CAUPORNIAl
FBH^GAME
California Fish and Game is a journal devoted to the conservation and
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Please direct correspondence to:
Robert N. Lea, Ph.D., Editor-in-Chief
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u
1
0
V
VOLUME 73
OCTOBER 1987
NUMBER 4
Published Quarterly by
STATE OF CALIFORNIA
THE RESOURCES AGENCY
DEPARTMENT OF FISH AND GAME
—LDA—
STATE OF CALIFORNIA
GEORGE DEUKMEJIAN, Governor
THE RESOURCES AGENCY
GORDON VAN VLECK, Secretary for Resources
FISH AND GAME COMMISSION
ALBERT C. TAUCHER, President
Long Beach
ABEL C. GALLETTI, Vice President JOHN A. MURDY III, Member
Rancho Palos Verdes Newport Beach
ROBERT A. BRYANT, Member E. M. McCRACKEN, JR., Member
Yuba City Carmichael
HAROLD C. CRIBBS
Executive Secretary
DEPARTMENT OF FISH AND GAME
PETE BONTADELLI, Acting Director
1416 9th Street
Sacramento 95814
CALIFORNIA FISH AND GAME
Editorial Staff
Editorial staff for this issue:
fVlarine Resources Peter L. Haaker, Robert N. Lea,
and Paul N. Reilly
Wildlife Bruce E. Deuel and William E. Grenfell, Jr.
Editor-in-Chief Robert N. Lea. Ph.D.
195
CONTENTS
Page
Movement and Dispersion of Red Abalone, Haliotis rufescens,
in Northern California Jerald S. Auit and John D. DeMartini 196
The Use of Baited Stations by Divers to Obtain Fish
Relative Abundance Data Daniel W. Gotshall 214
Survival and Recovery Rate Estimates of Northern
Pintail Banded in California, 1948-79 Warren C. Rienecker 230
Management of Midges and Other Invertebrates for Waterfowl
Wintering in California Ned H. Euliss, Jr. and Gail Grodhaus 238
NOTES
Yellowtail Chafing on a Shark: Parasite Removal? Bruce E. Coblentz 244
Atypical Plumage of a Female California Quail J. A. Crawford,
P. J. Cole, K. M. Kilbride and A. Fairbrother 245
BOOK REVIEWS 248
INDEX TO VOLUME 73 250
196 CALIFORNIA FISH AND CAME
Calif. Fish and Came 73(4): 1 9&-2 13 1 987
MOVEMENT AND DISPERSION OF RED ABALONE,
HAUOTIS RUFESCENS, IN NORTHERN CALIFORNIA
JERALD S. AULT
Rosenstiel School of Marine and Atmospheric Science
University of Miami
4600 Rickenbacker Causeway
Miami, FL 33149
and
JOHN D. DEMARTINI
Department of Biological Sciences
Humboldt State University
Areata, CA 95521
Tagging of red abalone in Northern California over an 11 year period showed that
movement and dispersal occurred at high frequency. The high incidence of movement
contributed to continuous population flux. However, not all tagged abalone were ob-
served to have moved. Some abalone exhibited no apparent movement for short pe-
riods of time, and occasionally maintained the same site for prolonged periods. Red
abalone densities apparently remained constant through time even though much
emigration and immigration occurred. Adults tended to reposition on scars, regard-
less if the scars were previously theirs. Stimuli for movement may have been food
supply, although physical disturbance was identified in prompting movement.
INTRODUCTION
Red abalone, Haliotis rufescens, are widely distributed along the coast of Cal-
ifornia and support important commercial and sport fisheries. During the past
two decades, red abalone stocks have declined in abundance throughout major
portions of their range. Declines are attributed to possible overexploitation, hab-
itat degradation, and increasing sea otter predation in the traditionally productive
abalone beds off central California (Leighton et al. 1981, Ault 1985a).
Studies providing insight into the spatial and temporal mechanisms that influ-
ence rates of repopulation are fundamental to the evaluation of haliotid popu-
lation dynamics (Hancock 1979, Clavier and Olivier 1984). Worldwide, several
species of abalone are known to move frequently, and some species for con-
siderable distances ( Newman 1 966, Poore 1 972, Shepherd 1 973 ) . The published
literature on red abalone implies that very little movement or dispersion occurs
(Bonnot 1948, Cox 1960, 1962, Mines and Pearse 1982).
An extensive tagging program of subtidal abalone in Northern California was
undertaken to assess growth, general distribution along the coast, food supply,
movement, and to identify predators and competitors. Ault (1985a) provides a
synopsis of these findings. This report deals with movement and presents both
qualitative and quantitative evidence for movements and dispersion by individ-
ual red abalone.
STUDY SITE
This study was conducted at Point Cabrillo Marine Ecological Reserve, Eort
Bragg, California (Figure 1 ). The reserve is closed to commercial and sport take
Accepted for publication March 1987.
MOVEMENT AND DISPERSION OF RED ABALONE
197
of red abalone. The coastline at Point Cabrillo is highly exposed. Wave action has
eroded the marine terrace such that the coastline is very irregular and bordered
by reefs (Figure 2). Thus, the study site contained various degrees of exposure
and depths resulting in a spectrum of habitats utilized by different size classes of
red abalone.
Pocific
Ocean
Caipor
Point
FIGURE 1. Location of Point Cabrillo marine ecological reserve.
PACIFIC OCEAN
linch-SOyordi
FIGURE 2. Depiction of the eleven tag and release zones developed for studying red abalone
movements at Point Cabrillo marine ecological reserve. Depths indicated are in feet.
1 98 CALIFORNIA FISH AND CAME
METHODS
Longterm Movements
The study began in January, 1971, and continued until 1975. Over this period
3,877 red abalone were tagged by scuba divers. Periodic observations continued
through June, 1982. We divided the subtidal waters and seabed constituting the
Reserve into eleven well-defined zones ( c.f. Figure 2 ). During the course of each
tagging session, within each study zone, most available red abalone greater than
50 mm long were collected. Shorter specimens were generally not taken, be-
cause tagging them fractured the shell between respiratory pores. Abalone were
removed by a metal bar or a lever, placed in a mesh bag and taken to shore. Each
abalone collected was measured for shell length and width in millimeters,
weighed in grams, sexed if possible, and a numbered metal tag attached to stain-
less steel wire was wound through two respiratory pores. While tagging, de-
pending upon prevalent weather conditions, care was taken to keep only a few
abalones out of the water at any time, minimizing shock and exposure. All tagged
abalone were free of deep cuts. Abalone were returned to the area of collection.
Caution was taken to place animals in crevices, on developed "scars" (a clean
area of rock approximately the size of their foot and usually devoid of
macrobiota), or under secure boulders. Subsequent observations and collections
of tagged abalone were made by divers at time intervals of varying length (usu-
ally 4 weeks) over the 11 year study. During these surveys the tag number and
the location were recorded. An abalone was classified to have "moved" only if
it was captured in a zone other than it's release area. The location noted was the
midpoint of the zone for the dive. Distances moved were determined by cal-
culating a minimum least-linear distance between midpoints of the zones. The
null hypothesis that P|X = x | Y = yj is equal to P|X = x| was tested at the a =
0.05 level throughout using row by column and multiway tests of independence
following methods presented in Snedecor and Cochran (1980).
Nocturnal and Short-term Movements
From July 22, 1974, to September 9, 1974, monitorings of nocturnal and short-
term movements were conducted at three specific sites (i) South Channel, (ii)
Slot, and (iii) Outer Surge Channel with depths from 25-35', 40-50', and 55-65',
respectively ( Figure 2 ) . Early in the morning of the first day of study from 20 to
26 abalone ranging in size from 170 to 200 mm long were tagged in situ at each
site. The position that day of each specimen was mapped. Divers returned daily
to determine the total number of abalone at a particular location, the number
tagged, the numbers that had moved, and an estimate of the movement.
Model
For the apparent loss of tagged abalone at a given site, a model was developed
to quantify observed dispersion rates of red abalone at the three short-term
movement study sites. The fraction of tagged abalone present in a search of an
entire zone at time t was expressed as:
T, = T„e-ut +i
where, T^ = initial number of abalone tagged and released at a site.
T, = number of tagged abalone resighted at time t.
u = coefficient of loss (u = M 4- d).
MOVEMENT AND DISPERSION OF RED ABALONE 1 99
M = instantaneous rate of natural mortality,
d = instantaneous rate of dispersion,
t = time elapsed post tagging.
. ^ = error term associated with nonidentification and tag loss.
For the duration of the short-term study (45 days), it was assumed that losses
due to natural mortality and to tagging mortality were zero (i.e., M = 0) ; thus
the coefficient of loss reduced to:
u = M + d = d
where d is the modeled dispersion rate.
The observed rate of loss to the tagged population would then be strictly due
to dispersion. It was also assumed that there was no active predation on tagged
abalone, or that if predation did occur, then it affected both the tagged and untag-
ged populations at equivalent rates. The model was fitted to data utilizing
nonlinear least squares regression by the methods suggested in Draper and Smith
(1981) using an algorithm by Marquardt (1963).
RESULTS
Qualitative Assessments of Movements
Information attesting to movements involved some unmarked specimens. Ob-
viously, seeing animals moving is satisfactory evidence that movements occur.
We noted that different size classes usually occupied different habitats. Diurnally,
juveniles preferred the dark undersides of boulders and the recesses of crevices.
Specimens <50 mm long were found diurnally under boulders having a clean
veneer of crustose coralline algae on the boulder's undersides. No small abalone
were found under boulders bearing sediments or colonial invertebrates like
sponges and bryozoans. At Point Cabrillo, boulders occurred in waters <8m
deep. Specimens between about 50 mm and 100 mm long commonly occurred
diurnally in crevices and under large boulders. Specimens approaching or ex-
ceeding sport legal size ( > 178 mm long) were generally exposed but also uti-
lized large crevices and undersides of large boulders. Suitable habitat was found
between 5-20 m (16-65 ft) deep. Exposed specimens were generally attached
to scars. Scars are produced by abalone occupying a site for prolonged periods,
resulting in the death of the covering macrobiota. Scars, in varying stages of for-
mation, were noted many times, but scars were particularly prevalent during the
summer. Rocky surfaces predisposed to scar formation were those bearing col-
onies of the polychaete worm Dodecaceria concharum. Diurnally, exposed
specimens were rarely observed moving. In a few instances movement was as-
sociated with the twenty-rayed star Pycnopodia helianthoides making contact
with exposed abalone. We observed this sea star at times eating red abalone
within the study area. Large abalone were also observed traversing sand. This fact
may possibly indicate a means for repopulation of rocks lacking juvenile habitat,
but possessing habitat for adults.
Disturbance of boulders serving as abalone habitat caused the immediate
movement of juveniles and adults. When boulders are being rolled or disturbed,
abalone will drop from the boulder to prevent crushing. Circumstantial evidence
suggests that, in the winter, movement may be prompted by rock and boulder
habitat being displaced by heavy storm seas. Following periods of heavy seas we
200 CALIFORNIA FISH AND CAME
observed large boulders having been rolled, and some bore many unoccupied
scars. During these periods many abalone were found attached, but not to scars.
After periods of high seas, broken shells, of those abalone presumably crushed,
were found strewn along the bottom. Some abalone bore evidence of broken
shell repair.
Further evidence for movement is the fact that red abalone size classes were
stratified according to habitats. The degree to which adults were exposed varied
seasonally. More empty scars were noted during the winter, a period charac-
terized by heavy seas, an extreme paucity of food, and the lack of cryptic cover
provided by the low and attached algae. Occasionally abalone were found with
their shells covered by the biota typical of another depth or specific microhabitat.
Most frequently the shells of these mobile specimens were covered by the coral-
line alga, Calliarthron tuberculosum, characteristic of exposed and well lighted
areas found 8 m deep or less (Abbott and Hollenberg 1976). Some extant ab-
alone shells bearing C. tuberculosum were found as deep as 25 m; C.
tuberculosum does not survive at such depth. Specimens bearing the alga at vary-
ing stages of degeneration indicated that these specimens had been present at
these greater depths for periods ranging from days to probably weeks.
Nocturnal and Short Period Movements
During the afternoon of the first day of the short term movement study we re-
turned to each specific study site and observed that no specimens had moved.
However, upon returning the following morning we noted that a number of the
specimens had moved, and some were on the scars previously occupied by
other abalone. In some cases vacant scars were noted which had been occupied
the day before (Table 1). Evidently movement was nocturnal and specimens
tended to home back to scars, but not necessarily the one they previously oc-
cupied. Distances traversed by individual abalone ranged from 1.0 to 6.0 m per
day. Some tagged abalone moved at least eight times during the course of our 45-
day investigation, although some specimens apparently did not move to new lo-
cations at all. In a few cases tagged abalone apparently occupied the same scar
for weeks. In numerous cases abalone moved, and were either not observed
again, or were subsequently observed in the study area or the near vicinity at ir-
regular intervals. Movements were probably greater than we resolved by mea-
surement (Table 2). Over the period that the diurnal investigation was con-
ducted, a general pattern of diffusion of the tagged population was indicated at
all depths and locations studied ( Figure 3 ) . Modeled dispersion rates were great-
est in South channel (u.^ = 9.39 X 10~^) which was the shallowest and most
dynamic area studied, and least (Uq, = 2.61 X 10~^) in the Outer Surge Chan-
nel, the deepest and presumably most stable area with respect to physical dy-
namics. Factor (s) inducing movement were not completely ascertained. How-
ever, areas of strong current and greater sea exposure apparently create more
dynamic and fluctuating environments and may hence prompt a greater inci-
dence of movement in and out of these locations. Deeper water sites were gen-
erally more stable with respect to currents, but not necessarily to food supply.
Movements of red abalone both horizontally and vertically along the seabed
were common at all depths.
MOVEMENT AND DISPERSION OF RED ABALONE
201
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MOVEMENT AND DISPERSION OF RED ABALONE
203
South Channel
N-20
DAYS POST TAG (f)
FIGURE 3. Dispersion model curves fitted for the three short-term movement study sites at Point
Cabrillo reserve.
Long-term Movements
A total of 3,877 red abalone was tagged and released in the 1 1 zones of the
study area. A frequency distribution of the total numbers of abalones tagged and
released by 5 mm size classes is shown (Figure 4). Tagged specimens ranged
from 20 to 230 mm long, and except for abalone < 100 mm long, included a fair
representation of the population size structure. The mode of the sampled dis-
tribution occurred at 110 mm shell length. Difficulty in sighting abalone smaller
than 100 mm long resulted in less tagging of these animals. The distribution of
sizes tagged in most areas was broad ( Figure 5 ) , although actual sample sizes by
zone varied considerably. Approximately 4% of the total abalone tagged and re-
leased over the 4-yr tagging period were mortalities subsequently collected mor-
ibund ( Figure 6) . Some of these were collected as tagged shells only. Total mor-
talities recovered relative to the total numbers tagged showed two discrete
groups of recoveries; (i) those < 150 mm in shell length Xg = 7 11 , n.s.), and
(ii) those > 150 mm (X5 = 3.31, n.s.), suggesting differential mortality. Those
< 50 mm and > 210 mm were discounted because of the small sample sizes in
these size intervals. These patterns were interesting because the recovery of mor-
talities suggests that abalone >150 mm shell length had a mortality rate 1.42
times greater than abalone < 150 mm (17.9% vs. 7.4% respectively). Two pos-
sible reasons for the division in mortality rates were that: ( 1 ) the rate was actually
higher in abalone > 150 mm, or that (2) probably more likely, the shells of ab-
alone > 150 mm were more easily located by divers, and for specimens < 150
mm their shells could much more easily be overlooked or lost, crushed by rolling
boulders, eaten or otherwise destroyed. Additionally, the frequency of multiple
204
CALIFORNIA FISH AND GAME
resightings for abalone > 150 mm was higher and thus may have contributed to
the higher mortality observed in larger animals. In general, we believe that tag-
ging had a negligible effect on survival or behavior because most shell margins
of those animals recovered showed evidence of new growth. For this reason we
believe that mortality rates associated with tagging were low and probably equiv-
alent across all size classes.
N- 3,877
PERCENT OF
TOTAL TAGS ^
-1 ■ I — ' — I — ' — r — ' — I — ' — I — ' — I — ' — I — ■ — I — • — I — ■ — I — ■ — 1 — ' — I ■ I — • — I — =V
30 40 50 60 70 80 90 100 110 120 130 140 150 160 170 160 190 200 210 220 230
SHELL LENGTH (mm)
FIGURE 4. Frequency distribution by 5 mm size classes for total number of red abalone tagged and
released for all zones combined.
From the 3,877 abalone tagged and released in our study, 2,247 (58%) were
either resighted or recovered at least one time. A total of 4,302 individual resight-
ings of tagged abalone was recorded. Some specimens were resighted only once,
while others were resighted several times and thus confirmed differential resight-
ing rates among size classes (X^, = 74.03, p<.001). Abalone >110 mm long
had the highest frequency of resighting ( c.f. Figure 5 ) . The greatest fraction of re-
captures for all tagged size classes occurred during the first year after tagging;
subsequently, resighting rates declined exponentially for abalone placed in all
study zones (Table 3). Approximately 42% of the tagged abalone were never
resighted after their release. For those abalone resighted, their recovery sug-
gested statistical homogeneity of four contiguous groupings of size classes: (i)
31-70 mm (X^ = 0.53, n.s.); (ii) 71-110 (X; = 6.23, n.s.); (iii) 111-160 mm
(X4 = 7.42, n.s. );and (iv) 161-230mm (Xe = 7.47, n.s.); however, these group-
ings were significantly different from each other (Xn = 34.03, p<.001 ). The
probability of resighting an abalone at least one time after tagging was: for ab-
alone 161-230 mm, 67.4% of total tagged; for abalone 111-159 mm, 58.9%; for
abalone 71-110 mm, 51.6%; and for abalone 31-70 mm, 32.6%, respectively.
This may explain the disparity in recovery of shells of the various moribund size
classes. If the resighting rates of tagged dead abalone are roughly proportional to
those of live tagged abalone then these statistics bear out differential resighting
frequencies of the size classes. Assuming that natural mortality affected the
MOVEMENT AND DISPERSION OF RED ABALONE
205
□ Tagged a Released
North Cove
N = I248
Non-duplicated Resightings of
at least one time
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30 40 50 60 70 80 90 100 110 120 130 140 150 160 170 ISO 190 200 2I0 220 230
SHELL LENGTH (mm)
FIGURE 5. Size frequency distribution of tagged and released abalone (open area); and subse-
quent non-duplicated resightings of those specimens of at least one time by release
zone.
206
CALIFORNIA FISH AND CAME
8
7H
PERCENT OF g
TOTAL
MORTALITIES 5
OBSERVED
4
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50 60 70 80 90 100 110 120 130 140 150 160 170 180 190 200 210
SHELL LENGTH (mm)
FIGURE 6. Frequency distribution by 5 mm size classes of tagged red abalone recovered as mor-
talities for all zones combined.
resighted and non-resighted groups equally ( not a robust assumption ) , then non-
resighted group mortality was 110 abalone. This leaves some 1,520 tagged ab-
alone in an undetermined status after their release. The disappearance of these
tagged abalone may be attributed to the following factors:
1 . The fraction of tagged population loss due to natural mortality was higher than
that estimated by the recovery of shells with tags attached (i.e., some of these
empty shells, particularly those of the small size classes, went unrecovered).
2. Some tag loss could be attributed to weakening of the wire securing the tag
to an abalone through breaks or corrosion, or fracturing shells < 100 mm.
3. All tagged abalone in a particular zone were not recovered. The paucity of re-
coveries could have been influenced seasonally by heavy algal growth, and
intra-annually by the inherently cryptic nature of juveniles and subadults.
4. Tagged abalone emigrated from the study area.
Rate of Movements
About 11 percent of the tagged abalone that were resighted at least once
moved out of their release zone. Abalone in some zones exhibited a wider range
of movement than others (Table 4). Median distance moved was 87 m for
resighted abalone (Figure 7). Movements out of the respective release zones
varied temporally. The trend for movement was from shallow to deep water in
the summer, and from deep to shallow water in winter. Movements were ex-
tensive for all size classes. Distances of movement varied from 1m to over 150
m per month per individual (Table 5).
Evidently large distances were traversed by some red abalone. There were 29
records of abalone which had each travelled > 350m along the seabed off Point
Cabrillo over periods ranging from 3 to 61 months. Larger abalone apparently
tended to move more frequently and further (Figure 8). Two abalone, observed
3 months after tagging, had moved distances greater than 0.5 km least-linear dis-
MOVEMENT AND DISPERSION OF RED ABALONE
207
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CALIFORNIA FISH AND CAME
tances from the point of their release. One tagged abalone released in the Inner
Surge Channel was recovered alive approximately 9 yr later by a sportdiver near
Caspar State Beach, a distance 2.4 km north of the study site in least-linear
transect from the point of release. In addition, a shell from a tagged abalone re-
leased in the Inner Surge Channel was found 3 yr after that release near Caspar
State Beach. Other evidences for extensive movements by abalone were cor-
roborated by recorded observations that showed specimens released in one tag
zone were subsequently identified as having moved into another zone, then later
located, after another move, in the zone on the other side of their original release
zone.
130 -|
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MEDIAN DISTANCE
MOVED (-eTm)
100 150 200 250 300 350 400 450 500 550 600 650
MINIMUM DISTANCE MOVED (meters)
FICURE 7. Least-linear distance traversed by numbers of abalone observed to have moved for all
zones combined.
TABLE 5. Distribution of Tagged Abalone by Four Size Intervals, Number of Recoveries, and
Their Rates of Movement.
5/>e C/ass (mm)
<100
100—130
131 — 160
>160
Number Tagged
988
1,079
685
1,125
3,877
Number Moved
Out of
Designated
Tag Zone
75
73
73
203
424
% Moving
7.59%
6.77%
10.66%
18.04%
Range of Elapsed
Time Post- Tag
(Months)
3—61
3—86
4—86
3—86
Range of Movement
Per Month
(Meters /Month)
1.05- 59.6
0.89-153.8
0.80- 66.5
1.02-153.8
10.94%
3—86
0.89-153.8
MOVEMENT AND DISPERSION OF RED ABALONE
209
Numbtr
of
Ob««rvation«
Oittanc* Movad (m)
650
Oiatonc* Moved (m)
650
FIGURE 8. Three dimensional surface plot of numbers of observations as a function of abalone shell
length and the distance moved.
Tagged population abundance in the study area followed approximately an ex-
ponential decline. Our findings indicate that dispersal rates vary among size
classes (X ,^, = 74.03, p<.001 ). For those animals resighted at least one time,
two discrete (X j = 64.45, p<.001 ), but contiguous, groups of abalone that
were known to have moved were apparent: (i) specimens >150 mm
(X 5 = .24, n.s.), and (ii) specimens < 150 mm (X I = 10.91, n.s.). Abalone
>150 mm shell length had the highest rates of movement; 27.4% of those
resighted at least once had moved out of their release zone. This compared with
13.6% of those abalone < 150 mm. Generally, less than 20% of those abalone
released within a particular zone would still be located in the same zone two
years after tagging. Apparently the red abalone population at Point Cabrillo was
in a constant state of flux due to movement and dispersal, with the new members
entering the area as well as those leaving the area. However, some abalone re-
mained in the same general vicinity for relatively long periods. We probably have
underestimated the extent of movements as it is likely that some abalone left the
zone in which they were released, only to return to the same general vicinity in
which they were released before our next observation.
210 CALIFORNIA FISH AND GAME
DISCUSSION
The incidence of movement among red abalone at Point Cabrillo was high.
Large distances were traversed by individual abalone in relatively short periods
of time. These results contrast somewhat from those of Bonnot (1948), Cox
(1962), and Mines and Pearse (1982) on H. rufescens. This is perhaps because
of: (i) limited sample sizes; (ii) general problems of sampling marine systems
(Dayton andTegner 1984); (iii) time constraints used in sampling programs; and
(iv) possible effects of higher predation rates by sea otters not present in our
study site. We observed movements by abalones up and down the coastline, per-
haps in response to physiological and environmental stress. Further, these find-
ings augment field and laboratory observations of others that have suggested reg-
ular translocation and very active movement recorded for other haliotids
(Stephenson 1924, 1 no 1952, Newman 1966, Momma and Sato 1969, Poore 1972,
Shepherd 1973).
At Point Cabrillo predation is limited. Population pressure on red abalone
tends to be intraspecific; positioning for available current and food, minimization
of disturbance, and the facilitation of reproduction and recruitment are the pri-
mary concerns for vitality (Ault 1985b). Movement may involve avoiding the
hazards of climate and food shortage during unfavorable environmental events,
at the costs of the hazards of migration and ultimate survival in a new area.
The incidence of movement in the red abalone population varied from year to
year, and directly and indirectly depended upon sea conditions. Since red ab-
alone feed primarily on drift kelp, currents coupled with food supply, light, and
season in the nearshore area probably dictate the amount and quality of the algae
to which a particular abalone might have access. In the winter suitable food sup-
ply is found at relatively shallow depths. However, there is a tradeoff because tur-
bulence and wave action can disrupt the boulder habitat to which abalone ad-
here. Storm conditions increase the probability of abrasion, crushing and
detachment of abalone, and severe sea conditions apparently force movement
to deeper water or safer domain. Several authors have stated that haliotid move-
ments are prompted by physical disturbance (Graham 1941, Sinclair 1963, Poore
1972, Shepherd 1973). Poore (1972) stated that movement of juvenile H. iris in
New Zealand was seasonal, being greatest in the fall and winter when rough wa-
ter disturbed the habitat more frequently. Red abalone will leave sites on the
sandline when threatened by smothering sand, drift and debris. In the present
study the recovery of tagged abalone was higher below 8 m. Presumably rough
weather had less effect on abalone positioned in deeper waters.
In northern California a clear distinction between the microhabitats of juvenile
and adult red abalone was observed, and is similar to that reported for southern
California (Leighton, 1968) and central California (McLean, 1962). Due to size-
stratified differences in dietary requirements, and the cryptic nature of juvenile
abalone versus the more exposed positioning of adults, some migration between
juvenile to adult habitat must occur as abalone grow. Migration between habitat
types for juvenile and adult abalone has been reported by Newman (1966) for
H. midae, and by Shepherd (1973) for H. iris. In general, older red abalone oc-
cupy deep depressions on the surface of rocks indicating the dearth of move-
ment from that particular spot. These spots generally occupied by larger abalone
appear to be prime feeding locations. The most important single factor ensuring
MOVEMENT AND DISPERSION OF RED ABALONE 21 1
an adequate food supply is the abalone's preference for resting places on open
rock where drifting algae are carried or deposited. At Point Cabrillo, due to the
lack of predation by sea otters, being exposed on open rock face was allowable.
It is generally accepted that most abalone participate in nocturnal feeding ex-
cursions, moving out after dark to graze on surrounding algae (Graham 1941,
Bonnot 1 948, Sinclair 1 963, Leighton 1 968, Momma and Sato 1 969 & 1 970, Poore
1972, Shepherd 1973). However, there is considerable debate as to whether
these foraging abalone return to their "home scar". Some abalone apparently
spend their entire life on small isolated stones, boulders or rocks from which they
do not move ( Cox 1 962 ) . The persistence of the abalone's foot on a specific area
for prolonged periods of time contributes to scar formation. Scars can become
very deep, especially in soft mudstone or sandstone, as periodic twisting of the
shell by an abalone may cause abrasion of the substrate. Scar formation led to the
assumption of homing according to Sinclair (1963) and Tunbridge (1967). Hom-
ing to a fixed particular scar is well known in limpets (prosobranch relatives of
haliotids), and is influenced by the size of an animal, the texture and stability of
the homesite rock, and the availability of food (Branch, 1981 ). Bonnot (1948)
stated that H. rufescens forage during the night and will sometimes travel con-
siderable distances, returning to their "home spot" by day break. By contrast,
Leighton (1968) stated that his tagging observations indicated that homing is not
universal in adult H. rufescens and is virtually unestablished in young juveniles,
as no scar is present under these abalone. Other California haliotids, H. corrugata
and H. sorenseni, are believed to move as much as several meters at night, some-
times returning to previously occupied scars by dawn (Tutschulte 1968, 1976).
Both Forster (1962), with H. tuberculata, and Shepherd (1973), with H. ruber,
stated that abalone normally live in a retreat from which nocturnal feeding ex-
cursions are made, though these abalones may not invariably return to the same
retreat before morning. If homing by red abalone occurs it is probably a means
of regulating population density '■elative to food abundance. A shortage of food
may be the key factor increasing the observed incidence of non-homing abalone.
Thus, dispersion away from a neighbor, if movement is linked to food shortage,
can spread the population and reduce competition for food and space. Our work
demonstrates that red abalone do move, but they do not necessarily home to the
same scar which they had previously occupied. Furthermore, an animal might
spend years in the same general area, only to depart suddenly and be found else-
where at a later date. Homing is not an important factor in red abalone; however,
the acquisition of an unoccupied scar could be of primary importance in that it
provides a site for good attachment and food procurement.
Our observations also suggest that hunger may stimulate movement. The ex-
tent to which an abalone moves probably depends upon available food supply.
In the presence of sufficient food, abalone movements were meager. These ob-
servations are congruent with those of Hines and Pearse (1982) who stated that
abalone in their central California study site, in the presence of sea otters, ha-
bitually remained within their respective cracks, and that movement appeared to
be in positive response to the presence of drifting pieces of the giant kelp,
Macrocystis pyrifera. The fraction of the population that disperses outside the
boundaries of the original home territory is expected to be high when environ-
mental conditions locally eliminate a particular food supply and create new sup-
plies in other places (Cohen 1967, Vadas 1977).
212 CALIFORNIA FISH AND CAME
In areas of regular food supply little movement occurs. At Fort Bragg in the
winter, movement might be induced by the seasonal paucity of the kelps. If nec-
essary, adult red abalone will scrape benthic diatoms with their radula to survive
during periods of scarce food supply. MacGinitie and MacGinitie (1966) re-
ported, from laboratory observations, that starvation did not stimulate H.
corrugata to move. Indeed, this scenario may be the impetus for red abalone
movement. Translocation to areas which possibly afford less protection, and the
unknown probability of successful food procurement, could be detrimental to a
particular mobile abalone and therefore mal-adaptive.
SUMMARY
Movement and dispersal in a northern California red abalone population oc-
curred. However, some individual abalone exhibited no apparent movement
during our study. The general stimulus for movement may be due to limited food
supply, although physical disturbance may also prompt movement. A general
tendency exists for adult abalone to reposition on scars after movement, regard-
less of whether or not the scar was theirs previously. The apparent trend for
movement was from shallow to deep water in the spring-summer, and from deep
to shallow water in fall-winter in response to the highly seasonal and depth lim-
ited abundance of algae. Intraspecific competition may limit population density
by density dependent mortality, or by influencing the rate of emigration in re-
lation to food availability. The incidence of movements and resulting population
size flux is great. Densities of abalone remained fairly constant through time for
our particular location because much emigration and immigration occurred.
Small dense groups of subadult abalone may be capable of replenishing exploited
stocks of larger abalone in their vicinity by movement to these favorable but ex-
ploited reefs. Although strong inferences may be made here, further studies must
be conducted to determine the relationship between dispersion and natural mor-
tality before any absolute assessment of the fraction of population loss or gain
due purely to movements can be ascertained.
ACKNOWLEDGMENTS
This work is a result of research sponsored in part by NOAA, National Sea
Grant College Program, Department of Commerce, under Grant #04-5-158-28,
through the California Sea Grant College Program, and in part by the California
State Resources Agency. The U.S. Government is authorized to reproduce this
document and distribute for governmental purposes.
The authors also gratefully acknowledge the assistance of Steven Schultz and
Richard Burge, formerly with the Marine Resources Region of the California De-
partment of Fish and Game, and Donald Heacock while he was a graduate stu-
dent at Humboldt State University.
LITERATURE CITED
Abbott, I. A. and C). Hollenberg. 1976. Marine Alga of California. Stanford University Press. Stanford, California.
827p.
Ault, J.S. 1985a. Species profiles; life histories and environmental requirements of coastal fishes and invertebrates
(Pacific Southwest)— black, green and red abalones. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.32) U.S. Army
Corps of Engineers, TR EL-82-4. 19p.
Ault, J.S. 1985b. Some quantitative aspects of reproduction and growth of the red abalone, Haliotis rufescens
Swainson. ). World Maricult. Soc. 16: 398-^25.
MOVEMENT AND DISPERSION OF RED ABALONE 213
Bonnot, P. 1948. The abalones of California. Calif. Fish Game 34 (4) :1 40-1 69.
Branch, CM. 1981. The biology of limpets: physical factors, energy flow, and ecological interactions. Oceanogr.
Mar. Biol. Ann. Rev., 19:235-380.
Clavier, J. and R. Olivier. 1 984. Experimental study of the movements of the ormer ( Haliotis tuberculata) in nature.
Rev. Trav. Inst. Peches marit. 46(4) :31 5-325.
Cohen, D. 1967. Optimization of seasonal migratory behavior. Amer. Natur. 101:5-17.
Cox, K.W. 1960. Review of the abalone in California. Calif. Fish and Game 46(4):381^06.
Cox, K.W. 1962. California abalones, family Haliotidae. Calif. Fish and Game Bull. No. 118, 133p.
Dayton, P.K. and M.J. Tegner. 1984. The importance of scale in community ecology: a kep forest example with ter-
restrial analogs. In "A New Ecology: Novel Approaches to Interactive Systems", Chap. 17, P.W. Price, C.N.
Slobodchikoff and W.S. Gaud (eds.) Wiley, New York.
Draper, N.R. and H. Smith. 1981. Applied Regression Analysis. John Wiley and Sons, New York. 709p.
Forster, G.R. 1%2. Observations on the ormer population of Guernsey. J. mar. biol. Assoc. U.K. 42:493-498.
Graham, D.H. 1941. Breedinghabitsof twenty-two species of marine Mollusca. Trans. Royal. Soc. of New Zealand
71:152-159.
Hancock, D.A. 1979. Population dynamics and management of shellfish stocks. Rapp. P. -v. Reun. Cons. int. Expor.
Mer 175:8-19.
Hines, A.H. and J.S. Pearse. 1982. Abalones, shells, and sea otters: dynamics of prey populations in central Cali-
fornia. Ecology 63 ( 5) :1 547-1 560.
Ino, T. 1952. Biological studies on the propagation of Japanese abalone, genus Haliotis. Tokai-Ku Suisan Kenkyoju
Hokoku 5:1-102.
Leighton, D.L. 1968. A comparative study of food selection and nutrition in the abalone, Haliotis rufescens
(Swainson) and the sea urchin, Stronglyocentrotus purpuratus (Stimpson). Dissertation. Univ. of Calif., San
Diego. 197p.
Leighton, D.L., Byhower, M.J., Kelly, J.C, Hooker, G.N. and D.E. Morse. 1981. Acceleration of development and
growth in young green abalone, Haliotis fulgens, using warmed effluent seawater. J. World. Maricul. Soc.
12(1):170-180.
Marquardt, D.W. 1%3. An algorithm for least squares estimation of nonlinear parameters. J. Soc. Ind. Appl. Math.
11:431-^M1.
MacGinitie, N. and G.E. MacGinitie. 1966. Starved abalones. Veliger 8:313.
McLean, J. 1962. Sublittoral ecology of the kelp beds of the open coasts near Carmel, California. Biol. Bull.
122:95-114.
Momma, H. and R. Sato. 1969. The locomotion behavior of the disc abalone, Haliotis discus hannai Ino, and the
Siebold's abalone, Haliotis seiboldii Reeve, in the fishing grounds. Tohoku J. Agric. Res. 20(3):150-157.
Momma, H. and R. Sato. 1970. The locomotion behavior of the disc abalone, Haliotis discus hannai Ino, in a tank.
Tohoku J. Agric. Res. 21 (1 ):20-25.
Newman, G.C. 1 966. Movements of the South African abalone, Haliotis midae. Investl. Rep. Div. Sea Fish. S. Africa
56: 1-20.
Poore, G.C.B. 1972. Ecology of New Zealand abalones, Haliotis species (Mollusca: Gastropoda) 2. Seasonal and
diurnal movement. New Zealand J. Mar. Fresw. Res. 6(3):246-258.
Shepherd, S.A. 1973. Studies on southern Australian abalone (genus Haliotis): I. Ecology of five sympatric species.
Aust. J. Mar. Freshw. Res. 24(3):217-257.
Sinclair, M. 1%3. Studies on the paua, Haliotis iris Martyn, in the Wellington district, 1945-1946. Zool. Publ. from
Victoria Univ. of Wellington 35, 16p.
Snedecor, G.W , and W.G. Cochran. 1980. Statistical Methods. The Iowa State University press. Ames, Iowa. 507p.
Stephenson, T.A. 1924. Notes on Haliotis tuberculata. J. mar. biol. Assoc. U.K. 13(2):480-495.
Tunbridge, B.R. 1967. Feeding habits of paua. New Zealand Mar. Dept. Fish Tech. Rept. 20:1-18.
Tutschulte, T.C. 1968. Monitoring the nocturnal movements of abalones. Underwater Naturalist 5(3):12-15.
Tutschulte, T.C. 1976. The comparative ecology of three sympatric abalones. Dissertation. Univ. of Calif., San
Diego. 335 p.
Vadas, R.L. 1977. Preferential feeding: an optimization strategy in sea urchins. Ecological Monographs 47:337-371.
214 CALIFORNIA FISH AND GAME
Calif. Fish and Came 73 ( 4 ): 2 1 4-229 1 987
THE USE OF BAITED STATIONS BY DIVERS TO OBTAIN
FISH RELATIVE ABUNDANCE DATA ^
DANIEL W. COTSHALL
California Department of Fish and Game
Marine Resources Division
2201 Garden Road
Monterey, California 93940
Divers were used to count fishes at 317 baited stations at Cojo Anchorage near
Point Conception, California between September 1980 and June 1981. This method
was tested in order to develop a quantitative technique to assess impacts, on fish
populations, of a proposed liquefied natural gas terminal. The counts were con-
ducted in depths 30 to 70 feet (9.2-21.4m) on bedrock substrate. Forty-three iden-
tifiable species were attracted to these baited stations. To test the effectiveness of
the method to detect temporal and spacial changes in abundances, five species plus
total combined fishes were selected to compare differences in counts among seasons
within each study area and among four study areas for each season. The baited sta-
tion counts yielded significant differences in seasonal counts for rainbow surfperch,
Hypsurus caryi; kelp bass, Paralabrax clathratus; black surfperch, Embiotoca
jacksoni; and onespot fringehead, Neoclinus uninottus, at one or more of the four
study areas. There were also significant differences for these same species when
counts between study areas were compared.
INTRODUCTION
Biologists have been utilizing scuba for a number of years to obtain quantitative
data on shallow-water fishes and invertebrates. Various non-destructive methods
have been used in attempts to obtain this data. For example, Miller and Geibel
( 1 973 ) used permanent 30-m transects in Monterey Bay to obtain counts of kelp-
bed fishes in order to determine daily and seasonal fluctuations in abundance.
Ebeling, Larson, Alevision, and Bray (1980) used an underwater movie camera
to produce their "cinetransects" in kelp forests off Santa Barbara to obtain spe-
cies composition and annual variability in numbers of fishes between canopy and
bottom habitats. Another approach has been used by Jones and Thompson
(1978) who counted fish species rather than individuals of a particular species,
during specific time periods, while "swimming around" coral reefs off Florida.
Their method was designed to compare species abundance quantitatively
through time or between areas. All of these methods provide indices of abun-
dance that work better for some species than others.
All of the non-destructive observational methods utilizing divers are subject to
uncontrolled factors that would influence variation, including difficulty in the
diver's ability to objectively judge distances and sizes of fish along the transect,
and to accurately identify fishes, particularly those at the outer edge of visibility,
and the varying behavior of species either to be attracted to or repelled by divers.
In addition cryptic and crevice dwelling species are usually missed by divers. The
use of all of the techniques in central and northern California has been further
questioned due to the role that surge and turbidity play in hampering the diver's
ability to concentrate on identifying and counting fishes. Also, heavy surge can
' Accepted for publication January 1987.
BAITED FISH STATIONS 2 1 5
cause many fishes that live near or on the bottom to seek shelter in caves and
crevices v^here the divers may not see them. Finally, there is the problem of ac-
curacy of diver counts of large schools that may contain 50 or more fish. Because
of all of these factors none of the diver survey methods can yield accurate spe-
cies composition data.
An alternative method, which involves counts of fishes attracted to baited sta-
tions, required testing to determine v^hether at least some of the variability in-
herent in the three methods discussed above could be reduced. The objective
of this study was therefore to develop such a method to attempt to produce rel-
ative abundance (catch-per-unit-of-effort, CPUE) data which would be useful in
assessing impacts on fish populations. In this case, the sport and commercial spe-
cies in the vicinity of a proposed liquefied natural gas (LNG) terminal. Two null
hypotheses were tested: (i) there were no significant seasonal changes in abun-
dance of the dominant fishes that live on or near the bottom; and ( ii ) there were
no significant differences in abundance of the dominant fishes between the pro-
posed LNG terminal area and two control areas.
DESCRIPTION OF STUDY AREA
The present study was conducted at Cojo Anchorage, just south of Pt. Con-
ception, California ( Figure 1 ) . The study area at Cojo Anchorage consists of large
areas of relatively flat bedrock interspersed with sand patches and channels. Six
study areas were established: a shallow (CW30) and deep (CW60) control area
west of the proposed LNG terminal site, a shallow (T30) and deep (T60) area
at the terminal site, and a shallow (CE30) and deep (CE60) control area east of
the proposed LNG terminal site (Figure 1 ). The west end of the study area con-
tains large sandy areas, particularly in depths greater than 14 m. Most of the sub-
strate in the proposed LNG terminal area is low-relief ( 1 m) of flat bedrock with
one major sand channel. The eastern portion of the study area consists of a mix-
ture of sand and bedrock in waters shallower than 14 m and low-relief bedrock
in deeper water (14-18 m). Much of the bedrock substrate supports beds of gi-
ant kelp, Macrocystis pyrifera, and the brown alga, Pterygophora californica.
METHODS
Sampling was conducted during four quarters beginning in September 1980
and concluding in June 1981. Counts were originally made at the six locations
mentioned above. The deep west control area (CW60) and shallow east control
area (CE30) were deleted from the sampling plan during the spring and summer
1981 surveys, because of lack of bedrock substrate. Seven random locations
within each study area were selected to be sampled each quarter. During the
spring and summer 1981 surveys, random stations were increased from 7 to 11.
Counts at the baited stations were conducted between one hour after sunrise and
one hour before sunset; however most counts were done between 0800 and
1500 h.
At each random station two divers descended to the bottom with a canvas bag
containing two lengths of 2-m chains connected in the middle. A bait container
filled with roe and guts from four to six sea urchins (Strongylocentrotus) was at-
tached to the center of the chains after the chains were laid out in the form of
a cross. The arms of the cross formed the radii of a circle two meters in diameter.
In effect the baited station acted as a trap with virtually unlimited access for
216
CALIFORNIA FISH AND GAME
SCU£:
0 1000
iiobothi in feci
FIGURE 1. Locations of study areas for visual fish observations at baited stations: CW60 = deep
west control, CW30 = shallow west control, T60 = deep trestle site, T30 = shallow
trestle site, CE60 — deep east control, CE30 = shallow east control.
fishes. At the signal of the diver team leader, each diver began recording the
nunnbers of each species of fish that entered the circle and within one meter of
the bottom. The counts were recorded on a minute-by-minute basis for ten min-
utes. The ten minute observation period was selected by using pre-survey data
to plot numbers of fishes (all species) observed each minute for ten minutes at
all stations. The resulting curve peaked at five minutes (Figure 2). Based on this
curve we selected the ten minute observation period as a compromise between
increasing the number of fishes that might occur with a longer observational pe-
riod and the number of stations the divers could complete on a single tank of air.
The divers also kept track of and recorded the total number of each species that
entered the circle during the ten minute count. At the completion of the first
count, the divers laid out a 30-m transect line due north of the first station. At the
terminus of the 30-m transect, a second ten minute count was conducted. At the
shallow study areas a third ten minute count was conducted 30-m west of the
second station. At each station the divers also recorded depth, substrate type,
presence of Macrocystis and Pterygophora, and horizontal visibilities as mea-
sured on the transect tape. Immediately upon completion of the dives, the divers
BAITED FISH STATIONS
217
compared counts to resolve any differences. These composite counts were re-
corded on a separate data sheet. Water temperatures were taken at the surface
with Martek VI water quality analyzer.
3
4
Q
UJ
_
_
_
_
t—
Z
ZD
—
-
_
_
O
u
3
—
X
in
"
u.
■
u.
o
a:
2
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OD
r-
-
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z
a
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-
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n
1 1 1 1 1 1 1 1 1
4 5 6 7 8
TIME INTERVAL IN MINUTES
10
11
FIGURE 2.
Mean number ( ± one standard error) of fishes, all species combined, observed
each) minute at 21 pre-survey baited stations, Cojo Anchorage, June-July 1980.
Target Species
To test the effectiveness of the baited stations to reflect any changes in abun-
dance, five species were selected from among the ten most frequently observed
species (Table 1): rainbow surfperch, Hypsurus caryi; kelp bass, Paralabrax
clathratus; black surfperch, Embiotoca jacksoni; onespot fringehead, Neoclinus
uninotatus; and smooth ronquil, Rathbunella hypoplecta. I also selected for test-
ing the total fishes (all species combined) observed at the stations. The results
of the fish counts at Station CW60 and at Station CE30 are not included in this
report because the final sampling plan was based on observations only on bed-
rock substrate.
Quarterly distributions of the count-per-station for each of five species, and to-
tal fishes, for each study area were tested for normality using the Komogorov-
Smirnov goodness of fit test ( Sokal and Rholf 1 969 ) . The Kruskal-Wallis test ( K-
W) (Sokal and Rohlf 1969) was used, at a significance level of p<0.05, to
determine if differences in average counts were significant among study areas
and quarters. Dunn's Multiple Comparisons (Dunn 1964) were used to locate
the significant differences. An experimental error rate of p < 0.10 was selected for
these tests.
Pearson's correlation (Sokal and Rohlf 1969) was used to test the null hy-
pothesis that there was no correlation between species counts and bottom tem-
peratures recorded at each observation site.
Kendall's correlation coefficient (Sokal and Rohlf 1969) was used to test the
null hypothesis that there was no correlation between the counts of two species
at an observation site.
218 CALIFORNIA FISH AND CAME
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BAITED FISH STATIONS
219
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220 CALIFORNIA FISH AND CAME
Means and standard errors are used in the graphs (Figure 3-8) to show changes
in abundance.
The Mann- Whitney test (Sokal and Rohlf 1969) was used to test the null hy-
pothesis that there was no difference between the shallow stations (T30 and
CW30) with the deep stations (T60 and CE60) for each species to see if depth
was a factor in the differences in abundance between stations.
RESULTS
A total of 317 visual fish counts was connpleted at the four bedrock study areas
(T60, T30, CE60, and CE30). Forty-seven species were identified, of which four
could only be taken to genus (Table 1 ).
Fishes that occurred on or near the bedrock included swell shark,
Cephaloscyllium ventriosum; smooth ronquil, Rathbunella hypoplecta; flatfishes,
Bothidae and Pleuronectidae; fringeheads, Neoclinus spp.; sculpins, Cottidae;
surfperches, Embiotocidae; greenlings, Hexagrammidae; wrasses, Labridae; and
kelp bass, Paralabrax clathratus. The sandy areas were frequented by California
lizardfish, Synodus lucioceps; midshipmen, Porichthys spp.; Pacific angel shark,
Squatina californica; and flatfishes. The highest number of species was observed
at study area CE60 (Table 1 ). The most frequently observed species at each of
the four study areas were: the smooth ronquil at T60, kelp bass at T30, sanddabs
at CW30, and rainbow surfperch at CE60. Only a few rockfishes were observed
at any of the study areas.
The results of the Kolmogorov-Smirnov tests of the counts were significantly
different from normal and non-parametric tests were then used to analyze the
data.
Comparison of Counts Among Seasons
The Kruskal-Wallis test for the four sampling periods (seasons) showed sig-
nificant differences of counts for the following species: rainbow surfperch (at
study areas T30, CE60, and CW30), kelp bass (T30, CE60, and CW30), black
surfperch (T30, CE60, andCW30) (Table 2). Generally these three species were
most abundant during summer or fall sampling periods (Figures 3, 4, and 5).
Onespot fringehead showed a different trend, these cryptic fish were more abun-
dant during the winter sampling period ( Figure 6); differences in counts were sig-
nificant only at T30 (Table 2) . Counts of smooth ronquil did not produce any sig-
nificant differences between sampling periods (Table 2, Figure 7). When the
comparisons of total fish counts at each study area were made the seasonal dif-
ferences between these total fish counts were significant at all of the study areas
(Table 2, Figure 8).
Dunn's Multiple Comparisons indicated where significant seasonal differences
occurred between sampling periods. These differences occurred at all four study
areas between combinations of summer/winter or spring and fall/winter or
spring (Table 3).
BAITED FISH STATIONS
221
»~ 10
=)
o
o a
(14) (17) (33) (33)
+
T30
2
(2) (11) (33) (33)
+
CW30
(13) (13) (22) (22)
+
T60
(13) (14) (22) (22)
CE60
F W Sp S F W Sp S
FIGURE 3. Mean counts per station { one stanciard error) for rainbow surfperch observecJ by
divers at baitecd stations, Cojo Anchorage, October 1980 to June 1981, (N) = number
of stations.
4.
8
K
O 6
O
4-
T30
1
+ •
(14) (17) (33) (33)
4-
(1
T60
3) (13) (22) (22)
CW30
(2) (11) (33) (33) •
+ .
1 -(-
(1
CE60
3) (14) (22) (22)
F W Sp S F W Sp S
QUARTER
FIGURE 4. Mean counts per station ( ± one standard error) of kelp bass observed by divers at
baited stations, Cojo Anchorage, October 1980 to June 1981, (N) = number of stations.
222
CALIFORNIA FISH AND GAME
K 6
8 '
(14) (17) (33) (33)
T30
(2) (11) (33) (33)
+
■f
CW30
(13) (13) (22) (22)
+ _ +
T60
4
(13) (14) (22) (22)
+
CE60
F W Sp S F W Sp S
QUARTER
FIGURE 5. Mean counts per station ( ± one standard error) of black surfperch observed by
divers at baited stations, Cojo Anchorage, October 1980 to June 1981, (N) = number
of stations.
o
O 20
I
+
(14) (17) (33) (33)
CW30
12) (11) (33) (33)
(13) (13) (22) (22)
CEeo
(13) (14) (22) (22)
F W Sp s F W Sp S
QUARTER
FIGURE 6. Mean counts per stations ( ± one standard error) of onespot fringehead observed by
diversatbaitedstations, Cojo Anchorage, October 1980 to June 1981, (N) = number
of stations.
BAITED FISH STATIONS
223
1.2
.a
I 0
o
O 12
.8
.4
T30
+
+
(14) (17) (33) (33)
CW30
+ +
(2) (11) (33) (33)
T60
(13) (13) (22) (22)
CE60
(13) (14) (22) (22)
F W Sp S F W Sp S
QUARTER
FIGURE 7. Mean counts per station (± one stanciar(d error) of smooth ronquil observecJ by
(divers at baited stations, Cojo Anchorage, October 1980 to June 1981, (N) = number
of stations.
ie
TJO
T60
-
-
12
--
a
-.
(1<) (17) (33) (33)
4
+
(13) (13) (J2) (22)
-H
-1-
0
36
CW30
CE60
32
o
o
24
20
16
(13) (14) (22) (22)
12
+
-
a
1
■
4
0
4-
(2) 111) (33) (33)
-(-
F W Sp S F W Sp S
QUARTER
FIGURE 8. Mean counts per station ( ± one stan(dar(J error) of total fishes observecJ by divers
at baited stations, Cojo Anchorage, October 1980 to June 1981, (N) = number
stations.
224
CALIFORNIA FISH AND GAME
TABLE 2. Significance Levels of Kruskal-Wallis Tests of Comparisons of Quarterly Counts Per
Station of Fishes at Each Study Area, Cojo Anchorage, October 1980 to June 1981.
Species
Rainbow surfperch
Kelp bass
Black surfperch
Onespot fringehead
Smooth ronquil
Total fishes
* Significant at p < 0.05
T60
T30
CE60
CW30
0.053
< 0.001 •
< 0.001 ♦
< 0.001
0.340
< 0.001 •
< 0.001 *
< 0.001
0.135
< 0.001 •
< 0.001 •
< 0.001
0.294
0.001 *
0.074
0.152
0.574
0.589
0.051
0.226
< 0.001 •
< 0.001 *
< 0.0001 •
< 0.001
TABLE 3. Dunn's Multiple Comparison Values for Tests of Diver Fish Counts Between Sam-
pling Periods, (for six combinations), Cojo Anchorage, October 1980 to June 1981.
Rainbow
Surfperch
Kelp
Bass
+ '
+ '
+ '
+
+
+
+
+
+
+
Sampling Period
Combinations
T30
Summer/Fall
Summer/Winter
Summer/Spring
Fall/Winter
Fall/Spring
Winter/Spring
T60
Summer/Fall
Summer/Winter
Summer/Spring
Fall/Winter
Fall/Spring
Winter/Spring
CW30
Summer/Fall
Summer/Winter
Summer/Spring
Fall/Winter
Fall/Spring
Winter/Spring
CE60
Summer/Fall
Summer/Winter
Summer/Spring
Fall/Winter
Fall/Spring
Winter/Spring
• Positive signs indicate significant differences (p < 0.10)
Black
Surfperch
+ •
+
+
+
+
+
+
+ •
Onespot
Fringehead
+ •
+ •
Smooth
Ronquil
Total
Fishes
+
+
+
+
+
+
+ *
+ •
+ •
+ •
+
+
TABLE 4. Significance Levels of Kruskal-Wailis Tests of Comparisons Between Study Areas,
of Counts Per Station of Fishes, Cojo Anchorage, October 1980 to June 1981.
Species
Rainbow surfperch
Kelp bass
Black surfperch
Onespot fringehead
Smooth ronquil
Total fishes
* Significant at p < 0.05
QUARTER
Fall
0.004
0.001
0.001
0.130
0.010
0.026
Winter
0.209
0.779
0.430
0.607
0.058
0.157
Spring
no fish
0.176
0.535
0.635
0.062
0.307
Summer
0.001 *
0.001 *
0.001 *
0.002*
0.033 *
0.001 •
All Quarters
Combined
0.001 *
0.001 *
0.001 *
0.026 *
0.001 *
0.002*
BAITED FISH STATIONS 225
Comparison of Counts Between Study Areas
Significant differences in counts occurred between study areas for rainbow
surfperch, kelp bass, black surfperch, smooth ronquil, and total fishes during the
fall (Table 4). There were no significant differences between study areas for any
of the tested species during the winter and spring. Conversely, all of the tested
species showed significant differences in counts for the summer sampling period
and for all sampling periods combined. Most of these differences for the all sea-
sons combined data were between T30 and CE 60, T60 and CE 60, and CW30
andCE60 (Table 5).
TABLE 5. Dunn's Multiple Comparison Values for Tests of Diver Fish Counts Between Study
Areas (for six combinations). All Sampling Periods Combined, Cojo Anchorage,
October 1980 to June 1981.
Study Rainbow Kelp Black Onespot ** Smooth Total
Areas Surfperch Bass Surfperch Fringehead Ronquil Fish
T30/T60 __-_--
T30/CW30 - - - - + * -
T30/CE60 + * - + * - - + *
T60/CW30 - - + * - + *
T60/CE60 + * + * + * - - + *
CW30/CE60 + * + • - - - + *
* Positive values indicate significant differences (p < 0.10).
** Pairwise differences were not significant because of the experimental error rate although
overall comparisons was significant (TABLE 4).
DISCUSSION
A baited station can be visualized as a trap with a diameter of 2 m and d height
of 1 m. This trap allows unlimited access by fishes to the bait and allows the diver
to record those species that might not have been able to find the entrance to a
conventional trap or those that were able to escape.
The successful use of CPUE (relative abundance) data for calculating popu-
lation size assumes that the catchability does not change due to seasonal changes
in abundance or the fishes' behavior, and there is no difference in individual vul-
nerability (Ricker 1975). Recruitment, natural mortality, immigration and emi-
gration can also introduce error into population estimates that use CPUE data.
The purpose of this study was to determine if CPUE data from baited stations
would show seasonal changes in abundances and differences in abundance due
to depth and habitat type.
CPUE from trap data have been used to determine changes in abundances of
a number of marine species (e.g., Dungeness crab. Cancer magister, Gotshall
1 978) . Baited traps have also been used extensively on land to sample insects for
population estimates (Southwood 1966).
It was assumed that individuals of each species of fish that would be attracted
to the baited stations would be attracted at some constant rate that reflected their
abundance in the study area. It is implied in the use of baited stations that not all
species of fishes in the study area would be attracted to the bait.
226 CALIFORNIA FISH AND GAME
Differences Among Seasons
All but one of the species (smooth ronquil) tested showed significant differ-
ences in mean counts among the four seasons. In all cases, except for the onespot
fringehead, the lowest counts occurred during the spring quarter. It is assumed
that there was no difference in the attractiveness of the bait during the four sea-
sons and that the numbers of fishes that visited the observation site represented
their true abundance in the study area. Based on our counts at the baited stations
and observations during dives in the study area it was concluded that some of the
fish tested left the study area during late winter and spring. Unfortunately, I have
only one year of observations, so I cannot say whether this apparent decline in
abundance is an annual event. However, similar reductions in abundance of kelp
bass, rainbow surfperch, and other kelp bed fishes during winter and spring
months were observed by Miller and Geibel (1973). Laur and Ebeling (1983),
consider rainbow surfperch as "transients" at their study area at Naples Reef off
Santa Barbara; these surfperch arrive in late spring and depart in the fall. The
movement out of the study area may be due to several factors. Kelp bass tagging
studies showed that at least some of these fish moved away from the tagging site;
3% of 410 recovered tagged kelp bass moved five or more miles and 5% moved
up to four miles from the original tagging site (Collyer and Young 1953). Kelp
bass spawn from late spring into late summer (Frey 1971 ) and their movement
may be associated with spawning activity. The shallow waters of the Cojo An-
chorage area became very turbid during periods of winter and spring storms; this
turbidity may have affected either the food supply or the ability of some fishes
to obtain food and forced them to move offshore.
Both black surfperch and rainbow surfperch are viviparous; the young are born
during late summer and early fall (Behrens 1977). Dave Behrens (PG&E, Avila
Beach, pers. comm.) speculates that mating probably occurs during late fall.
Behrens (1977) also noted that rainbow surfperch moved out of his study area
( Half Moon Bay, central California) in October and November. Thus, the move-
ment of these two species could be related to mating and/or pregnancy and
birth. Onespot fringehead showed a different pattern, their greatest abundance
occurred during the winter quarter (except at CE30) . John Stephens (Occidental
College, pers. comm.) has observed similar movements of this species in the
Redondo Beach area. Studies in Monterey Bay (Lindquist 1981) indicated a
spawning season for onespot fringehead from January to September. Thus,
onespot fringehead at Cojo Anchorage may move into deeper water during the
spawning season (Figure 6).
Changes in temperature may act as a signal to fishes indicating arrival of
spawning season or poor feeding conditions in inshore waters. The Pearson's
correlation test yielded significant p values ( <0.05) for rainbow surfperch, kelp
bass, black surfperch, and total fishes. Temperature accounted for 23% of the
variation in counts for rainbow surfperch, 30% for kelp bass, 24% for black
surfperch and 36% for total fishes. From these results, I conclude that temper-
ature probably is not a major factor influencing movements of those fishes that
were tested.
BAITED FISH STATIONS 227
It was also thought that predatory fishes might inhibit smaller species from en-
tering the baited stations. Significant Kendall's correlation coefficients (p <0.05)
were obtained between rainbow surfperch and kelp bass, black surfperch and
kelp bass, and rainbow surfperch and onespot fringehead, however the r values
were all less than 0.30. Thus, there is little evidence that the presence of kelp bass
inhibits that of adults of small species. It is more likely that microhabitat selection
of kelp bass and black and rainbow surfperches is similar. A positive correlation
may also reflect the greater mobility of these relatively large species. Negative
correlation between rainbow surfperch and onespot fringehead probably reflects
the latter species microphabitat preference of flat bedrock containing pholad
clam holes.
The smooth ronquil showed some differences in relative abundance at all the
stations but none were significant. Based on these data, I believe that this spec-
ifies is a permanent resident of the area.
Differences Among Stations
The significant differences in abundance of species between the four stations
may reflect the difference in microhabitat of each of the stations. For example,
kelp bass were most abundant at T30 and CE60, (both of these areas contained
medium profile reefs, 0.5-1.0 m), while T60 and CW30 were almost devoid of
any type of reef structure. Both rainbow surfperch and black surfperch were most
abundant at CE60 and T30 and the presence of medium profile reefs may also
have been responsible for their abundance at these two study areas. Onespot
fringehead were most abundant at CE30 (apparently the bedrock substrate here
provided more pholad holes for them to live in). The fact that smooth ronquils
were most abundant at T60 probably is due to their preferring deeper water. The
presence of medium profile reefs at CE60 may account for the larger number of
fishes being counted at this study area.
The comparison between shallow stations and deep stations for each species
was significant for onespot fringehead, smooth ronquil, and total fishes. This sig-
nificant difference in mean counts between depths for onespot fringehead and
smooth ronquil may be a reflection of microhabitat as well as depth differences.
The significant difference obtained for total fishes probably is due to microhabitat
selectivity as well as a depth range preference by some species.
There is little doubt that some species are attracted to divers or bait. In previous
studies at Cojo Anchorage (R. Dixon, Calif. Dept. Fish and Game, unpublished
data), we tried using a method where two divers descended to a certain spot and
counted all fishes that they could observe around them for a period of five min-
utes. This method was dropped because not enough fishes were attracted to the
divers.
CONCLUSIONS
The use of baited stations to obtain CPUE data on those fishes attracted to bait
has been shown to have both advantages and disadvantages. Based on our initial
tests of baited stations, before this present study was begun, we knew that there
would be difficulty in counting large numbers of fishes (e.g., seiiorita, Oxyjulis
californica) during each one-minute time period. Thus making it very difficult in
228 CALIFORNIA FISH AND GAME
keeping track of individual fish as they entered and left the observation site during
the entire ten-minute count. We also knev^ that the method could not be used
to calculate population numbers for a particular species unless the fishes at-
tracted to the station were killed or removed from the population. In order to
make such an estimate the exact area influenced by the bait for each species
would also have to be determined.
Factors that might cause variations in baited station counts include:
(i) the time of day the stations are occupied;
( ii ) the length of time between when the bait is set and the divers begin count-
ing;
(iii) strength and direction of currents;
(iv) differences in the gear divers use, i.e., colors, regulator type; etc., and
(v) increased response to baited station, particularly if the fish were able to
obtain some of the bait from the baited canister.
The initial tests suggested several advantages, including: (i) the method allows
divers to complete more stations in a small area during a working day than swim-
ming transects (a pair of divers can complete two to three stations in the same
area covered by a transect per dive, thus increasing the number of samples to be
used for statistical analysis), (ii) the baited station functions as a fish trap and has
the advantage over an actual trap in that all fishes attracted within the boundaries
of the station are "captured" by observation of the diver ( standard fish traps only
capture those fishes that enter the trap and do not escape), (iii) this method re-
duces the bias of how much time divers swimming a transect spend looking for
midwater, demersal, or cryptic fishes, (iv) divers are able to concentrate on
counting fish rather than being distracted by the efforts of swimming along a
transect, particularly during periods of surge or low visibility, and (v) the baited
stations attract some cryptic species that might be missed by divers swimming a
transect line.
I believe that the results of this study show that baited stations offer a stan-
dardized, controlled, and repeatable method of obtaining CPUE data for fishes
attracted to baited stations. The method provides good quantification for the spe-
cies tested and is operationally simple. Individuals of each species probably re-
spond to the bait at different rates. If this response rate does not fluctuate sig-
nificantly during the year, then the CPUE data reflect the relative abundance for
that species. In order to obtain relative abundance data on fishes not attracted to
baited stations, one would have to use some other method.
The data from this study showed that there were significant differences in sea-
sonal abundances for kelp bass, rainbow surfperch, and onespot fringehead.
Other studies in central and southern California have shown similar changes in
seasonal abundance. The comparison of counts among study areas indicated sig-
nificant differences that reflect the well accepted concept that high profile reefs
support a more diverse and abundant fish fauna. Future baited station studies
could consider using a mean of the ten one-minute counts of each species or the
maximum one minute count for statistical calculations. Gary Davis (National
Park Service, pers. comm). recently has used maximum one minute counts for
statistical analysis of baited station data collected from the Channel Islands Na-
tional Park. Separate studies could be conducted to determine which bait or
combination of baits will work best in a particular area and for particular species.
BAITED FISH STATIONS 229
It would also be useful to test the area influenced by a particular bait for each spe-
cies and the best time of day to conduct sampling.
The data indicate that the shallow waters of Cojo Anchorage support a wide
diversity of fishes, but in most cases the species observed in the study area were
not very abundant. The abundances of several species changed seasonally. Hab-
itats in the four study areas were probably responsible for the significant differ-
ences in CPUE between the study areas for four of the species tested.
ACKNOWLEDGMENTS
This project could not have been conducted were it not for the contributions
of the following divers: P. Reilly, K. Henderson, K. Miller, L. Ley, K. Matthews,
K. Shannon, and G. Stone who conducted the fish counts. L. L. Hahn and S.
Dostaiek typed the original manuscript. B. Hammer helped design computer
forms and was responsible for entering all field data into the computer. J. Geibel
and A. MacCall advised on data analysis and design of the study. P. Law con-
ducted the statistical analysis. R. N. Lea and anonymous reviewers provided
many useful suggestions for improving the manuscript.
LITERATURE CITED
Behrens, D. W. 1977. Fecundity and reproduction of the viviparous perches Hypsurus caryi (Agassiz) and
Embiotoca jacksoni (Agassiz). Calif. Fish and Came 63(4): 234-252.
Collyer, R. D. and P. H. Young. 1953. Progress report on a study of the kelp bass, Paralabrax clathratus. Calif. Fish
and Game 39(2): 191-208.
Dunn, O. J. 1964. Multiple comparisons using rank sums. Technometrics (6): 241-254.
Ebeling, A. W., R. J. Larson, W. 5. Alevision, and R. N. Bray. 1980. Annual variability of reef-fish assemblages in kelp
forests off Santa Barbara, California. Fishery Bull. 78(2): 361-377.
Frey, H. W. 1971.California'slivingmarineresourcesand their utilization. Calif. Dept. Fish and Game, Sacramento,
148 p.
Gotshall, D. W. 1 978. Catch-per-unit-of-effort studies of northern California Dungeness crabs. Cancer magister. Ca-
lif. Fish and Game. 64(3): 189-199.
Jones, R. 5. and M. J. Thompson. 1978. Comparison of Florida reef fish assemblages using a rapid visual technique.
Bull. Mar. Sci. 28(1): 159-172.
Laur, D. R. and A. W. Ebeling. 1983. Predaior-prey relationships in surfperches. Env. Biol. Fish. 8(3/4): 217-229.
Lindquist, D. G. 1981. Reproduction of the onespot fringehead, Neoclinus uninotatus in Monterey Harbor, Cali-
fornia. Bull. Calif. Acad. Sci. 80(1): 12-22.
Miller, D. J. and J. J. Geibel. 1973. Summary of blue rockfish and lingcod life histories; a reef ecology study: and
giant kelp Macrocystis pyrifera, experiments in Monterey Bay, California. Calif. Dept. of Fish and Game, Fish.
Bull. 158: 137 p.
Ricker, W. E. 1975. Computation and interpretation of biological statistics of fish populations. Dept. of the Envi-
ronment Fisheries and Marine Service. Bull. 191: 382 p.
Sokal, R. R. and F. J. Rohlf. 1969. Biometry. The principles and practices of statistics in biological research. W. H.
Freeman and Co. San Francisco, CA., 776 p.
Southwood, T.R.E. 1966. Ecological Methods with Particular Reference to the Study of Insect Populations. Methuen
and Co. Ltd. London. 391 p.
230 CALIFORNIA FISH AND GAME
Calif. Fish and Came 73 (4) : 230-237 1987
SURVIVAL AND RECOVERY RATE ESTIMATES OF
NORTHERN PINTAIL BANDED IN CALIFORNIA, 1948-79^
WARREN C. RIENECKER
California Department of Fish and Came
1416 Ninth Street
Sacramento, CA 95814
Estimates of survival and recovery of northern pintail. Anas acuta, banded
preseason (N=168,763) and postseason (N=59,165) at seven stations in California
were analyzed. Postseason banded pintails followed the same pattern of survival rate
estimates as those from preseason bandings. Adults had higher survival and lower re-
covery rate estimates than did immatures. Adult males had higher survival and re-
covery rate estimates than did adult females. Pintails banded in Imperial Valley have
lower survival rates than those banded in northern California.
INTRODUCTION
Most survival studies in the past two decades have been done on the mallard,
Anas platyrhynchos, whereas only one (Anderson and Sterling 1974) has been
compiled on the northern pintail. Based on wintering population and total har-
vest, northern pintail are the most important species of waterfowl in California
( Pacific Flyway midwinter waterfowl surveys, USFWS waterfowl parts collection
surveys ) . An average of 2 million migrate there annually from northern breeding
grounds, accounting for approximately half of the total ducks wintering in Cal-
ifornia (Bellrose 1976). About 56% of the U.S. harvest of pintails occurs in the
Pacific Flyway of which over half takes place in California (USFWS waterfowl
parts collection surveys). Females are harvested closer to the breeding grounds
and show a greater homing instinct to the area of banding than do males
(Reinecker 1987). Males tend to range wider and are more likely to be recovered
in Mexico, Central America or on one of the other flyways than are females.
The purpose of this report is to examine and compare survival and recovery
rate estimates for pintail banded at seven stations in California (Figure 1 ).
METHODS
A total of 245, 174 northern pintail was banded in California from 1948 to 1979.
Of these, 227,928 were used to determine survival and band recovery rate es-
timates for the major banding stations (Table 1 ). In the 1950's and 1960's pintail
were banded on many waterfowl concentration areas in California. Some of
these bandings were exploratory and lasted only a year or two. On the more im-
portant areas, banding was nearly continuous through the 1950's. Thereafter,
only Klamath Basin NWRs and Gray Lodge Wildlife Area were used as banding
stations. All pintails to be banded were caught in baited, wire, swim-in traps or
on baited cannon net sites. Traps were checked daily, caught pintail were banded
with standard FWS aluminum leg bands and released. Only preseason ( 1 july-30
September, N = 168,763) and postseason (16 January-15 March, N = 59,1 65)
banded pintails recovered as direct or indirect recoveries through 1979 were
used. Direct recoveries are banded birds recovered during the first hunting
Accepted for publication November 1986
SURVIVAL AND RECOVERY OF NORTHERN PINTAIL
231
-Klomath Basin
Gray Lodge
Suisun
Sa San Francisco
Bay
Los Bonos
Imperial Vaiiey-
FICURE 1. California pintail banding stations used in determining survival and recovery rate esti-
mates.
season after banding (Anderson 1975). Indirect recoveries are band recoveries
in subsequent years following the year of banding. All recoveries were wild birds
shot or found dead during the hunting season.
Survival rate is defined as the probability that a bird alive at the approximate
midpoint of the banding period in one year survives until the midpoint of banding
the following year. Recovery rate is defined as the probability that a banded bird
alive at the midpoint of the banding period in one year will be shot or found dead
the following hunting season and the Bird Banding Laboratory notified. It is as-
sumed that band reporting rates do not change during the study period.
Models of recovery data from all banding stations were examined, and the
model that best fit the data was presented by age and sex groups for each banding
station. A Z test (Brownie et al. 1978) was used to test for differences between
survival and recovery rates for different time periods. The level of significance
was P < 0.05.
232
CALIFORNIA FISH AND CAME
TABLE 1. Summary of 227,928 Pintails Banded in California from which Survival and Band Re-
covery Rates Were Estimated.
PRESEASON
Year
Number banded
Station
AM
IM
AF
IF
Total
Klamath Basin
Honey Lake
Gray Lodge
Suisun
So. S.F. Bay
Los Banos
Imperial Valley
1948-79
1950-59
1949-79
1951-58
1954-58
1948-64
1951-59
29,922
5,796
15,446
3,889
1,810
9,961
4,161
13,384
5,466
11,824
2,798
1,377
7,891
7,078
3,910
2,228
7,535
828
198
2,090
2,151
4,915
4,073
9,500
1,200
704
4,913
3,715
52,131
17,563
44,305
8,715
4,089
24,855
17,105
TOTAL
70,985
49,818
18,940
29,020
168,763
POSTSEASON
Cray Lodge
Los Banos
Imperial Valley
1954-79
1948-58
1951-73
15,672
2,277
17,523
8,607
1,491
13,595
24,279
3,768
31,118
TOTAL
35,472
23,693
59,165
RESULTS AND DISCUSSION
Adult and immature female recovery data from the South San Francisco Bay
station were insufficient to estimate survival and recovery rates. Also, no rea-
sonable model fit could be obtained for data sets of adult and immature females
from the Klamath Basin 1964-1979 and for adult and immature males from the
Suisun station. Estimates of parameters on an annual basis are subject to large
sampling variances and therefore average survival and recovery rates are pre-
sented (Table 2).
Adult males generally had higher survival rates than females. Survival rate es-
timates for preseason banded adult males ranged between 75.8 ± 8.2% on the
South San Francisco Bay station to 64.3 ± 4.0% at Imperial Valley. Survival rate
estimates for preseason banded adult females ranged between 65.6 ± 6.3% at
Honey Lake to 48.7 ± 7.3% at Imperial Valley. Survival rates for immature
pintails also indicate that males generally had higher survival rates than females.
Survival rate estimates for immature males ranged between 62.9 ± 7.4% for
birds banded at Gray Lodge to 49.5 ± 5.1% from Imperial Valley. Average sur-
vival rate estimates for immature females ranged between 69.0 ± 5.0% at Gray
Lodge to 36.0 ± 12.3% at Suisun. Fewer immature females were banded than
other age and sex classes, resulting in greater variance in survival rate estimates
for immature females than other classes. Adult males were banded and recov-
ered in large numbers; thus estimates for them are more precise.
Postseason banded pintail followed the same pattern of survival rates as those
from preseason bandings (Table 2). Survival rate estimates for postseason males
ranged between 77.0 ± 7.3% at Gray Lodge to 66.0 ± 2.2% at Imperial Valley.
Female survival ranged between 65.0 ± 2.6% at Cray Lodge and 50.8 ± 4.6%
at Los Banos.
Comparing banding stations, the average survival rate estimates for adult
males, adult females and immature males were lowest at Imperial Valley (Table
2). Most pintail banded in the Imperial Valley were from a population separate
from those banded in northern California and were more closely linked to the
Central and Mississippi flyways (Rienecker 1987). This suggests that pintail out-
side of the Pacific Flyway may have a lower survival rate. Conversely, Anderson
SURVIVAL AND RECOVERY OF NORTHERN PINTAIL 233
and Sterling (1974) found no difference in survival of adult male pintail banded
in south-central Saskatchewan (1955-1958) and recovered in Texas (69.11%)
and California (70.77%). However, their analysis assumed that birds wintering
in one area (e.g., Texas) did not shift to other areas (e.g., California) in subse-
quent years. California bandings showed that some birds used both areas
(Rienecker 1987).
Average estimated recovery rates for California banded adult male pintails
were 5% compared to 4% for adult females, 9% for immature males and 7% for
immature females (Table 2). Recovery rates are an index to harvest rates (Henny
and Burnham 1976).
Studies of mallard, American wigeon. Anas americana, and ring-necked duck,
Athya collaris, have also shown that survival and recovery rates of females are
lower than those of males (Anderson 1975, Rienecker 1976, Conroy and
Eberhardt 1983). The assumption is that females have higher non hunting mor-
tality, particularly during the nesting season, than that of males, thus resulting in
the lower survival and recovery rates. Males, in turn, are more intensively har-
vested and thus have greater band recovery rates. Differential migration (males
moving into higher harvest areas earlier than females), hunter selectivity for
males and fewer females available to hunters because of predation on the breed-
ing grounds are possible causes for higher band recovery rates of males.
Several factors could contribute to the variation in band recovery rates among
the banding stations but differences in hunting pressure, sample size and report-
ing rate are probably the main causes. Lower band recovery rates from
postseason vs. preseason banded birds occur because of the time period be-
tween banding and hunting seasons. Postseasons banded birds go through spring
migration and breeding before the hunting season starts and thus are subjected
to a greater preseason mortality resulting in fewer band recoveries from hunters.
Survival rates are based on the assumption that the same population is banded
each year. Many preseason banded birds are only passing through when banded,
and several subpopulations are probably banded at the same station during the
same period (Rienecker 1987). Thus, preseason banded samples may not pro-
vide accurate estimates for specific wintering populations. These data suggest
that postseason bandings were more representative of pintails wintering in the vi-
cinity of the banding station and less likely to contain several subpopulations.
The only significant difference between survival rates was for Klamath Basin
adult males that had higher survival rates during the 1950's (81%) than during
the 1960's (72%, Table 3). Recovery rates for Klamath Basin male pintails were
significantly lower during the 1960's (AM 2.7%-IM 6.2%) than during either the
1950's (AM 3.9%-IM 10.0%) or the 1970's (AM 3.7%-IM 7.8%). For all age
and sex classes banded both preseason and postseason at Gray Lodge, the only
significant difference was a higher immature male recovery rate in the 1950's
(8.9%) than in the 1960's (7.0%).
Mixing of subpopulations of pintails on the wintering grounds make manage-
ment by subpopulations difficult. The problem with managing several sub-
populations as a unit is that they could have varying harvest rates and/or
nonhunting mortality rates. While one subpopulation might be able to sustain an
increase in harvest, others might not. Thus, special management measures on
wintering populations would have to be formulated cautiously.
234
CALIFORNIA FISH AND GAME
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SURVIVAL AND RECOVERY OF NORTHERN PINTAIL
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SURVIVAL AND RECOVERY OF NORTHERN PINTAIL 237
Resumption of pintail banding in California is recommended to monitor sur-
vival and recovery rates in addition to monitoring migration and distribution pat-
terns. Banding was terminated in 1979 at a time when the population was trend-
ing downward. Biotelemetry of pintail in the Central and Imperial Valleys would
improve knowledge of daily and seasonal movements within these Valleys and
provide data on the area used by each wintering subpopulation.
ACKNOWLEDGMENTS
I thank members of the Waterfowl Studies Project, California Department of
Fish and Game, who trapped and banded pintails and to the staff of the Klamath
Basin NWRs and Salton Sea NWR for cooperation in the banding programs.
Thanks is also given to the U.S. Fish and Wildlife Service for furnishing computer
programs on survival and recovery rates, to R. Carpenter, California Department
of Fish and Game, who furnished a computer summary of recovery data on sur-
vival and to J. Nichols, USFWS Patuxent Wildlife Research Center, for sugges-
tions and interpretation of the data sets and for reviewing the manuscript. I also
wish to thank J. Bartonek, D. Gilmer, P. Law, D. Sharp, M. Miller and J. Fleskes
for review of this manuscript.
LITERATURE CITED
Anderson, D.R. 1975. Population ecology of the mallard: V. Temporal and geographic estimates of survival, re-
covery and harvest rates. U.S. Dept. inter.. Fish and Wildl. Serv. Resour. Publ. 125. 110 p.
, and R.T. Sterling. 1974. Population dynamics of molting pintail drakes banded in south-central
Saskatchewan. J. Wildl. Manage., 38(2): 266-274.
Belirose, F.C. 1976. Ducks, geese and swans of North America. Stackpole Books, Harrlsburg, PA. 544 p.
Brownie, C, D.R. Anderson, K.P. Burnham and D.S. Robson. 1978. Statistical inference from band recovery data —
a handbook. U.S. Fish and Wildl. Serv., Resour. Publ. 131. 212 p.
Conroy, M.J. and R.T. Eberhardt. 1983. Variation in survival and recovery rates of ring-necked ducks. J. Wildl. Man-
age., 47: 127-137.
Henny, C.J. and K.P. Burnham. 1976. A mallard reward band study to estimate band reporting rates. J. Wildl. Man-
age., 40(1): 1-14.
Rienecker, W. C. 1976. Distribution, harvest and survival of American wigeon banded in California. Cal. Fish and
Game, 62(2): 141-153.
. 1987. Migration and distribution of pintails banded in California. Cal. Fish and Game. 73(3): 139-155.
238 CALIFORNIA FISH AND CAME
Calif. Fish and Came 71{A): 238-243 1987
MANAGEMENT OF MIDGES AND OTHER INVERTEBRATES
FOR WATERFOWL WINTERING IN CALIFORNIA^
NED H. EULISS, JR.2
Department of Fisheries and Wildlife
Oregon State University
Corvallis, Oregon 97331
and
CAIL CRODHAUS
California Department of Health Services
Berkeley, California 94704
A review of recent waterfowl food habit studies showed that invertebrates are of
major dietary importance to ducks wintering in California. However, current wetland
practices are directed at production of plant foods and seldom consider the prop-
agation of invertebrates. We suggest that invertebrate repopulation of seasonally
flooded marshes will occur more rapidly if an inoculum of invertebrates is provided
via small ponds flooded several weeks before general marsh flooding in fall. Man-
agers will require considerably more information before management of aquatic in-
vertebrates can be fully developed.
INTRODUCTION
Invertebrates, principally midge larvae (Chironomidae) are important
waterfowl foods during the breeding season (Bartonek 1972, Krapu 1974,
Landers et al. 1977, Reinecke and Owen 1980, Sugden 1973, Swanson and
Bartonek 1970, Swanson et al. 1977). It is less well known that northern pintails.
Anas acuta acuta, mallards, A. platyrhynchos, and green-winged teal, A. crecca
carolinensis, consume significant amounts of midge larvae and other inverte-
brates during the non-breeding period in California (Beam and Gruenhagen
1980, Connelly and Chesemore 1980, Pederson and Pederson 1983, Euliss and
Harris 1987, Miller 1987). Consequently, it may be desirable to increase inver-
tebrate production in California marshes as a means of raising the carrying ca-
pacity of wetlands for waterfowl. This is especially important because of con-
tinued wetland losses (Tiner 1984) and the need to manage remaining wetlands
more efficiently (Bellrose and Low 1978). Although midges and other inverte-
brates have been the subject of many studies, the management of midge pop-
ulations for waterfov^l has received little attention.
The objectives of this paper are to assess the potential of managing inverte-
brates for waterfowl, to summarize the pertinent literature, and to offer prelim-
inary suggestions for v^^etland management designed to increase production of in-
vertebrates in seasonally flooded and permanently flooded marshes in
California's Central Valley. Additionally, general midge ecology, vegetative sub-
strates that are suitable for invertebrate colonization, and human health consid-
erations are discussed. Lastly, "brood-stock ponds" (BSP's) are introduced as a
conceptual method that managers may use to increase repopulation rates of
aquatic invertebrates in seasonally flooded marshes.
^ Accepted for publication May 1987.
2 Present address: U.S. Fish and Wildlife Service, Northern Prairie Wildlife Research Center, 6924 Tremont Road,
Dixon, California 95620
MANAGEMENT OF INVERTEBRATES FOR WATERFOWL 239
GENERAL ECOLOGY OF MIDGES
Midges and other aquatic invertebrates use a wide range of microhabitats in
marshes. Plant substrates, however, offer the best management potential because
invertebrates could be produced along with waterfowl food plants. Previous
workers have suggested that the abundance and diversity of aquatic invertebrates
increase with plant biomass (Krecker 1939, Berg 1949, McGaha 1952, Rosine
1955, Darby 1962, Krull 1970, Magy et al. 1970, Lamberti and Resh 1984). Fur-
ther, the density and diversity of invertebrate populations fluctuate seasonally
and according to plant species and physiological state. With certain exceptions
(e.g. obligate benthic forms), most marsh-inhabiting midge larvae are epiphytic
and forage mainly on epiphytic algae and metaphyton. Foraging midges can sig-
nificantly reduce epiphytic algae biomass (Cattaneo 1983) while interfering little
with the growth of macrophytes. Structurally complex plants are preferable to
simple ones, because the former provide more suriface area for colonization of
epiphytes and generally harbor greater numbers of midges.
Although midge abundance on specific plants has been studied, relatively few
workers have examined the abundance of these insects on common waterfowl
food plants. High densities of midge larvae have been observed in stands of sago
pondweed, Potamogeton pectinatus, horned pondweed, Zannichellia palustris,
southern naiad, Najas guadalupensis, common burhead, Echinodorus
cordifolius, and common widgeongrass, Ruppia maritima, (Gerry 1954, Darby
1962, Magy etal. 1970, Lamberti and Resh 1984, Grodhaus, unpubl. data). Darby
(1962) reported that living stands of tule bulrush, Scirpus acutus, common cat-
tail, Typha latifolia, and common barnyardgrass, Echinochloa crusgalli, were rel-
atively unproductive of midges. However, Euliss (1984) observed that midges,
Cricotopus sp., were abundant in stands of senescent common barnyardgrass.
High densities of midge larvae have been reported in decomposing substrates
provided by other plant species (Danell and Sjoberg 1979, Pederson and
Pederson 1983).
MANAGEMENT CONSIDERATIONS
General
Wetland managers should develop management plans directed at particular
species of midges because some species are much more productive than others.
For example, there were at least 7 midge species present on Kern National Wild-
life Refuge (NWR), California (Euliss 1984) yet only 4, Chironomus stigmaterus,
C. decorus, Cricotopus sp., and Apedilum subcinctum, dominated the midge
biomass in waterfowl diets. Similar findings were reported by Pederson and
Pederson (1983) on Lower Klamath NWR, California. Therefore, we encourage
managers of waterfowl areas to identify local midge populations as a basis from
which to develop management efforts.
Surveys to determine general invertebrate community structure in marshes
would be profitable. Survey techniques and invertebrate identification keys are
included in Usinger (1956), Ward and Whipple (1959), Pennak (1978), and
Merritt and Cummins (1984). The keys to genera of Chironomidae in
Wiederholm (1983) and Coffman and Ferrington (1984) cover wide geographic
areas. Several useful keys to midge species are available which cover specific
habitats or geographic areas (Roback 1957, Darby 1962, Grodhaus 1967, Mason
1968, Oliver et al. 1978, Simpson and Bode 1980).
240 CALIFORNIA FISH AND GAME
Seasonally Flooded Marshes
In seasonal marshes, the time required for midges to establish populations is
extremely variable and unpredictable. The life cycle of most midges includes
standing water and freshly laid eggs (Oliver 1971 ). This suggests that the rate of
repopulation in wetlands would depend on the availability of suitable species in
nearby wetlands at the time of flooding. In seasonal marshes, repopulation is a
passive process in which viable midge eggs and larvae are introduced into freshly
flooded marshes via the water used for ponding and/or from adults flying in from
surrounding areas. In either case, there may be considerable variation in the spe-
cies and numbers available to colonize marshes from year to year. Midge eggs
or larvae may be present in some water sources but not others. In years of severe
drought, wells may be the only water source and the establishment of adequate
midge populations may be delayed because this water is free of midges. The
availability of midges and other invertebrates in purchased water sources is un-
certain. During normal and drought years, wetland basins surrounding seasonal
marshes may be dry, and few midges would be available to lay eggs in freshly
flooded marshes. In years of above normal precipitation and runoff, midge spe-
cies available in nearby wetlands may not be well suited to the particular habitats
flooded on a waterfowl area or they may be present in insufficient densities to
enable rapid repopulation.
As an alternative to the passive repopulation of invertebrates just described,
we suggest the use of brood-stock ponds (BSP's) as an active restocking ap-
proach. We define BSP's as subunits of main ponds that are flooded 1-2 months
before remaining habitats. These ponds should have the same basic vegetative
composition as main ponds and they would serve as culturing sites for inverte-
brates that invade from outside sources. Ideally, BSP's would be established
within a main pond that is used to convey water to other ponds of similar veg-
etative composition. Thus, when remaining habitats are flooded, the inverte-
brates present in BSP's would be introduced into freshly flooded wetlands. In
waterfowl areas that flood a variety of habitat types, we recommend that BSP's
be established in each of the habitat types provided. This should allow a more
rapid colonization of freshly flooded wetlands than occurs presently because it
would provide an inoculum of invertebrates that are specifically adapted to par-
ticular vegetative types in managed wetlands.
In California, seasonal marshes are generally flooded during late summer or
early fall when water temperatures are high. With the onset of winter, midge de-
velopment and production of egg-laying adults slows because of lower water
temperatures (Oliver 1971 ). Thus, BSP's may increase the biomass of midge lar-
vae produced in seasonal marshes during the winter because more eggs would
be deposited in marshes before the onset of cold weather. The objective of this
plan would be to maximize midge biomass in initial generations when water tem-
peratures are favorable. Warm water temperatures (24 C) may allow adult
midges to develop from eggs in as little as 2 weeks (Euliss 1984).
Seasonal marshes in California are normally flooded 6-8 months each year.
The winter diets of pintails and green-winged teal consist of a substantial pro-
portion of midge larvae about 2 months after the ponds are flooded in the fall
(Connelly and Chesemore 1980, Euliss and Harris 1987). This delay is likely
caused because midge populations are low initially and available adults are in-
sufficient to saturate the marshes with eggs. Assuming a 6 month period of sea-
MANAGEMENT OF INVERTEBRATES FOR WATERFOWL 241
sonal inundation, BSP's have the potential to increase the availability of midge
larvae over an additional 33% of the wintering period.
Research is needed to evaluate BSP's and to identify specific features of prac-
tical innportance to wetland managers. Because of the great reproductive po-
tential of midges (Oliver 1971 ), relatively few adults are required to generate
large populations of larvae. Thus, a relatively small area may be required for BSP's,
but the exact size relative to seasonal marsh types is uncertain.
Permanently Flooded Marshes
The management of invertebrates in permanent marshes can also provide ad-
ditional foods for waterfowl. Both nonvegetated (i.e. phytoplankton dominated)
and vegetated (i.e. submersed aquatic macrophyte dominated) habitats can be
extremely productive of midges and other invertebrates. Most species of sub-
mersed waterfowl food plants provide large surface areas for invertebrate col-
onization. However, the invertebrates present in deep-water marshes may not be
highly available to waterfowl. Lowering of water levels to provide numerous shal-
low areas would concentrate invertebrates and enhance their availability to dab-
bling ducks. The objective of water level manipulation would be to increase the
availability of invertebrates during periods of high waterfowl use and when the
nutritional demands of waterfowl for animal foods are high. However, complete
drawdowns have detrimental effects on aquatic invertebrates (Kadlec 1962) and
a reestablishment period would be required after reflooding.
Permanently flooded marshes often contain abundant fish populations that
may include predators on midge larvae. Mosquitofish, Cambusia affinis, do not
consume significant numbers of midges (Bay and Anderson 1966), but it is likely
that most other eurythermal fish prey on chironomid larvae. Threespine
stickleback, Gasterosteus aculeatus, carp, Cyprinus carpio, and goldfish, Caras-
sius auratus, are particularly efficient midge predators (Bay and Anderson 1965,
Fleming and Schooley 1984) and should be discouraged from permanent marsh
impoundments.
HUMAN HEALTH AND NUISANCE CONSIDERATIONS
The goal of wetland managers should be to enhance the productivity and avail-
ability of midge larvae and pupae rather than to produce adult insects. As men-
tioned previously, relatively few adults are required to produce large populations
of larvae. There are certain adverse consequences of excessive numbers of adult
midges near human activities. The allergenic potential of inhaled fragments
( Bauer et al. 1 983 ) and the possibility of highway accidents due to obscured vis-
ibility (Mortenson et al. 1967) are important problems, but property defacement
is the most frequent unwanted outcome of excessive midge production (AM
1980).
The most serious insect problem associated with waterfowl management is
mosquito production. Plans to enhance populations of midges and other inver-
tebrates on wintering areas in California are not expected to create serious con-
flicts with mosquito-control interests. Although there is some overlap with the
mosquito season during the fall and spring, the management of midges would be
conducted mostly during the winter months when mosquito populations are typ-
ically low. Moreover, plans to enhance midge production (e.g. BSP's) would not
require flooding main ponds any earlier than currently practiced. However,
242 CALIFORNIA FISH AND CAME
wetland managers should incorporate sound mosquito-control practices into
management programs. Because of the diversity of mosquito species and habitat
requirements, we recommend that wetland managers coordinate their efforts
with local mosquito abatement districts to minimize mosquito production.
CONCLUSION
The management of midges and other aquatic invertebrates is in its infancy and
many aspects of specific strategies have not been developed. However, the po-
tential benefit for wintering waterfowl and other wildlife is great. Considerable in-
novation by both managers and researchers will be required to develop practical
and effective invertebrate management programs. We encourage the various
agencies to obtain accurate records of midge and other invertebrate usage of spe-
cific plant types and in areas where specific management strategies are practiced.
Plans to enhance invertebrate populations appear feasible and results from recent
research efforts should enable managers to develop initial plans at a fairly rapid
pace.
ACKNOWLEDGMENTS
We wish to thank R. Alls, T. Charmley, J. Houk, G. Kramer, S. Mulligan, and
R. Parman for assistance; F. Bellrose, B. Coblentz, D. Connelly, B. Euliss, D.
Gilmer, S. Harris, J. Hicks, G. Lamberti, M. Miller, E. Mortenson, P. O'Halloran,
G. Pederson, P. Pederson, C. Smith, P. Springer, G. Swanson, and R. Yescott for
editorial review; and Northern Prairie Wildlife Research Center at Dixon, Cali-
fornia for providing travel services and technical support.
LITERATURE CITED
All, A. 1980. Nuisance chironomids and their control: a review. Bull. Entomol. Soc. Amer. 26:6-16.
Bartonek, J. C. 1 972. Summer foods of American widgeon, mallards, and a green-winged teal near Great Slave Lake,
N.W.T. Can. Field-Nat. 86:373-376.
Bauer, X., M. Dewair, K. Haegele, H. Prelica, A. SchoN, and H.Tichy. 1983. Common antigenic determinants of he-
moglobin: a basis of immunological cross reactivity between chironomid species (Diptera: Chironomidae).
Clin. Exp. Immunol. 54:599-607.
Bay, E. C, and L. D. Anderson. 1965. Chironomid control by carp and goldfish, Mosq. News 25:310-316.
1966. Studies with the mosquitofish Cambusia affinis as a chironomid control. Ann. Ent. Soc. Am.
59:150-153.
Beam, )., and N. Cruenhagen. 1980. Feeding ecology of pintails (Anas acuta) wintering on the Los Banos Wildlife
Area, Merced County, California. Calif. Dep. Fish and Came, Fed. Aid Wildl. Restor. Prog. Rep., Proj. W-40-
D-1. 23pp.
Bellrose, F. C, and ). B. Low. 1978. Advances in waterfowl management research. Wildl. Soc. Bull. 6:63-72.
Berg, C. 0. 1949. Limnological relations of insects to plants of the genus Potamogeton. Trans. Am. Microscop. Soc.
68:279-291 .
Cattaneo, A. 1983. Grazing on epiphytes. Limnol. Oceanogr. 28:124-132.
Coffman, W. P., and L. C. Ferrington. 1984. Chironomidae. Pages 551-652 in R. W. Merritt and K. W. Cummins
(eds.). An introduction to aquatic insects of North America. 2nd ed. Kendall Hunt Publ., Dubuque, Iowa.
Connelly, D. P., and D. L. Chesemore. 1980. Food habits of pintails. Anas acuta, wintering on seasonally flooded
wetlands in the northern San Joaquin Valley, California. Calif. Fish and Game 66:233-237.
Danell, K., and K. Sjoberg. 1979. Decomposition of Carex and Equisetum in a northern Swedish lake: dry weight
loss and colonization by macro-invertebrates. J. Ecol. 67:191-200.
Darby, R. E. 1962. Midges associated with California rice fields, with special reference to their ecology. Hilgardia
32:1-206.
Euliss, N. H., )r. 1984. The feeding ecology of pintail and green-winged teal wintering on Kern National Wildlife Ref-
uge. M. S. Thesis, Humboldt State Univ., Areata. 188pp.
, and S. W. Harris. 1987. Feeding ecology of northern pintails and green-winged teal wintering in Cali-
fornia. J. Wildl, Manage. 51: 724-732.
MANAGEMENT OF INVERTEBRATES FOR WATERFOWL 243
Fleming, K. J., and J. K. Schooley. 1984. Foraging patterns and prey selection by marsh fish. Proc. Calif. Mosq. Vector
Control Assoc. 52:139.
Gerry, B. I. 1954. Ecological conditions which influence control of mosquito-like pests. Mosq. News 14:145-149.
Grodhaus, G. 1967. Identification of chironomid midges commonly associated with waste stabilization lagoons in
California. Calif. Vect. Views 14:1-12.
Kadlec, ). A. 1962. Effects of a drawdown on a waterfowl impoundment. Ecology 43:267-281.
Krapu, C. L. 1974. Foods of breeding pintails in North Dakota. J. Wildl. Manage. 38:408-417.
Krecker, F. H. 1939. A comparative study of the animal population of certain submerged aquatic plants. Ecology
20:553-562.
Krull, J. N. 1970. Aquatic plant-macroinvertebrate associations and waterfowl. J. Wildl. Manage. 34:707-718.
Landers, J. L., T. T. Fendley, and A. S. Johnson. 1977. Feeding ecology of wood ducks in South Carolina. ]. Wildl.
Manage. 41:118-127.
Lamberti, G. A., and V. H. Resh. 1984. Seasonal patterns of invertebrate predators and prey in Coyote Hills Marsh.
Proc. Calif. Mosq. Vector Control Assoc. 52:126-128.
Magy, H. I., G. Grodhaus, J. D. Gates, and J. Montez. 1970. Pondweed — a substrate for chironomids, especially
Paralauterborniella subcincta. Proc. 37th Ann. Conf. Calif. Mosq. Assoc, Jan. 27-29, 1969. 37:115-119.
Mason, W. T. 1968. An introduction to the identification of chironomid larvae. Fed. Water Pollut. Control Adm. Dist.
by NTIS, U. S. Dept. of Commerce, Springfield, VA 87pp.
McGaha, Y. J. 1952. The limnological relations of insects to certain aquatic flowering plants. Trans. Am. Microscop.
Soc. 71:355-381.
Merritt, R. W., and K. W. Cummins (eds.). 1984. Aquatic insects of North America. 2nd ed. Kendall Hunt Publ.,
Dubuque, Iowa. 441 pp.
Miller, M. R. 1987. Fall and winter foods of northern pintails on three northern California refuges. J. Wildl. Manage.
51:403-^12.
Mortenson, E. W., G. Grodhaus, C. M. Myers, and D. C. White. 1967. A report on the San Luis midge problem,
Merced County, California. Department of Public Health, Bureau of Vector Control, 10pp. (mimeographed).
Oliver, D. R. 1971. Life history of the Chironomidae. Ann. Rev. Entomol. 16:211-230.
, D. McClymont, and M. E. Roussel. 1978. A key to some larvae of Chironomidae (Diptera) from the
Mackenzie and Porcupine River watersheds. Ottawa, Ontario. Canada. Fish, and Mar. Serv. Tech. Rep. No.
791. 73pp.
Pederson, G. B., and R. L. Pederson. 1983. Feeding ecology of pintails and mallards on Lower Klamath marshes.
Final rept. on U. S. Fish and Wildl. Serv. contract 14-16-001-79106. Humboldt State Univ. Foundation, Areata.
89pp.
Pennak, R. W. 1 978. Fresh-water invertebrates of the United States. 2nd ed. John Wiley and Sons, New York. 803pp.
Reinecke, K. J., and R. B. Owen, Jr. 1980. Food use and nutrition of black ducks nesting in Maine. J. Wildl. Manage.
44:549-558.
Roback, S. S. 1957. The immature tendipedids of the Philadelphia area. Monogr. Acad. Nat. Sci., Philadelphia
9:1-152.
Rosine, W. N. 1955. The distribution of invertebrates on submerged aquatic plant surfaces in Muskee Lake, Col-
orado. Ecology 36:308-314.
Simpson, K., and R. Bode. 1980. Common larvae of Chironomidae (Diptera) from New York streams and rivers,
with particular reference to the fauna of artificial substrates. Bull. New York State News. 439:1-105.
Sugden, L. G. 1973. Feeding ecology of pintail, gadwall, American widgeon and lesser scaup ducklings in southern
Alberta. Can. Wildl. Serv, Rept. Series No. 24. 43pp.
Swanson, G. A., and J. C. Bartonek. 1 970. Bias associated with food analysis in gizzards of blue-winged teal. J. Wildl.
Manage. 34:739-746.
, G. L. Krapu, and J. R. Serie. 1977. Foods of laying female dabbling ducks on the breeding grounds. Pages
47-57 in T. A. Bookhout, (ed.). Waterfowl and wetlands: An integrated review. Proc. 1977 Symp., Madison,
Wis., N. Cent. Sect., The Wildl. Soc.
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Washington, D.C. 59pp.
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Ward, H. B., and G. C. Whipple. 1959. Fresh-water biology. 2nd ed. John Wiley and Sons, New York. 1248pp.
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244 CALIFORNIA FISH AND GAME
Calif. Fish and Came 73(4): 244-247 1987
NOTES
YELLOWTAIL CHAFING ON A SHARK: PARASITE REMOVAL?
Ectoparasites are commonly found on a variety of pelagic fishes. That they
cause irritation has not been determined through experimentation, but has pro-
vided the basis of explanations that: ( i ) attribute leaping by mantas ( Mobulidae)
and billfishes (Istiophoridae) to attempts at dislodging parasites (Bohike and
Chaplin 1968, Walford 1974) and (ii) attribute chafing on substrates in at least
30 families of fishes to removal of sources of irritation (Wyman and Walters-
Wyman 1985). In many pelagic fishes vigorous leaping is not observed. Further-
more, they do not have ready access to substrates or cleaning fishes or inver-
tebrates (usually restricted to inshore areas). Although there does not appear to
be any published documentation of chafing in pelagic fishes there are various ob-
jects in the pelagic environment which might serve as "chafing posts" against
which parasites could be dislodged.
On 30 August 1986 at approximately 1500 h I observed a group of 8-15
yellowtail, Seriola lalandi, (Carangidae), at the surface of Bahia San Hippolito on
the west coast of Baja California, Mexico. They were swimming alongside a small
( 1 .5-1 .6 m ) blue shark, Prionaceglauca, and apparently using the shark as an ob-
ject for chafing. The yellowtail were continuously bunched along both sides of
the posterior half of the shark. They repeatedly rushed up to the shark, made
side-to-side contact and swam with an obvious rubbing motion toward the an-
terior end of the shark. The chafing was always disengaged prior to passing the
shark's pectoral fin.
That dislodging of ectoparasites was the purpose of the behavior could not be
determined with certainty; however, I ruled out aggression because the shark
continued swimming slowly throughout the entire 3-4 min that the behavior was
observed. No obvious change in swimming speed or direction on the part of the
shark resulted from physical contact by the yellowtail, but minor lateral displace-
ment caused by the chafing itself was evident. Examination of several (ca. 30)
yellowtail caught that afternoon in the same location revealed that ectoparasitic
copepods were present in low numbers (1 or 2) on approximately 60-70% of
the yellowtail.
ACKNOWLEDGMENTS
This study was supported by a grant from the L. L. Stewart Faculty Develop-
ment Fund, OSU Foundation, and by Oregon Agricultural Experiment Station
Project ORE00925. I thank C. E. Bond, D. M. Markle, C. B. Schreck, and P. C.
Sikkel for suggestions that improved the quality of the manuscript. This is Oregon
Agricultural Experiment Station Tech. Report No. 8083.
LITERATURE CITED
Bbhike, J.E. and C.C.C. Chaplin. 1968. Fishes of the Bahamas and adjacent tropical waters. Livingston Publishing
Co.: Wynnewood, Pennsylvania. 771 p.
Walford, L.A. 1974. Marine game fishes of the Pacific coast from Alaska to the Equator. T.F.H. Publications:
Neptune, New Jersey. 205 p
Wyman, R.L., and MP. Walters-Wyman. 1985. Chafing in fishes: occurrence, ontogeny, function and evolution.
Env. Biol. Fish. 12:281 -289. versity, Nash 104, Corvallis, OR 97331-3803. Accepted for publication May 1987.
— Bruce E. Coblentz, Department of Fisheries and Wildlife, Oregon State Uni-
versity, Corvallis, OR 97331-3803. Accepted for publication May 1987.
NOTES 245
ATYPICAL PLUMAGE OF A FEMALE CALIFORNIA QUAIL
Reversal of secondary sex characteristics, most noteably plumage color and
pattern, has been reported in several species of gallinaceous birds, including ring-
necked pheasants, Phasianus colchicus (Bent 1932), Japanese quail, Coturnix
coturnix japonica (Kannankeril and Domm 1968), and northern bobwhites,
Colinus virginianus (Brodkorb and Stevenson 1934, Buchanan and Parkes 1948).
There is, however, a paucity of information relating to the reversal of plumage in
other Phasianidae. Herein we report on an adult, female California quail, Cal-
lipepla californica, with mixed male and female plumage characters that was col-
lected on 7 November 1 986 at the E.E. Wilson Wildlife Management Area, 1 5 km
north of Corvallis, Benton County, Oregon.
The plumage of this bird, which had completed her annual molt, contained
some feathers that were typically female, some that were male-like, and others
that were intermediate between the sexes (Figure 1 ). The throat patch was a
mixture of black (male-like) and grayish-brown (female-like) feathers sur-
rounded by a white stripe that contained some brown feathers and lacked the
distinct edge of the male. The crown was light chestnut with some streaking and
the forehead was brown with a dirty white background. A white stripe extended
from the forehead to the black auricular feathers. The nape was female-like in ap-
pearance except that the brown edges of the feathers were darker than normal.
Three of the topknot feathers were elongated and intermediate in length between
male and female; the remaining feathers of the crest were female-like. The breast
was a mixture of brown (female-like) and gray (male-like) feathers. The upper
abdominal feathers were light tan (intermediate between male and female) and
the borders of the "scaled" abdominal feathers were black (male-like). The
lower abdomen lacked the chestnut patch characteristic of males.
Plumage of female gallinaceous birds is controlled, or at least influenced, by
female sex hormones, which are produced largely by the ovary (Voitkevich
1966). Male-like plumage in females has been associated with pathogenic re-
gression (Witschi 1961 ), atrophication with age (Bent 1932), and abnormal en-
largement (Buchanan and Parkes 1948) of the ovary, as well as sinistral
ovarectomization (Kannankeril and Domm 1968). Gross and histological exam-
ination of the reproductive tract of this female California quail revealed the pres-
ence of an oviduct (left only) of normal size and cellular structure for a non-
laying hen. No gonads or other accessory structures were found. The ovary
apparently either regressed to a size that we were unable to locate or was absent.
The scant amount of information about the reversal of plumage characters in
female quail indicates this phenomenon is unusual. This bird represents the only
example we have observed of a California quail with atypical plumage among ap-
proximately 450 birds from this study area that have been examined from 1975
through 1986.
The skin of this California quail was deposited in the wildlife collection of the
Department of Fisheries and Wildlife, Oregon State University (Specimen No.
FW 5153). Appreciation is expressed to R.L. Jarvis for his review of the manu-
script. This paper is Technical Publication No. 8120 of the Oregon Agricultural
Experiment Station.
246
CALIFORNIA FISH AND GAME
FIGURE 1. Dorsal and ventral views of typical male (left), typical female (right), and female Cal-
ifornia quail with mixed male and female plumage characters (center) taken in Benton
County, Oregon, 7 November 1986.
NOTES 247
LITERATURE CITED
Bent, A.C. 1932. Life histories of North American gallinaceous birds. U.S. Nat. Mus. Bull, 162. 490 p.
Brodkorb, P. and J. Stevenson. 1934. Additional northeastern Illinois notes. Auk, 51:100-101.
Buchanan, F.W. and K.C. Parkes. 1948. A female bob-white in male plumage. Wilson Bull., 60:119-120.
Kannankeril, J.V. and L.V. Domm. 1 968. The influence of gonadectomy on sexual characteristics in Japanese quail.
). Morphol., 126:395-412.
Voitkevich, A.A. 1966. The feathers and plumage of birds. Sidgwick and Jackson, London. 331 p.
Witschi, E. 1 961 . Sex and secondary sexual characters. Pages 1 1 5-1 68 in A.J. Marshall, ed. Biology and Comparative
Physiology of Birds. Academic Press, London.
— / A. Crawford, P. J. Cole, and K. M. Kilbride, Department of Fisheries and
Wildlife, Oregon State University, Corvallis, Oregon 97337 and A. Fair brother,
U.S. Environmental Protection Agency, Corvallis, Oregon 97330. Present ad-
dress for P. J. Cole: Idaho Department of Fish and Game, Jerome, Idaho 83338.
Accepted for publication June 1987.
248 CALIFORNIA FISH AND GAME
BOOK REVIEWS
MARINE MAMMALS (OF THE EASTERN NORTH PACIFIC AND ARCTIC WATERS)
Edited by Delphine Haley. Second Edition, Revised. Pacific Search Press, Seattle, Washington,
1986, 2% p.; $22.95 paper.
This brief treatment of each of the marine mammals of the eastern North Pacific is very good. It
could function nicely as a text for an introductory course in marine mammals of the eastern North
Pacific, and would be a good reference book for students of these mammals at all levels.
The book is abundantly illustrated with photographs and range maps. The photographs, while gen-
erally splendid, have apparently sometimes lost a bit of their crispness in the reproduction process.
Occasionally some of the detail that is described as clearly visible, clearly is not.
The list of contributors would provide a good start on a "who's who" in marine mammal research.
One can feel the excitement that many of the authors have for their work. Although a "protectionist"
philosophy occasionally permeates a section to the extent that real resource conflicts are relegated
to ranting by overzealous fishermen, on the whole the book objectively presents information, point-
ing out adequacies and inadequacies.
I recommend the book to anyone interested in marine mammals.
—Jack A. Ames
DISEASES AND PARASITES OF MARINE FISHES
by H. Moller and K. Anders, Verlag Moller, Kiel, 1986, 365 p. Illustrated, cloth 50 DEM (ca. $25
US).
This book represents an ambitious effort to provide basic information to the layman and still be
of interest to the professional. I believe that Moller and Kiel have succeeded. The authors had the
foresight to have specialists review the various sections, thus avoiding obvious mistakes. There are
many good line drawings and more than 200 photographs, some in color, which should help the lay-
man identify micro- and macroparasites and the causes of skeletal abnormalities and tumors. The
book includes sections on the techniques used in fish parasitology and lists of current fish health text-
books and journals. Workers in fish disease and parasitology will appreciate the bibliography which
follows each section and the many tables and figures.
The chapters include such topics as parasites as biological tags, human pathogens transferred by
fish and spoilage of fish due to parasites. I found the sections on epidemiology and pollution, the
authors' specialties, especially informative. The case study of fish disease in the Elbe Estuary dem-
onstrates the complex relationship between pollution, disease and natural parameters.
As a suggestion for future editions, the authors might include a section on the fast-growing field
of fish immunology. I recommend this book; it is well worth the price.
— Mike Moser
EIDER DUCKS IN CANADA
Edited by Austin Reed. 1986. Canadian Wildlife Service Report Series Number 47, Ottawa.
1977. $23.50.
This is another in the excellent series of reports by the Canadian Wildlife Service dealing with var-
ious aspects of wildlife biology and management in Canada. This publication consists of 18 separate
papers which cover eider status and ecology, most concentrating on the Common Eider, Somateria
mollissima, with some information presented on King Eider, 5. spectabilis. Twelve of the papers are
presented in English and five in French, with abstracts in the second official language. The final paper
is written in both languages.
The report is organized into six parts. The eight papers of Part I cover distribution and abundance,
with each paper discussing a separate population in geographical order from east to west. Part II con-
sists of two papers on identification and distribution of eastern races of Common Eiders. The single
paper in Part III discusses winter numbers and distribution of Gulf of St. Lawrence eiders. Part IV con-
tains four papers on ecology, primarily on the breeding grounds, with one devoted to Inuit knowledge
of Common Eider ecology. There are two papers on use of eiders by people in Part V, and Part VI,
titled Conclusions, contains a single paper summarizing population size and status of Common Eiders
in eastern North America. It incorporates the findings of the other papers in the publication to create
a population model for six subpopulations of Common Eiders, with recommendations for further re-
search and management.
BOOK REVIEWS 249
This is a technical report with copious maps, figures and tables. It is well written and up to the usual
scholarly standards of the Canadian Wildlife Service. However, except for someone with an all-
consuming interest in eiders, because of the high cost of the report most readers might want to peruse
a copy from their local government repository library, rather than buy it.
— Bruce £ Deuel
OCEAN FORUM
By Ron I. Jackson and William F. Royce; Fishing News Books Ltd, Farnham-Surrey-England
1986; 240 p. $31.50
An exceptional amount of information on North Pacific resources and fisheries from the fur trade
period to the present is contained in this 240 page book. The authors provide "an interpretive history
of the International North Pacific Fisheries Commission." From the chaos of war amid national dif-
ferences in fishing methods, policies, and objectives emerged the International Convention for the
High Seas Fisheries of the North Pacific Ocean among Japan, Canada, and the United States in 1952.
The Convention established the International North Pacific Fisheries Commission (INPFC) to pro-
mote and coordinate scientific studies to ascertain conservation measures to secure the maximum
sustained productivity and each nation would carry out such recommendations. A feature of the
Convention was the unprecedented principle of abstention, contrary to the prevailing concepts of
freedom of the seas. Japan and Canada agreed to abstain from fishing in named Convention areas;
the major abstention was by Japan for salmon east of 175° West Longitude.
Confrontations in negotiations and renegotiation sessions and in annual meetings are described as
are the collaborations and cooperation in the massive fishery research programs carried out by mem-
ber nations. Interwoven in the history of INPFC are early dissatisfactions of national fishing groups,
the reemergence of Japan as the world fishing leader, new principles of law of the seas, extended ju-
risdictions, the naturation of fishery science, the explosive growth of north Pacific fisheries, changing
business practices and markets, and the development of respect and trust among participants.
The focus in early years of INPFC was on salmons, halibut, and herring. In latter years the tre-
mendous groundfish resources of the north Pacific greatly influenced the actions of INPFC.
Details are provided on individuals, their perspectives and their roles in forging the direction of
INPFC. Summaries of fisheries, fishery biology anhd oceanography in the north Pacific are succinct
and bring the reader abreast of the past and current major fisheries from California to the Bering Sea,
with emphasis on the North Pacific.
The appendices contain the 1953 the 1979 Conventions, past and present INPFC commissioners
and secretariats, biographies of major participants, and a list of INPFC publications. The preface and
epilogue are concise summaries of the formation, activities, accomplishments, and future of the
INPFC. The details and comprehensive features of this book add to the reader's understanding of
INPFC and its role.
Both authors had substantive roles in INPFC. Jackson was the first permanent Executive Director
and Royce served as an U.S. advisor to commissioners and as an expert on the Commission's Biology
and Research Committee. They contend that INPFC, the Ocean Forum, will be needed now more
than ever with current dynamic changes in fisheries, national jurisdictions, and fish businesses.
Many current fisheries from central California to the Bering Sea now have multi-national process-
ing and marketing features. Anyone involved with fisheries of the northeastern and North Pacific will
benefit from reading this book.
— Tom Jow
250 CALIFORNIA FISH AND GAME
INDEX TO VOLUME 73
AUTHORS
Allen, Sarah C: see Webber and Allen, 60-61.
Asay, Christopher E.: Habitat and Productivity of Cooper's Hawks Nesting in California, 80-87.
Ault, Jerald S., and )ohn D. DeMartini: Movement and Dispersion of Red Abalone, Haliotis rufescens, in Northern
California, 196-213.
Brooks, Andrew J.: Two Species of Kyphosidae Seen in King Harbor, Redondo Beach, California, 49-50.
Coblentz, Bruce E.: Yellowtail Chafing on a Shark: Parasite Removal?, 244.
Cole, P.J.: see Crawford, Cole, Kilbride, and Fairbrother, 245-247.
Compagno, Leonard ).V.; see Ebert, Compagno, and Natanson, 117-123.
Crawford, J.A., P.J. Cole, K.M. Kilbride, and A. Fairbrother; Atypical Plumage of a Female California Quail, 245-247.
DeMartini, John D.: see Ault and DeMartini, 196-213.
Dole, Jim W.: see Perry, Dole, and HoH, 156-162.
Duhamel, G.E.: see Kent, Duhamel, Foott, and Hedrick, 99-105. Ebert, David A., Leonard J.V. Compagno, and Lisa
J. Natanson: Biological Notes on the Pacific Sleeper Shark, Somniosus pacificus (Chondrichthyes: Squalidae),
117-123.
Erickson, Daniel L. and Ellen K. Pikitch: First Oregon Record for the Cowcod, Sebastes levis, 192.
Euliss, Ned H. Jr., and Gail Grodhaus: Management of Midges and Other Invertebrates for Waterfowl Wintering
in California, 238-243.
Fairbrother, A.: see Crawford, Cole, Kilbride, and Fairbrother, 245-247.
Foott, J.S.: see Kent, Duhamel, Foott, and Hedrick, 99-105.
Fritzsche, Ronald A.: see Ward and Fritzsche, 175-187.
Gilbert, Barrie K.: see Hastings and Gilbert, 188-191.
Gotshall, Daniel W.: The use of Baited Stations by Divers to Obtain Fish Relative Abundance Data, 214-229.
Grodhaus, Call: see Euliss and Grodhaus, 238-243.
Hartmann, A. Rucker: Movement of Scorpionfishes (Scorpaenidae: Sebastes and Scorpaena) in the Southern Cal-
ifornia Bight, 68-79.
Hastings, Bruce C, and Barrie K. Gilbert: Extent of Human-Bear Interactions in the Backcountry of Yosemite Na-
tional Park, 188-191.
Hedrick, R.P.: see Kent, Duhamel, Foott, and Hedrick, 99-105.
Hemmer, M.J.: see Russell, Middaugh, and Hemmer, 169-174.
Holl, Stephan A.: see Perry, Dole, and Holl, 156-162.
Ivey, Gary L.: Winter Foods of American Coots in the Northern San Joaquin Valley, California, 45-48.
Kent, M.L., G.E. Duhamel, J.S. Foott, and R.P. Hedrick: Chronic Branchitis (Hamburger Gill Disease) of Channel
Catfish in California and its Possible Mysosporean Etiology, 99-105.
Kilbride, KM.: see Crawford, Cole, Kilbride, and Fairbrother, 245-247.
Knutson, Arthur C. Jr.: Comparitive Catches of Ocean Sport-Caught Salmon using Barbed and Barbless Hooks and
Estimated 1984 San Francisco Bay Area Charterboat Shaker Catch, 106-116.
Lea, Robert N., and Richard H. Rosenblatt: Occurrence of the Family Notacanthidae (Pisces) from Marine Waters
of California, 51-53.
Lea, Robert N.: On the Second Record of Barbourisia rufa, the Velvet Whalefish, from California, 124.
Marshall, William H.: see Miller and Marshall, 37^t4.
Middaugh, D.P.: see Russell, Middaugh, and Hemmer, 169-174.
Miller, Kathy Ann, and William H. Marshall: Food Habits of Large Monkeyface Prickleback, Cebidichthys violaceus,
37^M.
Natanson, Lisa J.: see Ebert, Compagno, and Natanson, 117-123.
Perry, William M., Jim W. Dole, and Stephen A. Holl: Analysis of the Diets of Mountain Sheep from the San Gabriel
Mountains, California, 156-162.
Pikitch, Ellen K.: see Erickson and Pikitch, 192.
Randall, John E.: Refutation of Lengths of 11.3, 9.0, and 6.4 m Attributed to the White Shark, Carcharodon
carcharias, 1635-168.
Reilly, Paul N.: Population Studies of Rock Crabs, Cancer anfennarius, Yellow Crabs, C. anthonyi, and Kellet's
Whelks, kelletia kelletii. in the Vicinity of a Proposed Liquified Natural Gas Terminal at Little Cojo Bay, Santa
Barbara County, California, 88-98.
Rienecker, Warren C: Population Trends, Distribution, and Survival of Canada Geese in California and Western Ne-
vada, 1949-79, 21-36.
INDEX TO VOLUME 73 251
Rienecker, Warren C: Migration and Distribution of Northern Pintails Banded in California, 139-155.
Rienecker, Warren C: Survival and Recovery Rate Estimates of Northern Pintails Banded in California, 1948-79,
230-237.
Rosenblatt, Richard H.: see Lea and Rosenblatt, 51-53.
Russell, G. A., D. P. Middaugh, M. J. Hemmer.: Reproductive Rhythmicity of the Atherinid Fish, Colpichthys regis,
from Estero del Soldado, Sonora, Mexico, 169-174.
Seigel, Jeffrey A.: Record of the Twinpored Eel, Xenomystax atrahus (Anguilliformes: Congridae) from California
Waters, 57-59.
Spratt, Jerome D.: Variation in the Growth Rate of Pacific Herring from San Francisco Bay, California, 132-138.
Ward, David L., and Ronald A. Fritzsche: Comparison of Meristic and Morphometric Characters among and Within
Subspecies of the Sacramento Sucker [Catostomus occidentalis) Ayres, 175-187 .
Warner, Ronald W.: Age and Growth of Male Dungeness Crabs, Cancer magister, in Northern California, 4-20.
Webber, Marc A., and Sarah G. Allen: Resightings of Two Rehabilitated and Released Harbor Seals in California,
60-61.
Wicksten, Mary K.; Range Extensions of Offshore Decapod Crustaceans from California and Western Mexico,
54-56.
SUBJECT
Abalone, red: Movement and dispersion of, in northern California, 196-213
Baited stations: Use of, by divers to obtain fish relative abundance data, 214-229
Bear-human interactions: In the backcountry of Yosemite National Park, 188-191
Branchitis, chronic: Of channel catfish, 99-105
Coot, American: Winter foods of, 45-48
Cowcod: First Oregon record, 192
Crab, Dungeness: Age and growth of males, 4-20
Crab, rock: Population studies of, 88-98
Crab, yellow: Population studies of, 88-98
Crustaceans, decapod: Range extensions of, 54-56
Eel, twinpored: Record from California waters, 57-59
El Nirio: 50, 113, 134
Food habits: Of large monkeyface prickleback, 37^M
Foods, winter: Of American coots, 45-48
Geese, Canada: Population trends, distribution, and survival, 21-36
Gill disease, hamburger: Of channel catfish, 99-105
Hawk, Cooper's: Habitat and productivity of, 80-87
Herring, Pacific: Variation in the growth rate, 132-138
Kyphosidae: Two species seen in King Harbor, Redondo Beach, 49-50
Midges: Management of, for waterfowl wintering in California, 238-243
Notacanthidae: Occurrence from marine waters of California, 51-53
Pintail, northern: Migration and distribution of, banded in California, 139-155
Pintail, northern: Survival and recovery rate estimates of, banded in California, 230-237
Population studies: Of rock crabs, yellow crabs, and Kellet's whelks, 88-98
Population trends: Of Canada geese in California and western Nevada, 21-36
Prickleback, monkeyface: Food habits of, 37—44
Quail, California: Atypical plumage, 245-247
Range extension: Of offshore decapod crustaceans, 54-56
Reproductive rhythmicity: Of the atherinid fish, Colpichthys regis, 169-174
Salmon, ocean sport-caught: Comparative catches of, using barbed and barbless hooks, 106-116
Scorpion fishes: Movement in the Southern California Bight, 68-79
Seal, harbor: Resightings of two rehabilitated and released, 60-61
252
CALIFORNIA FISH AND CAME
Shark, Pacific sleeper: Biological notes on, 1 1 7-1 23
Shark: Yellowtail chafing on, 244
Shark, white: Refutation of lengths of 11.3, 9.0, and 6.4 m, 163-168
Sheep, mountain: Analysis of the diets of, 1 56-1 62
Sucker, Sacrannento: Comparison of meristic and morphometric characters among and within subspecies, 175-187
Waterfowl: Management of midges and other invertebrates for, 238-243
Whalefish, velvet: On the second record of, from California, 1 24
Whelk, Kellet's: Population studies of, 88-98
Yellowtail: Chafing on a shark, 244
SCIENTIFIC NAMES
Accipiter cooperii: 80-87
Adenostema fasciculatum: 160
Aglaophenia sp..- 41
Ahnfeltia plicata: 39
Anas acuta acuta: 238
Anas acuta: 46, 139-155
Anas americana: 26, 233
Anas crecca caroiinensis: 238
Anas platyrhynchos: 35 230-237, 238
Anisotremus davidsonii: 49
Anoplopoma fimbria: 124
Anser americana: 147
Anser caerulescens caerulescens: 26, 147
Anser crecca caroiinensis: 147
Apedilum subcinctum: 239
Aplidium sp..- 41
Archidistoma sp..- 41
Archidistoma ritteri: 41
Arctostaphylos sp..- 160
Aster alexis: 47
Atherinops: 169, 170
Athya collaris: 233
Avicennia germinans: 170
Axiidae: 54
Barbourisia rufa: 124
Barbourisiidae: 124
Bothidae: 220
Botryoglossum farlowianum: 38, 41
Branta canadensis maxima: 34
Branta canadensis minima: 35
Branta canadensis moffitti: 21-36
Branta canadensis occidentalis: 29
Callianassa goniophthalma: 55
Callianassidae: 55
Calliarthron tuberculosum: 200
Callipepla californica: 245
Callophyllis crenulata: 39
Callophyllis pinnata: 38
Callophyllis violacea: 38
Calocaris quinqueseriatus: 54
Cancer antennarius: 88-98
Cancer anthonyi: 88-98
Cancer magister: 4-20
Cancer productus: 96
Carangidae: 244
Carassius auratus: 241
Carcharodon carcharias: 163-168
Castilleja affinis: 159
Catostomus occidentalis: 175-187
Catostomus occidentalis humboldtianus: 175-187
Catostomus occidentalis lacusanserinus: 175-187
Catostomus occidentalis mnlotiltus: 175-187
Catostomus occidentalis occidentalis: 175-187
Ceanothus crassifolius: 160
Ceanothus leucodermis: 159
Cebidichthys violaceus: 37-44
Centroceros clavulatum: 39
Cephaloscyllium ventriosum: 220
Ceramium sp..- 39
Cercocarpus betuloides: 158, 159
Cercocarpus ledifolius: 159, 161
Chione: 170
Chironomidae: 46, 47, 239
Chironomus decorus: 239
Chironomus stigmaterus: 239
Ciccadellidae: 47
Cladophora columbiana: 38
Clupea harengus: 69
Clupea harengus pallasi: 132-138
Colinus virginianus: 245
Colpichthys regis: 169-174
Congridae: 57, 59
Corallina vancouveriensis: 39
Corixidae: 47
Costia: 102
Cottidae: 220
Coturnix coturnix japonica: 245
Cricotopus sp..- 239
Cryptopleura corallinara: 39
Cryptopleura lobulifera: 38
Cryptopleura violacea: 38, 41
Cryptosiphonia woodii: 38
Cyprinus carpio: 241
Decapoda: 54-56
INDEX TO VOLUME 73
253
Dilsea californica: 39
DisWplia occidentalis: 41
Distapiia sp.; 41
Distich/is spicata: 47
Oodecaceria cone ha rum: 199
Dvtiscidae: 47
Echinochloa crusgalli: 47, 239
Echinodorus cordifolius: 239
Embiotoca jacksoni: 217
Embiotocidae: 220
Encelia californica: 159
Endocladia murlcata: 39
Engraulis mordax: 107, 136
Enteromorpha intestinalis: 39
Enteromorpha linza: 39
Ephydridae: 47
Eriodictyon crassifolium: 160
Eriogonum fasciculatum: 158, 159
Eriogonum ovalifolium: 161
Eriogonum umbellatum: 161
Eriophyllum confertiflorum: 159
Erythrophylium delesserioides: 39
Farlowia mollis: 39
Fulica americana: 45—48
Calathea californiensis: 55
Calatheidae: 55
Cambusia affinis: 241
Garrya veatchii: 158, 159
Gasterosteus aculeatus: 241
Gastroclonium coulteri: 38, 41
Gelidium coulteri: 39
Celidium purpurascens: 39
Gigartina canaliculata: 39
Gigartina corymbifera: 39
Gigartina volans: 39
Gracilaria sjoestedtii: 39
Gymnogongrus linearis: 39
Ha Hot is corrugata: 211, 212
Haliotis iris: 210
Haliotis midae: 210
Haliotis ruber: 211
Haliotis rufescens: 196-213
Haliotis sorenseni: 211
Haliotis tuberculata: 211
Halymenia californica: 39
Heleochloa shoenoides: 47
Henneguya: 99-105
Henneguya exilis: 100, 103
Heptacarpus sp.; 41
Hermosilla azurea: 49
Heteromeles arbutifolia: 159
Hexagrammidae: 220
Hippolytidae: 54
Hymenena flabelligera: 38, 41
Hypsurus caryi: 217
Ictalurus nebulas us: 100
Ictalurus punctatus: 99-105
Idotea stenops: 41
Iridaea cordata: 38, 39, 41, 42
Iridaea flaccida: 39
Istiophoridae 244
Juncus sp..- 47
Kelletia kelletii: 88-98
Kyphosidae; 49-50
Kyphosus analogus: 49-50
Labridae; 220
Leptodactylon californicum: 159
Leuresthes sardina: 169, 174
Leuresthes tenuis: 169, 174
Lithodidae: 55
Loligo opalescens: 4
Lymaneidae: 47
Macdonaldia challengeri: 51
Macrocystis pyrifera: 89, 211, 215
Mastocarpus jardinii: 39
Mastocarpus papillatus: 38, 41
Menidia menidia: 169, 172
Mentzelia laevicaulis: 159
Merluccius productus: 57
Microcladia borealis: 38
Microcladia coulteri: 39
Microstomus pacificus: 51, 124
Mobulidae: 244
Morone saxatilis: 1 15
Munida hispida: 55
Muraenesocidae; 59
Najas guadalupensis: 239
Neoclinus spp..- 220
Neoclinus uninotatus: 217
Neoptilota californica: 39
Neoptilota densa: 39
Neoptilota hypnoides: 39
Nettastomidae: 59
Nienburgia andersonii: 39
Notacanthidae; 51-53
Notacanthus chemnitzii: 51-53
Octopoteuthis deletron: 121
Odonthalia floccosa: 39
Oncorhynchus kisutch: 106-116
Oncorhynchus tshawytseha: 106-116, 122
Ophiodon elongatus: 69, 72
Ovis canadensis nelsoni: 156-162
Oxyjulis californica: 227
Pandalidae: 54
Pandalopsis ampla: 54
Paralabrax clathratus: 217, 220
Paralithodes rathbuni: 55
Parapaguridae: 55
254
CALIFORNIA FISH AND CAME
Parapagurus haigae: 55
Paspalum distlchum: 47
Phasianidae: 245
Phasianus colchicus: 245
Phocavitulina: 122
Phoca vltulina richardsi: 60-61
Phyllospadix torreyi: 38
Pikea californica: 39
Pinus sabinlana: 82
Pithophora sp: 46, 47
Platanus racemosa: 82
Pleuronectidae: 220
Plocomlum cartilagineum: 38
Pogonophorella californica: 39
Polyacanthonotus challengeri: 51
Polycheria sp.; 41
Polyneura latissima: 38
Polysiphonia sp.: 39
Populus fremontii: 82
Porichthys spp.; 220
Porphyra lanceolata: 39
Porphyra perforata: 38, 40, 41
Potamogeton pectinatus: 239
Prionace glauca: 244
Prionitis lanceolata: 39
Prionitis lyallii: 38
Prunus ilicifolia: 158, 159
Psychrolutes phrictus: 51
Pterosiphonia dendroidea: 39
Pterygophora californica: 95, 215
Plilota filicina: 38
Pycnopodia helianthoides: 199
Quercus agrifolia: 82
Quercus duglasii: 82
Quercus lobata: 82
Quercus wislizenii: 82
Rathbunella hypoplecta: 217, 220
Rhamnus californica: 159
Rhamnus crocea: 159
Rhizophora mangle: 170
Rhodoglossum affine: 38
Rhodoglossum roseum: 38
Rhodomela larix: 39
Rhodymenia californica: 39
Rhus ovata: 159
Rumex crispus: 47
Ruppla maritima: 239
Salvia apiana: 158, 159
Sarcodiotheca gaudichaudii: 39
Schizymenia pacifica: 39
Scirpus acutus: 239
Sclrpus maritimus: 47
Scirpus spp.; 47
Scorpaena guttata: 69, 72
Scorpaena: 68-79
Scorpaenichthys marmoratus: 69, 72
Scorpaenidae: 68-79
Sebastes: 68-79
Sebastes spp.; 4
Sebastes atrovirens: 72
Sebastes auriculatus: 72
Sebastes carnatus: 72
Sebastes caurinus: 72
Sebastes chlorostictus: 72
Sebastes chrysomelas: 72
Sebastes constellatus: 72
Sebastes dalli: 72
Sebastes entomelas: 69, 72
Sebastes flavidus: 72
Sebastes goodei: 69
Sebastes hopkinsi: 72
Sebastes jordani: 69
Sebastes levis: 69, 192
Sebastes melanops: 72
Sebastes miniatus: 69, 72
Sebastes mystinus: 72, 137
Sebastes nebulosus: 72
Sebastes ovalis: 72
Sebastes paucispinis: 69, 72
Sebastes pinniger: 72
Sebastes rastrelliger: 72
Sebastes rosaceous: 72
Sebastes ruberrimus: 72
Sebastes rubrivinctus: 72
Sebastes serranoides: 72
Sebastes serriceps: 72
Sebastes umbrosus: 72
Sebastolobus alascanus: 121, 124
Sectator ocyurus: 49-50
Seriola lalandi: 244
Somniosus microcephalus: 1 18
Somniosus pacificus: 1 17- 123
Spirontocaris sica: 54
Squalidae: 117
Squatina californica: 220
Stenogramme interrupla: 39
Strongylocentrotus: 215
Synodus lucioceps: 220
Thaleichthys pacificus: 61
Thunnus alalunga: 121
INDEX TO VOLUME 73 255
Trichodina: 102 Ursus americanus: 188-191
Trichophyra: 102 Ursus arctos: 188
Tri folium sp.; 47 Viola purpurea: 159
Typha latifolia: 239 Zenomystax atrarius: 57-59
Ulva lobala: 38, 40, 41, 43 Yucca whipplei: 159
Ulva taeniata: 38 Zannichellia palustris: 239
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