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18

19

20

21

22



TABLE 1. Plankton Tow Station Data.

Bottom
depth

(m)

12
12

9

6
11
20
16
12

7
11
16
12

7
19
19
11
11
11

7
21
23
36



Inclusive trawl
dates

Feb. 1973-May 1974
Feb. 1973-Sept. 1974
Feb. 1973-Sept. 1974
Feb. 1973-Sept. 1974
Feb. 1973-May 1974
Feb. 1973-May 1974
Feb. 1973-May 1974
Feb. 1973-May 1974
Feb. 1973-May 1974
Feb. 1973-May 1974
Feb. 1973-May 1974
Feb. 1973-May 1974
Feb. 1973-May 1974
Feb. 1973-May 1974
Feb. 1973-May 1974
Apr. 1973-Aug. 1974
Nov. 1973-Aug. 1974
Jan. 1974-Aug. 1974
Mar. 1974-Aug. 1974
Mar. 1974-Aug. 1974
July 1974-Sept. 1974
July 1974-Sept. 1974



5 min. A calibrated rotometer, located inside the mouth of the net, was used to
determine the volume of water filtered during 40 of the standardized tows. The
mean volume of water strained was calculated to be 50.5 m 3 per tow (range,
43-57; standard deviation, 5.1; standard error of mean, 0.8). All samples were
taken during daylight hours, with the exception of one complete series. Samples
were preserved in 5% formalin solution and later sorted with the aid of a
dissecting microscope. Large samples were aliquoted with a Folsom Splitter.

RESULTS

Standard lengths, weights, and gonosomatic indices were computed for 359
female and 280 male £ mordax sampled during 1973 and 1974. The female-male
ratio of 1.28:1 is close to the estimate by Clark and Phillips (1952) for the
1947-1951 live-bait fishery (1.2:1), and the statewide estimate of 1.27:1 given
by Miller etal. (1955). However, Klingbeil (1977) suggests spatial and temporal
segregation of the sexes in anchovies which may lead to sampling bias when
specimens are obtained from commercial or bait fishermen's nets.

The highest individual GSI values (male, 524; female, 751) and the highest
mean GSI values occurred in February and March of both years and gradually
declined to the lowest values in September (Figure 2). A very rapid increase in
the GSI values occurred between January and February. Although fish with
seemingly ripe gonads (large, yolked eggs plainly visible in ovaries; testes en-
larged and cream-colored) were found throughout the year, the proportion of
fish with ripe gonads was highest in February and lowest in late summer. Female
anchovies as small as 81 mm (3.2 inches) sl were found with well developed
ova, and males as small as 78 mm (3.1 inches) sl were found with enlarged
testes. Anchovies within this size range are probably 1 year old (Collins 1969).



178



CALIFORNIA FISH AND CAME



6OO-1



500'



400-



x

LU

o

Z



u



300-



<

o

m
O

Z

o

O 200'



150-

100-
75-
50
25-



RANGE



FEMALE

T



.MALE



• MEAN




M



A



— r-
M



N



MONTH

FIGURE 2. Conosomatic indices for Engraulis mordax sampled from the Los Angeles-Long Beach
Harbor live-bait fishery (original data in Brewer 19756 ).



Although spawning was recorded in every month, eggs and larvae occurred
most frequently and were most numerous in February and March when 61.5%
of all eggs and 68.6% of all larvae were taken (Tables 2 and 3; Figure 3). Tows
made during the period between February and May captured 80% of all eggs
and 81.4% of all larvae. This same period included 50.5% of all trawls taken.
The greatest number of anchovy eggs taken during one trawl was 1720 (May
1974, station 20), equivalent to 34,059 eggs per 1000 m 3 of water filtered. The
greatest number of larvae taken during one trawl was 812 (March 1974, station
14), or 16,079 larvae per 1000 m 3 of water filtered. During the entire 20-month



REPRODUCTION AND SPAWNING OF NORTHERN ANCHOVY



179



towing period, San Pedro Bay yielded 536 eggs and 128 larvae per 1000 m 3 of
water filtered.

TABLE 2. Eggs and Larvae of Engraulis mordax taken in Los Angeles-Long Beach Harbor and
San Pedro Bay, California, February 1973-September 1974, Summarized by Month.



No. of

Month tows

January 36

February 51

March 68

April 72

May 92

June 30

July 37

August 37

September 38

October 32

November 34

December 34

TOTALS 561



No. of


No. of




Mean No.


occurrences


specimens


per ,


1000 m 3


Eggs


Larvae


Eggs


Larvae


Eggs


Larvae


2


2


4


2


2


1


31


24


4054


1098


1574


426


60


60


5189


1471


1511


428


42


37


863


377


237


104


45


35


3214


258


692


56


1





1





1





13


5


326


15


174


8


11


6


980


151


524


81


18


12


403


22


210


11


12


1


20


1


12


1


6


23


129


155


75


90


5


16


7


68


4


40



246



221



15190



3618



TABLE 3. Eggs and Larvae of Engraulis mordax taken in Los Angeles-Long Beach Harbor and
San Pedro Bay, California, February 1973-September 1974, Summarized by Station.



Station

1

2

3

4

5

6

7

8

9
10
11
12
13
14
15
16
17
18
19
20
21
22

TOTALS



No. of
tows

32
35
35
35
32
31
32
32
31
32
32
32
32
30
35
29
15
11
6
6
3
_3

561



No. of
occurrences



Eggs

10

18

14

14

15

15

10

10

10

15

13

19

14

20

19

5

5

4

5

5

3

_3

246



Larvae

12

16

14

10

9

13

14

10

14

14

14

19

14

20

19

5

8

4

4

5

2

_2

221



No. of

specimens

Eggs Larvae

128 79

466 209

434 229

361 122

267 105

370 175

272 62

736 70

247 105

192 127

417 98

1246 293

846 153

2927 930

3101 614

8 11

42 22

7 10

94 23

2195 130

348 29

486 22



Mean No.
per 1000 m 3
Eggs
79
264



246

204

165

236

168

455

153

119

258

771

523

1932

1754

5

55

13

310

7244

2297

3208



Larvae

49

118

130

69

65

112

38

43

65

79

61

181

95

614

347

8

29

18

76

429

191

145



15190



3618



Some of the increase in the number of eggs spawned during the summer
months resulted from increased effort during July, August, and September 1974,
with the initiation of off-shore stations 21 and 22 (Figure 1 ). Some 49% of all
eggs collected during the summer months of 1973 and 1974 were taken from a



180
lOOOn

500'



4

ioo-r



90-



< 80-




uu

°- 70-

LU

< 60-

>
>

O

I 50'
U

z
<

4<H



30-



20-



10-



CALIFORNIA FISH AND CAME



/T



RANGE



k MEAN




MONTH

FIGURE 3. Engraulis mordax larval abundance in Los Angeles-Long Beach Harbor and San Pedro
Bay based on number of larvae captured per tow per month.

total of only six trawls at stations 21 and 22. Mature, reproducing fish may move
to deeper waters offshore to escape warm waters nearshore.



DISCUSSION

Data on seasonal occurrence and distribution of £ mordax larvae from a large
area influenced by the California Current is available. According to Ahlstrom
(1967), the northern and southern extent of anchovy spawning fluctuates from
year to year in response to varying oceanic conditions and no average distribu-
tion of abundance of larvae could be found. However, the spawning distribution



REPRODUCTION AND SPAWNING OF NORTHERN ANCHOVY 181

is closely tied to water temperatures. During California's "warm water years"
in 1957, 1959, and 1964 (Jones 1971), Ahlstrom (1966) recorded a northward
shift in anchovy spawning off California. Similarly, during 1956, an abnormally
cold year for waters off California, an unusually high number of anchovy larvae
were captured off southern Baja California. While anchovy eggs have been taken
in water temperatures (at 10-m depth) ranging from 9.9 to 23.3 C (50 to 74 F),
over 90% were taken in water 13.0 to 17.5 C (55 to 64 F) (Ahlstrom 1956).
Ninety-six percent of all larvae captured during the period 1951 to 1964 were
taken between Point Conception, California, and Magdalena Bay, Baja Califor-
nia, with the highest yields between January and May and the lowest yields
between August and October (Ahlstrom 1967). Water temperatures (at 10-m
depth) between Point Conception and Magdalena Bay from January through
May range between 12 and 18 C (54 and 66 F). Maximum temperatures in the
same area occur in the months of August, September, and October (Lynn 1967).

Within restrictive temperature limits, other subtle environmental factors con-
trol reproduction of £ mordax. In San Pedro Bay, surface water temperatures
are generally between 1 3 and 1 8 C ( 55 to 66 F ) throughout the year ( Soule and
Oguri 1972-1976); nevertheless, most spawning occurs in the late winter and
early spring.

Day length influences gonad maturation in some fishes. Wiebe (1968) found
that spermatogenesis would proceed in Cymatogaster (Embiotocidae) in cold
water (10 C) if day lengths were long; high temperatures (20 C) and short
photoperiods inhibited gonad maturation, while high temperatures and long
photoperiods hastened gonad maturation. Kaya (1973) observed that ". . .
long and increasing photoperiods and elevated temperatures of spring . . ."
induced rapid gametogenesis in Lepomis (Centrarchidae). At first sight, anchovy
reproduction off southern California seems to fall within a similar pattern of
regulation, with intensive spawning occurring when water temperatures and day
lengths are increasing. However, other data discount the importance of
photoperiod or changing water temperatures as stimuli for reproduction.

Richardson (1973) has shown that spawning of £ mordax is delayed off
Oregon until June, July, and August when water temperatures are above 14 C
(57 F). Day lengths are maximum in June and are decreasing in July and August.

Leong (1971) maintained a spawning stock of £ mordax at the La Jolla
Fisheries-Oceanography Center throughout the year, under laboratory condi-
tions, which included a short day length (4 hr) and constant temperatures of
1 5 C ( 59 F ) . Adult anchovy held at temperatures between 1 2 and 1 8 C (54 and
64 F) in the laboratory by Brewer (19756 ) developed and maintained relatively
high GSI values regardless of the photoperiod. GSI values for acclimated labora-
tory fish were higher than those sampled from nature at the same time during
the summer and fall when the laboratory fish were maintained under photoperi-
ods of 8, 12, or 16 hr of light.

Indirect evidence suggests that the seasonal reproductive cycle of the an-
chovy may be imposed by limited availability of food.

Gametogenesis requires the storage of large amounts of lipids and proteins in
the gonads. The caloric requirements of reproductive fish must be substantially
above maintenance levels; these levels may not be available to the anchovy
throughout the year (Leong and O'Connell 1969). Clemens and Reed (1967)
and de Vlaming (1971) have induced gonadal regression in Carassius (Cy-



182 CALIFORNIA FISH AND CAME

prinodontidae) and Gillichthys (Gobiidae), respectively, by food deprivation.
Scott (1961 ) found that insufficient diet caused reduction in fecundity in Salmo
(Salmonidae). On the other hand, well fed northern anchovy in the laboratory
maintain high GSI values throughout the year (Leong 1971; Brewer 19756).

Data on zooplankton densities lend support to these speculations concerning
limited food availability. Surface zooplankton densities from San Pedro Bay were
recorded each month in 1 972 ( Harbors Environmental Projects 1 975 and unpub-
lished data). Assuming that station 15 (Figure 1 ) is representative of San Pedro
Bay waters and the dominant zooplankter Acartia tonsa, which comprise 57.9%
of all zooplankters is representative of the adult anchovy's food (Loukashkin
1970), trends in food availability coincide with the relative abundance of an-
chovy larvae (Table 4). Interestingly, 81.5% of all anchovy larvae were captured
in the first 5 months of the year and 82.1% of all Acartia were captured during
the same monthly period. Quarterly surveys of zooplankton in 1973 and 1974
by the Harbors Environmental Projects show similar trends in zooplankton
abundance.

TABLE 4. Abundance of Acartia tonsa in Relation to Abundance of Anchovy Larvae and

Mean GSI Values in San Pedro Bay.

Percentage Percentage

of Acartia of larvae Mean GSI values

Month captured captured female male

January 46.7 0.0 84 33

February 21.4 30.4 308 167

March 4.4 40.7 210 218

April 1.4 10.4 189 116

May 8.2 7.1 124 54

June 2.3 0.0 127 64

July 2.4 0.4 61 34

August 0.5 4.2 44 29

September 0.9 0.6 42 16

October 2.3 0.0 86 80

November 7.6 4.3 43 18

December 1.5 1.9 62 47

£ mordax may have the potential to breed all year, but is constrained to a
seasonal reproductive cycle by high and low temperatures (e.g., above 18 C and
below 13 C). Within this temperature-regulated cycle, the maximum expression
of the anchovy's fecundity may be limited by nutritional requirements that
outweigh the available environmental production.

This view holds that the number of spawnings per year and the number of eggs
spawned are variable. Within the geographical range of anchovy spawning,
multiple spawnings are possible only as long as temperatures are not restrictive,
and the number of eggs spawned is maximal only if the fish's diet is not limiting.

Data on larval abundance in San Pedro Bay (Table 4) where temperatures
generally are not limiting suggest a trimodal abundance of larvae (i.e., three
unequal batches of eggs spawned annually) . Low GSI values and repeat spawn-
ings during the second half of the year correspond to a period of low zooplank-
ton densities. Low zooplankton availability, combined with increased water
temperatures (and hence, increased metabolic requirements) during August,
September, and October, may not allow the increase in gonad weights necessary



REPRODUCTION AND SPAWNING OF NORTHERN ANCHOVY 183

for heavy spawning. The resorption of ovarian eggs noted by MacGregor (1968)
may be a manifestation of the anchovy's inability to obtain an adequate diet
during part of the year. Careful temperature and photoperiod controlled labora-
tory experiments on anchovy fed various rations could resolve these supposi-
tions.

ACKNOWLEDGMENTS

Basil G. Nafpaktitis, Gerald J. Bakus, John E. Fitch, Bernard W. Pipkin, and
Camm C. Swift criticized an earlier version of this manuscript. Without the
interest and cooperation of William Verna, live-bait dealer, this study would not
have been possible.

The work was funded, in part, by NOAA-Sea Grant (No. 04-3-158-36 and
04-3-158-45); the Army Corps of Engineers; the Resources Agency, State of
California; the Pacific Lighting Service Company; and a Grant-in-Aid of Research
from the Society of Sigma Xi.

REFERENCES

Ahlstrom, E.H. 1956. Eggs and larvae of anchovy, jack mackerel, and Pacific mackerel. Calif. Mar. Res. Comm.,
Calif. Coop. Oceanic Fish Invest. Prog. Rept., 1 April 1955-June 1956: 33—42.

1959. Vertical distribution of pelagic fish eggs and larvae off California and Baja California. U.S. Fish and

Wildl. Serv., Fish Bull., 60 (161 ):107-146.

1966. Distribution and abundance of sardine and anchovy larvae in the California Current region off

California and Baja California, 1954-64: A summary. U.S. Fish and Wildl. Serv., Spec. Sci. Rept., Fisheries,
534:1-71.

1967. Co-occurrences of sardine and anchovy larvae in the California Current region off California and

Baja California. Calif. Mar. Res. Comm., Calif. Coop. Oceanic Fish. Invest. Rept., 11:117-135.

Bolin, R.L. 1936. Embryonic and early larval stages of the California anchovy, Engraulis mordax Girard. Calif. Fish
Came 22(41:314-321.

Brewer, CD. 1975a. The biology and fishery of the northern anchovy in San Pedro Bay: Potential impact of
dredging and landfill, p. 23-43. In D. Soule and M. Oguri (eds.) Marine Studies of San Pedro Bay, California,
pt. VIII. Allan Hancock Found. & Sea Grant Progr., Univ. So. Calif. USC-SG-1-74.

1975&. The biology of the northern anchovy (Engaulis mordax), in relation to temperature. Ph.D.

Dissertation, Univ. So. Calif., 205 p.

1976. Thermal tolerance and resistance of the northern anchovy (Engraulis mordax). U.S., Fish. Bull.,



74(21:433-445.

California Department of Fish and Game. 1971. Northern anchovy, p 48-51. In H.W. Frey ed. California's living
marine resources and their utilization. The Resources Agency, State of California.

Clark, F.N., andJ.B. Phillips. 1952. The northern anchovy [Engraulis mordax mordax) in the California fishery. Calif.
Fish Game, 38(2):189-207.

Clemens, H.P., and C.R. Reed. 1967. Testicular characteristics of goldfish (Carasslus auratus) in nature and under
diet limitations. J. Morphol., 122:131-138.

Collins, R.A. 1969. Size and age composition of northern anchovies (Engraulis mordax) in the California anchovy
reduction fishery for the 1965-66, 1966-67, and 1967-68 seasons, p 56-63. In\. Messersmith ed., The northern
anchovy and its fishery, Calif. Dept. Fish and Game, Fish Bull., (147): 1-102.

de Vlaming, V.L. 1971. The effects of food deprivation and salinity changes on reproductive function in the
estuarine gobiid fish, Gillichthys mirabilis. Biol. Bull. (Woods Hole), 141:458-471.

Allan Hancock Foundation. 1975. Environmental investigations and analyses for Los Angeles-Long Beach Harbors.
A report to the U.S. Army Corps of Engineers, Contract No. DAWCD 9-73-0112.

Haydock, I. 1971. Gonad maturation and hormone induced spawning of the gulf croaker, Bairdiella Icistia. U.S.,
Fish. Bull., 69(11:157-180.

Jones, J.H. 1971. General circulation and water characteristics in the southern California bight. So. Calif. Coastal

Water Res. Prog., El Segundo, Calif., 37 p.
Kaya, CM. 1973. Effects of temperature and photoperiod on seasonal regression of gonads of green sunfish,

Lepomis cyanellus. Copeia, 1973(2):369-373.

Khanna, S.S., and M.C Pant. 1967. Seasonal changes in the ovary of asisorid catfish, Clyptosternum pectinopterum.
Copeia, 1967(1 ):83-88.



184 CALIFORNIA FISH AND CAME

Klingbeil, R.A. 1 977. Sex ratios of the northern anchovy, Engraulis mordax, off southern California. Calif. Fish Came
64(31:200-209

Leong, R.J.H. 1971. Induced spawning of the northern anchovy, Engraulis mordax. U.S., Fish. Bull., 69(21:357-360.

Leong, RJ.H., andC.P. O'Connell. 1969. A laboratory study of particulate and filter feeding of the northern anchovy
{Engraulis mordax.) Can., Fish. Res. Bd., )., 26(3):557-582.

Loukashkin, A.S. 1970. On the diet and feeding behavior of the northern anchovy, Engraulis mordax Gnaxd. Calif.
Acad. Sci., Proc, Fourth Ser, 37(131:419-458.

Lynn, R.J. 1967. Seasonal variation of temperature and salinity at 10 meters in the California Current. Calif. Mar.

Res. Comm., Calif. Coop. Oceanic Fish, Invest. Rept., 11:157-186.
MacCregor, ).S. 1968. Fecundity of the northern anchovy, Engraulis mordax Guard. Calif. Fish Came, 54(41:281-

288.

Mclnerney, ).E., and DO. Evans. 1970. Action spectrum of the photoperiod mechanism controlling sexual matura-
tion in the three-spine stickleback, Casterosteus aculeatus. Can., Fish. Res. Bd., J., 27(4):749-763.

Mais, K.F. 1974. Pelagic fish surveys in the California Current. Calif. Dept. Fish and Came, Fish Bull., ( 1 62) :1— 79.

Miller, D.J., A.E. Daugherty, F.E. Felin, and J. MacCregor. 1955. Age and length composition of the northern
anchovy catch off the coast of California in 1952-53 and 1953-54, p. 36-66. In Age and length determination
of the northern anchovy. Calif. Dept. Fish and Came, Fish Bull., (1011:1-66.

Moser, H.J. 1967. Reproduction and development of Sebastodes paucispinis and comparison with other rockfishes
off southern California. Copeia, 1967(41:773-797.

Richardson, S.L. 1973. Abundance and distribution of larval fishes in waters off Oregon, May-October 1969, with
special emphasis on the northern anchovy, Engraulis mordax. U.S., Fish. Bull., 71 (31:697-711.

Scott, D.P. 1961. Effect of food quantity on fecundity of rainbow trout, Salmo gairdneri. Can., Fish. Res. Bd., J.,
19(41:715-731.

Smith, P.E. 1972. The increase in spawning biomass of northern anchovy, Engraulis mordax. U.S., Fish. Bull.,
70(31:849-874.

Soule, D.F., and M.Oguri (eds.l. 1972-1976. Marine Studies of San Pedro Bay, California, pts. 1-12. Allan Hancock
Found. & Sea Grant Progr., Univ. So. Calif.

Wiebe, J. P. 1968. The reproductive cycle of the viviparous seaperch, Cymatogaster aggregata Gibbons. Can. J.
Zool., 46(61:1221-1234.

Wood, R., and A.R. Strachan. 1970. A description of the northern anchovy live-bait fishery, and the age and length
composition of the catch during the seasons 1957-58 through 1964-65. Calif. Fish Game, 56(11:49-59.



LARCEMOUTH BASS HOOKING MORTALITY 185

Calif. Fish and Came 64 ( 3 ) : 1 85-1 88. 1 978

HOOKING MORTALITY OF JUVENILE

LARGEMOUTH BASS, MICROPTERUS SALMOIDES"

by

RONALD J. PELZMAN

Inland Fisheries Branch

California Department of Fish and Came

Sacramento, California 95819

Mortalities of juvenile largemouth bass due to hooking were studied as part of an
overall assessment of the value of minimum length limit regulations. Significant
mortality occurred only among fish hooked in the esophageal area. Results indicate
that direct mortality due to hooking is not a factor which materially reduces the value
of size limit regulations.

INTRODUCTION

In 1972 the California Fish and Game Commission imposed an experimental
305-mm (12.0-inch) size limit on largemouth bass at Merle Collins Reservoir,
Yuba County. This action was based on high angler harvest rates recorded by
Rawstron and Hashagen (1972) and length frequency of sport catch as deter-
mined from census (Hashagen 1973). Subsequently, size limits have been im-
posed at several reservoirs to control overharvest of largemouth bass. Other
states also use size restrictions for retaining desirable predator-prey structure and
to control bass harvest (Funk 1974).

Hooking mortality will diminish the effectiveness of size restrictions and it is
essential that the magnitude of immediate and delayed mortality of sublegal fish
be determined. The effects of hooking on largemouth bass less than 305 mm
(12.0 inches) were investigated by Rutledge (1974) with inconclusive results.
Therefore, as part of an evaluation of the size limit at Merle Collins Reservoir,
a companion study was initiated in January 1976 to assess hooking mortality of
sublegal largemouth bass.

METHODS AND MATERIALS

The experimental design of this study involved the hooking (by hand) of
hatchery-reared bass held in tanks on the grounds of the Department's experi-
mental management facility (Field Station) in Sacramento. To determine the
degree of pull on embedded hooks under actual fishing conditions, anglers
caught fish of test size on tackle that included a spring scale between the rod
tip and the hook. Anglers caught the fish, played them for 30 seconds, and lifted
them waist high. During the course of this action, an attachment to the spring
scale recorded the maximum reading. The mean of these readings (24 observa-
tions) was doubled and this value, 346 g (12.2 oz), was applied as pressure to
all experimental fish hooked by hand. All hooking experiments (angling and by
hand) were conducted over the period January 15 through March 21, 1976.

A No. 4, medium-shanked "Compac" hook was embedded to the depth of
the barb at designated locations within the fish's mouth cavity. The cavity was
separated into six major areas, each of which was further divided into subareas.

' This work was performed as part of Dingell-)ohnson project California F-18-R, "Experimental Reservoir Manage-
ment", supported by Federal Aid to Fish Restoration funds. Accepted for publication December 1977.



186



CALIFORNIA FISH AND GAME



The number of subareas within each major area was positively correlated with
the size of the area and ranged from two to 12 (Figure 1 ). Individual hookings
were randomized by subarea within each area. Generally, 50 fish per area were
hooked.



300'



270^



240




£ ESOPHAGUS T TONGUE

R ROOF OF MOUTH G GILLS

F FLOOR OF MOUTH L LIP



FIGURE 1. Mouth cavity of largemouth bass divided into six major hooking areas (E,R,F,L,G,T)
and numbered subareas within each area.

Test fish were differentially fin punched by major hooking area and an addi-
tional group (also fin punched) served as a control. All fish were weighed and
measured. Following hook placement and the application of 346 g (12.2 oz) of
pressure in line with the hook shank, the fish was allowed to move about the



LARGEMOUTH BASS HOOKING MORTALITY



187



test tank for 30 seconds. It was then lifted to waist level by a monofilament line
attached to the hook. The hook was usually removed by hand although the use
of needle-nosed pliers was occasionally required. Fish were tossed back into the


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