at California ports, 1963- Zweifel, J., and R. Lasker. 1976. Prehatch and posthatch growth of
1970. Calif. Dep. Fish Game, Mar. Res. Tech. Rep. No. 2. fishes- a general model. Fish. Bull., U.S. 74:609-622.

98

You might also like