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OCR for page 39
An Assessment of Atlantic Bluefin Tuna
3
Transatlantic Movement of Atlantic Bluefin Tuna
INTRODUCTION
The degree of fish movement between fishing grounds in the eastern and western Atlantic Ocean and the degree of genetic mixing in the spawning areas in the eastern and western Atlantic Ocean are both important topics relevant to the management of Atlantic bluefin tuna. The review and reanalysis of tagging data constitute the committee's evaluation of the scientific basis for physical movement or mixing of fish between fishing grounds. These tagging data, however, do not provide information on the degree of genetic mixing.
TAG-RECAPTURE DATA
Tag-recapture data provide the strongest evidence available for transatlantic movement of fish. Atlantic bluefin tuna were marked and released in the western Atlantic Ocean from 1954 in varying numbers until 1987 (Parrack, 1990). The total number tagged by the National Marine Fisheries Service (NMFS) since the inception of the program has been over 15,000. Early releases were primarily of large (giant) fish, although more recently releases have included numerous small fish.
West to East
An overview of movement from the west to the east by size class (i.e., from age 0 fish to giant fish), as indicated by tag returns, appears in Suzuki (1990).
OCR for page 40
An Assessment of Atlantic Bluefin Tuna
TABLE 3-1 Synopsis of release and recapture (tagging experiments) of western Atlantic bluefin tuna. Data are from Mather (1980) and Brunenmeister (1980).
Tag locality
# Released
Size
# Recaptured
Bahamas
1,709
Giant fish
17
NW Atlantic Ocean
1,881
Giant fish
81
US Coast*
468
Medium fish
10
NW Atlantic Ocean
>17,700
Small fish
>2,180
Bahamas to eastern Atlantic Ocean
9 [giant fish]
US Coast* to eastern Atlantic Ocean
1 [medium fish]
NW Atlantic to eastern Atlantic Ocean
46 [small fish]
Trans-Atlantic giant fish released in Bahamas
9/17
52.9%
Trans-Atlantic giant fish released in W Atlantic Ocean
9/98
9.2%
Trans-Atlantic medium fish released off US Coast*
1/10
10.0%
Trans-Atlantic small fish released in NW Atlantic Ocean
46/2,180
2.1%**
* From Mather (1980): "mostly" medium fish were released north of 35°N and west of 60°W.
** More than 2,180 fish released; estimate of 2.1% is maximum.
Recaptures through 1978 of large fish released in the Bahamas in the 1960s, and of small fish released along the U.S. Atlantic coast, document unequivocally that transatlantic migration occurs (Table 3-1). Transatlantic migrants include nine giant fish taken off the Norwegian coast, one medium fish (recovered as a giant tuna over 10 years later) taken off the southern coast of Spain, and 46 small fish taken chiefly from the Bay of Biscay. Estimates of the total proportion of fish tagged in the western Atlantic Ocean in eastern Atlantic samples (Table 3-1) range from 9.3% for giant fish and medium fish to 2.1% for small fish. Estimates for giant and medium fish differ slightly from those given in Mather (1980), who found a total of 10 western migrants in 126 eastern Atlantic recaptures (10/126 = 7.9%). Suzuki (1990) estimated an "overall" proportion of western-tagged fish in eastern Atlantic samples of 3.2% as of 1988.
Estimates of west to east transatlantic movement are variable and depend on the size of fish tagged and tagging locality. All but one of the giant fish transatlantic recoveries were tagged near the Bahamas, and five of the nine migrants were recovered in Norwegian waters in the same year. These results indicate that giant bluefin tuna can travel long distances; only one of the remaining 91 giant tuna or medium fish tagged in the western Atlantic Ocean was recovered in the eastern Atlantic Ocean. This fish was tagged near Bermuda and recovered 10 years later. Mather (1980) hypothesized that the transatlantic migration of
OCR for page 41
An Assessment of Atlantic Bluefin Tuna
giant fish was not part of an annual pattern but rather represented irregular migration of varying numbers of fish in different years. This hypothesis deserves consideration because bluefin tuna tend to follow ocean currents (Sella, 1929, cited in Mather, 1980) and because of the proximity of the Bahamian tagging sites to the Gulf Stream. This possible transatlantic "route" for giant fish is strikingly similar to the one for larvae proposed by Murphy (1990). Finally, Mather (1980) also noted that the transatlantic movement of small bluefin tuna was variable and that virtually all transatlantic movements of small fish were from the Mid-Atlantic Bight to the Bay of Biscay. Mather concluded that the west to east transatlantic movement of small fish might be due to unusual stimuli and that the great majority of Atlantic bluefin tuna remain on one side of the Atlantic Ocean or the other. Alternatively, the "nonrandom" movement of small bluefin tuna from the Mid-Atlantic Bight to the Bay of Biscay may reflect the importance of both nursery areas for small bluefin tuna.
East to West
Eastern Atlantic/Mediterranean bluefin tuna have been tagged and released since 1911, although most of the tag and release data are from 1957 to the present (Cort and Liorzou, 1990c). As in the western Atlantic Ocean, most of the early releases were giant tuna, with the number of small fish released and recovered increasing over the past two decades. Numbers of fish released and recaptured from the eastern Atlantic Ocean and Mediterranean Sea from 1911 through 1992 are listed in Brunenmeister (1980), Cort and Liorzou (1990c), and Cort and de la Serna (1993). A synopsis of these listings is shown in Table 3-2. The proportion of eastern-tagged fish among western Atlantic recaptures varies from 0% for giant fish to 4.5% for small fish. Virtually all of the transatlantic migrants recaptured off the U.S. coast were released from the Bay of Biscay, and most were captured between 10 and 20 months after first released. The "total" proportion of eastern fish among western Atlantic recaptures is 4.0% (19/472) after adjustment for the greater number of small fish among recaptures. This estimate is similar to the estimate of 4.4% of Cort and Loirzou (1990c). Estimates of eastern Atlantic fish among Mediterranean recaptures are 11.3% for giant fish and 2.8% for small fish. The proportion of Mediterranean fish among eastern Atlantic recaptures is 2.8% for small fish.
The absence of documented east to west transatlantic movement of large fish is striking and merits further investigation. Considering only small fish, the proportion of transatlantic immigrants in western and eastern Atlantic samples differs: estimates made here suggest an overall proportion of small western fish among eastern Atlantic recaptures of 2.1% and a proportion of 4.5% eastern-tagged fish among western recaptures.
OCR for page 42
An Assessment of Atlantic Bluefin Tuna
TABLE 3-2 Synopsis of release and recapture (tagging experiments) of eastern Atlantic/Mediterranean bluefin tuna. Data are from Brunenmeister (1980), Cort and Loirzou (1990c), and Cort and de la Serna (1993).
Tag Locality
# Released
Size
# Recaptured
Eastern Atlantic Ocean
599-604
Giant fish
53
6,144
Small fish
418-420
107-232
Unknown*
3
Mediterranean Sea
3,993
Small fish
70
20-30
Unknown*
0
Eastern Atlantic to western Atlantic Ocean
19 [small fish]
Eastern Atlantic Ocean to Mediterranean Sea
13 [6 giant fish, 7 small fish]
Mediterranean Sea to eastern Atlantic Ocean
11 [small fish]
Trans-Atlantic giant fish released in eastern Atlantic Ocean
0
Trans-Atlantic small fish released in eastern Atlantic Ocean
19/419
4.5%
Mediterranean small fish to eastern Atlantic Ocean
11/70
15.7%
Eastern Atlantic giant fish to Mediterranean Sea
6/53
11.3%
Eastern Atlantic small fish to Mediterranean Sea
13/472
2.8%
* From studies carried out prior to 1940 in Italy and Portugal.
Reanalysis of Tagging Data
Tagging data suggest that physical movement or mixing of Atlantic bluefin tuna in fishing grounds from opposite sides of the Atlantic Ocean is significant. Thus, the committee reanalyzed the tagging data to estimate rates of transatlantic movement between fishing grounds. This analysis was undertaken to provide rigorous estimates of transfer across the Atlantic Ocean, which has not been attempted previously. Because it is not possible to estimate transfer rates without knowledge of the total mortality rates and the nonreporting rates (tags recovered but not reported), these quantities are also estimated.
METHODS
The problem of transfer rates is described by the rates of change in the population sizes, Nw for the west and Ne for the east. Equations for the transfer of fish from the west to the east, applicable to fish tagged with a single tag, are:
(1)
OCR for page 43
An Assessment of Atlantic Bluefin Tuna
(2)
M = annual rate of instantaneous natural mortality plus shedding. 1
Fw = annual rate of instantaneous fishing mortality in the west.
Fe = annual rate of instantaneous fishing mortality in the east.
Tw = annual rate of instantaneous transfer from west to east.
Integration of this pair of equations provides estimates of the population size at any time t, over the interval for which Fw is constant in the west and Fe is constant in the east. The solutions are given by:
where X = M + Fe and Y = Fw-Fe + Tw
Similarly, the equations for the caches are:
When t is sufficiently large:
When Pw and Pe are the proportions of tags recovered that are reported for the west and east, respectively, then:
(3)
Analogous equations (with the subscripts e and w reversed) are used to estimate the transfer rate of fish from the east to the west. Because the models given by Equations (1) and (2) do not admit roundtrip movement, any such return movement would cause the transfer rates estimated by Equation (3) to be underestimates of movement rates from one side of the ocean to the other. Also, if fishing mortality decreases with age, as it appears to do, the movement rates estimated by assuming a constant fishing mortality will be underestimated.
Tag-recovery information from Corte and de la Serna (1993) (Table 2; which is reproduced below as Table 3-3) and from NMFS archives (file name MRFISH)
1
Shedding refers to tags that have detached from the fish.
OCR for page 44
An Assessment of Atlantic Bluefin Tuna
TABLE 3-3 Spanish tagging data for Atlantic bluefin tuna in the Cantabrian Sea (Bay of Biscay) from 1976 to 1991.
Recoveries
Recoveries in the Eastern Atlantic With Known Years At Liberty
Year
N
0
1
2
3
4
Trans Atlantic
Med. Sea
East Atlantic*
No data
Total
1976
3
-
-
-
-
-
0
-
0
0
-
1977
10
-
-
-
-
-
0
-
0
0
-
1978
170
29
2
4
1
-
1
-
0
5
42
1979
101
1
10
-
-
-
2
-
0
1
14
1980
302
15
2
2
-
-
3
-
2
8
32
1981
293
3
5
-
1
-
0
2
0
2
13
1982
395
5
6
4
2
-
1
3
2
2
25
1983
370
2
1
1
-
-
0
1
0
1
6
1984
513
8
7
1
-
1
0
1
2
6
26
1985
407
12
2
-
2
-
1
-
0
4
21
1986
838
37
8
6
1
1
5
-
0
10
68
1987
-
-
-
-
-
-
0
-
0
0
-
1988
1,151
26
17
5
-
2
1
-
0
7
58
1989
122
2
1
2
2
-
0
-
0
0
7
1990
973
8
22
2
-
-
3
-
0
0
39
1991
15
4
1
-
-
-
0
-
0
0
5
Total
5,663
152
84
29
11
4
17
7
6
46
356
* This column refers to recoveries in the eastern Atlantic without known recovery year.
OCR for page 45
An Assessment of Atlantic Bluefin Tuna
was used to obtain values of catch (C). Some descriptions of qualitative features of the tagging data with respect to movement are given below in the Results section. Shedding (L = M -0.14) (0.14 was taken from the literature as the natural mortality rate; ICCAT, 1993) was estimated from these catch data using the method of Chapman, Fink, and Bennet (1965, equations 5, 6, and 7) to estimate L. This method was used rather than that of Bayliff and Mobrand (1972), because shedding rates do not appear to be constant and appear to be insensitive to type I (initial) shedding. Once the estimate of M (L + 0.14) was available, VPA analysis was done on the catches over time to obtain estimates of F.
Results
Tables 3-4, 3-5, and 3-6 show the number of fish tagged in the western Atlantic Ocean (from MRFISH) and the number of recoveries by year after tagging. Year 0 means fish caught in the same calendar year as tagged; year 1 means fish caught in the calendar year following tagging, etc. Table 3-7 (eastern Atlantic Ocean, ICCAT data) and Table 3-8 (western Atlantic Ocean, NMFS data) show the months in which the fish were tagged and the months in which the fisheries captured the tagged fish. Because most tagged fish were small (less than four years old), inferences about fishing times based on these tag recoveries should apply to small fish.
From 1971 to 1978, there was intensive tagging in the west with both single-and double tagged fish (a small number of double tagged fish actually had more than two tags). Table 3-9 shows the number of recoveries from single tagged fish (SS), the number of recoveries of fish with two tags from double tagged fish (DD), and the number of recoveries of fish with one tag from double tagged fish (DS), by year of recovery. Tagging time varied, so data from the west were sorted into periods of one-quarter year; that is, if a tag was recovered with the same quarter as the release, it was placed in quarter 1. If the recovery was made after more than 91 days but before 182 days, it was placed in quarter 2, and so on. Table 3-10 shows the recovery data for recoveries SS, DD, and DS by year of release and quarter of recovery.
The western data from Table 3-10 were used to estimate shedding rates (Table 3-11). Comparison of SS to DD (Db Db vs. Sing in Table 3-11) shows the most variability, and comparison of DD to DS (Db Db vs. Db Sing) shows the most stability. After examining Table 3-11, it was decided to try L = 0.26 for all ages and also to try L(1) = 0.50, L(2) = 0.40, L(3) = 0.30, and L(4+) = 0.20, corresponding to quarter 1, quarter 2, quarter 3, and all quarters 4 or more. These values are not much different from the value of 0.205 estimated by Baglin et al. (1980).
With the shedding rates mentioned above, and the catch vectors for SS and DD in Table 3-11, VPA analysis was performed showing that the two sets of
OCR for page 46
An Assessment of Atlantic Bluefin Tuna
TABLE 3-4 Atlantic bluefin tuna release and recovery data from the United States tagging program in the western Atlantic Ocean. Tag returns are by year tagged and years out for all tag types.
Year
Tag
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
Total
1954
193
1
-
-
-
-
2
-
-
-
-
-
-
-
-
-
-
-
-
-
3
1955
232
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1956
99
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1957
39
-
-
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1
1958
38
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1959
147
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1
1960
237
-
-
3
1
-
-
-
-
1
-
-
-
-
-
-
-
-
-
-
5
1961
188
2
2
3
2
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
9
1962
128
1
4
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
5
1963
223
11
3
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
15
1964
553
100
32
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
132
1965
1,812
164
63
36
3
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
267
1966
4,128
524
577
51
8
6
-
2
-
-
1
1
-
-
2
-
-
-
-
-
1,172
1967
718
98
60
16
13
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1
188
1968
521
88
19
9
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
116
1969
567
14
81
12
2
2
-
1
1
-
1
-
-
-
-
-
-
-
-
-
114
1970
729
52
119
9
7
3
-
-
-
-
-
-
-
-
-
-
-
-
-
-
190
1971
432
11
59
12
-
-
1
-
-
-
-
-
-
-
-
-
-
-
-
-
83
1972
284
10
56
6
2
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
74
1973
393
40
22
7
2
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
71
1974
1,752
75
126
51
14
7
-
-
-
-
1
1
1
1
-
-
1
-
-
-
278
1975
349
22
27
6
2
-
-
1
-
-
-
-
-
-
-
-
-
-
-
-
58
1976
2,460
185
63
75
8
-
1
-
-
1
-
-
-
-
-
-
-
-
-
-
333
1977
2,115
52
193
81
2
-
-
1
2
1
1
1
2
-
-
-
-
-
-
-
341
1978
1,680
24
110
34
7
7
3
1
1
2
-
1
2
-
-
-
-
-
-
-
192
1979
1,124
6
30
11
1
3
1
1
-
1
-
-
-
-
-
-
-
-
-
-
53
1980
3,074
141
100
6
3
4
2
2
4
-
2
-
-
-
-
-
-
-
-
-
268
1981
1,797
57
2
2
1
1
1
2
-
4
-
-
-
-
-
-
-
-
-
-
68
1982
210
1
1
1
2
-
-
-
-
2
-
-
-
-
-
-
-
-
-
-
5
1983
149
1
2
4
4
-
-
1
-
-
-
-
-
-
-
-
-
-
-
-
12
1984
86
1
2
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
3
1985
130
-
1
2
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
3
1986
51
-
1
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
2
1987
66
1
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
2
1988
96
-
2
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
2
1989
113
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1990
127
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
Note: Dash indicates zero value.
OCR for page 47
An Assessment of Atlantic Bluefin Tuna
OCR for page 48
An Assessment of Atlantic Bluefin Tuna
TABLE 3-5 Atlantic bluefin tuna release and recovery data from the U.S. tagging program in the western Atlantic Ocean. Tag returns are by year tagged and years out for fish tagged with a single tag.
Year
Tag
0
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
Total
1954
192
1
-
-
-
-
2
-
-
-
-
-
-
-
-
-
-
-
-
-
3
1955
231
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1956
99
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1957
39
-
-
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1
1958
38
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1959
147
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1
1960
237
-
-
3
1
-
-
-
-
1
-
-
-
-
-
-
-
-
-
-
5
1961
185
2
2
3
2
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
9
1962
128
1
4
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
5
1963
183
7
3
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
10
1964
544
100
32
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
132
1965
1,751
164
63
36
3
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
267
1966
4,114
523
575
51
8
6
-
2
-
-
1
1
-
-
2
-
-
-
-
-
1,169
1967
718
98
60
16
13
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1
188
1968
448
60
12
6
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
78
1969
547
14
77
12
1
2
-
1
1
-
1
-
-
-
-
-
-
-
-
-
109
1970
724
51
119
9
7
2
-
-
-
-
-
-
-
-
-
-
-
-
-
-
188
1971
115
2
11
3
-
-
1
-
-
-
-
-
-
-
-
-
-
-
-
-
17
1972
130
3
14
6
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
21
1973
105
4
8
1
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
14
1974
466
17
43
14
7
4
-
-
-
-
1
-
-
1
-
-
-
-
-
-
87
1975
239
20
16
4
-
-
-
1
-
-
-
-
-
-
-
-
-
-
-
-
41
1976
1,916
145
52
66
5
-
1
-
-
-
-
-
-
-
-
-
-
-
-
-
269
1977
1,590
39
129
50
1
-
-
-
1
1
1
1
1
-
-
-
-
-
-
-
224
1978
848
20
36
13
4
4
1
1
-
-
-
-
1
-
-
-
-
-
-
-
80
1979
1,110
5
29
10
1
3
1
1
-
-
-
-
-
-
-
-
-
-
-
-
50
1980
3,066
141
100
6
3
4
2
2
2
4
4
2
-
-
-
-
-
-
-
-
268
1981
1,787
55
2
2
1
1
1
1
-
2
-
-
-
-
-
-
-
-
-
-
65
1982
200
1
1
1
2
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
5
1983
146
1
1
4
4
-
-
1
-
-
-
-
-
-
-
-
-
-
-
-
11
1984
84
-
2
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
2
1985
128
-
1
2
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
3
1986
51
-
1
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
2
1987
66
1
1
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
2
1988
96
-
2
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
2
1989
112
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
1990
127
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
-
Note: Dash indicates zero value.
OCR for page 49
An Assessment of Atlantic Bluefin Tuna
OCR for page 68
An Assessment of Atlantic Bluefin Tuna
TABLE 3-20 Atlantic bluefin tuna tagged in the eastern Atlantic Ocean and recovered in the eastern Atlantic Ocean versus time (years) before recapture and a regression analysis of the natural logarithm of catch number against years out (Cort and LaSerna, 1993).
OCR for page 69
An Assessment of Atlantic Bluefin Tuna
TABLE 3-21 Atlantic bluefin tuna tagged in the western Atlantic Ocean from 1960 to 1981 with a single tag and recovered in the western Atlantic Ocean vs. years before recapture and a regression analysis of the natural logarithm of catch number against years out (data in file MRFISH provided by NMFS).
OCR for page 70
An Assessment of Atlantic Bluefin Tuna
TABLE 3-22 Atlantic bluefin tuna. VPA analysis on fish tagged in the eastern Atlantic Ocean (near Spain) from 1976 to 1991. Six different solutions.
Time
Initial pop
Catch
Fish. Mort.
Nat. Mort.
Shed Rate
Time Units
1
553.8
193.00
0.9608
0.1400
0.2600
0.50
2
280.5
107.00
0.6047
0.1400
0.2600
1.00
3
102.7
37.00
0.5605
0.1400
0.2600
1.00
4
39.3
14.00
0.5522
0.1400
0.2600
1.00
5
15.2
5.00
0.5000
0.1400
0.2600
1.00
TOTALS
356.00
2.6978
0.6300
1.1700
4.50
Time
Initial pop
Catch
Fish. Mort.
Nat. Mort.
Shed Rate
Time Units
1
587.3
193.00
0.8922
0.1400
0.2600
0.50
2
307.8
107.00
0.5351
0.1400
0.2600
1.00
3
120.8
37.00
0.4558
0.1400
0.2600
1.00
4
51.3
14.00
0.3954
0.1400
0.2600
1.00
5
23.2
5.00
0.3001
0.1400
0.2600
1.00
TOTALS
356.00
2.1325
0.6300
1.1700
4.50
Time
Initial Pop
Catch
Fish. Mort.
Nat. Mort.
Shed Rate
Time Units
1
753.3
193.00
0.6599
0.1400
0.2600
0.50
2
443.4
107.00
0.3419
0.1400
0.2600
1.00
3
211.2
37.00
0.2370
0.1400
0.2600
1.00
4
111.7
14.00
0.1640
0.1400
0.2600
1.00
5
63.5
5.00
0.0999
0.1400
0.2600
1.00
TOTALS
356.00
1.1729
0.6300
1.1700
4.50
OCR for page 71
An Assessment of Atlantic Bluefin Tuna
Time
Initial Pop
Catch
Fish. Mort.
Nat. Mort.
Shed Rate
Time Units
1
1,000.3
193.00
0.4765
0.1400
0.2600
0.50
2
645.4
107.00
0.2226
0.1400
0.2600
1.00
3
346.3
37.00
0.1381
0.1400
0.2600
1.00
4
202.2
14.00
0.0874
0.1400
0.2600
1.00
5
124.2
5.00
0.0499
0.1400
0.2600
1.00
TOTALS
356.00
0.7363
0.6300
1.1700
4.50
Time
Initial Pop
Catch
Fish. Mort.
Nat. Mort.
Shed Rate
Time Units
1
2,047.9
193.00
0.2193
0.1400
0.2600
0.50
2
1,502.6
107.00
0.0902
0.1400
0.2600
1.00
3
920.4
37.00
0.0499
0.1400
0.2600
1.00
4
586.9
14.00
0.0294
0.1400
0.2600
1.00
5
382.0
5.00
0.0160
0.1400
0.2600
1.00
TOTALS
356.00
0.2952
0.6300
1.1700
4.50
Time
Initial Pop
Catch
Fish. Mort.
Nat. Mort.
Shed Rate
Time Units
1
2,969.6
193.00
0.1485
0.1400
0.2600
0.50
2
2,257.3
107.00
0.0591
0.1400
0.2600
1.00
3
1,426.4
37.00
0.0319
0.1400
0.2600
1.00
4
926.1
14.00
0.0185
0.1400
0.2600
1.00
5
609.4
5.00
0.0099
0.1400
0.2600
1.00
TOTALS
356.00
0.1936
0.6300
1.1700
4.50
OCR for page 72
An Assessment of Atlantic Bluefin Tuna
TABLE 3-23 Atlantic bluefin tuna. VPA analysis on fish tagged in the western Atlantic Ocean (near USA) from 1960 to 1981. Five different solutions.
Time
Initial Pop
Catch
Fish. Mort.
Nat. Mort.
Shed Rate
Time Units
1
19,162.9
1,471.00
0.1768
0.1400
0.2600
0.50
2
14,361.5
1,386.00
0.1241
0.1400
0.2600
1.00
3
8,503.7
309.00
0.0450
0.1400
0.2600
1.00
4
5,449.3
59.00
0.0133
0.1400
0.2600
1.00
5
3,604.6
26.00
0.0087
0.1400
0.2600
1.00
6
2,395.3
7.00
0.0035
0.1400
0.2600
1.00
7
1,600.0
44.00
0.0113
0.1400
0.2600
10.00
TOTALS
3,302.00
0.3961
2.1700
4.0300
15.50
Time
Initial Pop
Catch
Fish. Mort.
Nat. Mort.
Shed Rate
Time Units
1
13,739.9
1,471.00
0.2507
0.1400
0.2600
0.50
2
9,924.0
1,386.00
0.1845
0.1400
0.2600
1.00
3
5,531.6
309.00
0.0700
0.1400
0.2600
1.00
4
3,457.1
59.00
0.0209
0.1400
0.2600
1.00
5
2,269.4
26.00
0.0139
0.1400
0.2600
1.00
6
1,500.3
7.00
0.0056
0.1400
0.2600
1.00
7
1,000.0
44.00
0.0184
0.1400
0.2600
10.00
TOTALS
3,302.00
0.6044
2.1700
4.0300
15.50
Time
Initial Pop
Catch
Fish. Mort.
Nat. Mort.
Shed Rate
Time Units
1
9,218.3
1,471.00
0.3859
0.1400
0.2600
0.50
2
6,223.0
1,386.00
0.3114
0.1400
0.2600
1.00
3
3,055.1
309.00
0.1305
0.1400
0.2600
1.00
4
1,797.4
59.00
0.0404
0.1400
0.2600
1.00
5
1,157.1
26.00
0.0276
0.1400
0.2600
1.00
6
754.5
7.00
0.0114
0.1400
0.2600
1.00
7
500.0
44.00
0.0386
0.1400
0.2600
10.00
TOTALS
3,302.00
1.1003
2.1700
4.0300
15.50
OCR for page 73
An Assessment of Atlantic Bluefin Tuna
Time
Initial Pop
Catch
Fish. Mort.
Nat. Mort.
Shed Rate
Time Units
1
6,945.2
1,471.00
0.5296
0.1400
0.2600
0.50
2
4,363.3
1,386.00
0.4771
0.1400
0.2600
1.00
3
1,815.0
309.00
0.2293
0.1400
0.2600
1.00
4
967.3
59.00
0.0768
0.1400
0.2600
1.00
5
600.5
26.00
0.0539
0.1400
0.2600
1.00
6
381.4
7.00
0.0224
0.1400
0.2600
1.00
7
250.0
44.00
0.0854
0.1400
0.2600
10.00
TOTALS
3,302.00
1.9787
2.1700
4.0300
15.50
Time
Initial Pop
Catch
Fish. Mort.
Nat. Mort.
Shed Rate
Time Units
1
5,570.1
1,471.00
0.6844
0.1400
0.2600
0.50
2
3,238.9
1,386.00
0.7082
0.1400
0.2600
1.00
3
1,069.4
309.00
0.4243
0.1400
0.2600
1.00
4
468.9
59.00
0.1646
0.1400
0.2600
1.00
5
266.6
26.00
0.1253
0.1400
0.2600
1.00
6
157.7
7.00
0.0554
0.1400
0.2600
1.00
7
100.0
44.00
0.3143
0.1400
0.2600
10.00
TOTALS
3,302.00
4.9628
2.1700
4.0300
15.50
OCR for page 74
An Assessment of Atlantic Bluefin Tuna
TABLE 3-24 Estimates of transfer rates from east to west in the Atlantic Ocean, using tagging data and assuming a natural mortality rate of 0.14, a tag shedding rate of 0.26, and an eastern catch of 339 and a western catch of 17, with a ratio of 0.0501.
Pe = 0.5244
Fe = 0.0430
Pe = 0.3616
Fe = 0.0656
Pe = 0.1330
Fe = 0.2606
Pw = 0.9147
Fw = 0.0256
Te = 0.0206
Te = 0.0216
Te = 0.0316
Pw = 0.6558
Fw = 0.0390
Te = 0.0194
Te = 0.0204
Te = 0.0298
Pw = 0.4400
Fw = 0.0710
Te = 0.0171
Te = 0.0179
Te = 0.0262
Pw = 0.3315
Fw = 0.1277
Te = 0.0141
Te = 0.0148
Te = 0.0217
Pw = 0.2659
Fw = 0.3202
Te = 0.0096
Te = 0.0101
Te = 0.0147
Pw = Fraction of captured tags which are reported in the west.
Pe = Fraction of captured tags which are reported in the east
Fw = Annual instantaneous fishing mortality rate in the west.
Fe = Annual instantaneous fishing mortality rate in the east.
Te = Annual instantaneous rate of transfer from the east to the west.
OCR for page 75
An Assessment of Atlantic Bluefin Tuna
TABLE 3-25 Estimates of transfer rates from west to east in the Atlantic Ocean, using tagging data and assuming a natural mortality rate of 0.14, a tag shedding rate of 0.26, and a western catch of 3037 and an eastern catch of 65, with a ratio of 0.0214.
Pw = 0.9147
Fw = 0.0256
Pw = 0.4400
Fw = 0.0710
Pw = 0.2659
Fw = 0.3202
Pe = 0.5244
Fe = 0.0430
Tw = 0.0099
Tw = 0.0131
Tw = 0.0358
Pe = 0.3616
Fe = 0.0656
Tw = 0.0098
Tw = 0.0130
Tw = 0.0356
Pe = 0.1330
Fe = 0.2606
Tw = 0.0096
Tw = 0.0127
Tw = 0.0347
Pe = 0.1037
Fe = 0.4739
Tw = 0.0089
Tw = 0.0119
Tw = 0.0324
Pe = 0.0978
Fe = 0.5995
Tw = 0.0085
Tw = 0.0114
Tw = 0.0311
Pw = Fraction of captured tags which are reported in the west.
Pe = Fraction of captured tags which are reported in the east.
Fw = Annual instantaneous fishing mortality rate in the west.
Fe = Annual instantaneous fishing mortality rate in the east.
Tw = Annual instantaneous rate of transfer from the west to the east.
OCR for page 76
An Assessment of Atlantic Bluefin Tuna
Western Atlantic Ocean
(1)
Pw = 0.9147
Fw = 0.0256
Zw = 0.4256
(2)
Pw = 0.6558
Fw = 0.0390
Zw = 0.4390
(3)
Pw = 0.4400
Fw = 0.0710
Zw = 0.4710
(4)
Pw = 0.3315
Fw = 0.1277
Zw = 0.5277
(5)
Pw = 0.2659
Fw = 0.3202
Zw = 0.7202
These estimates were used to compute a variety of transfer rates from east to west (Table 3-24) and from west to east (Table 3-25). Not all combinations of P values are used in Tables 3-24 and 3-25 because other results were inconsistent with VPA estimates of Chapter 4.
DISCUSSION
Shedding rates appear to be 0.20 or higher (Table 3-11). Rates estimated by comparing double-double returns to double-single returns should be the most robust estimates, because the two groups originated from the same group of tags and should have been exposed to similar histories. Rates appear to be higher in the first several quarters. Two different sets of shedding rates were used in the VPA analysis (Tables 3-12 to 3-15).
The results of this reanalysis of tagging data provide quantitative confirmation of the empirical results on transfer rates presented earlier in Chapter 3. Although the committee was unable to obtain variance estimates for these transfer rates due to time constraints, results are robust with respect to reporting rate, the parameter most likely to affect the results. Thus, the results are expected to be statistically significant. However, movement of fish from west to east seems to have varied from year to year (Table 3-16). Movement from west to east also seems to have a cumulative effect, with the percent caught in the east increasing as a function of the amount of time the tags were attached to the fish (Table 3-17).
Annual transfer rates estimated with the model (equations 1 and 2) are about 0.01 for west to east, about 0.02 for east to west, and 0.03 overall. These rates depend to a great extent on the nonreporting rate for captured tagged fish. The VPA analysis on the Spanish data is especially indicative of a high nonreporting rate. The nonreporting rate for the west could also be high. Little can be said about the estimate of natural mortality from these data. However, even though there is good reason to be skeptical about the nonreporting rate and natural mortality rate, the transfer rates are almost certainly low, because the number of tags recovered in the other area is small compared to the total tags recovered.
Information is not available to determine whether a transfer from one side
OCR for page 77
An Assessment of Atlantic Bluefin Tuna
FIGURE 3-1 Proportion of tagged bluefin tuna in area 2 (eastern Atlantic Ocean) that were originally released in area 1 (western Atlantic Ocean) for a hypothetical population with Tw = 0.03, F = 0.1, and M = 0.14.
to the other is permanent. The consequences of returning would give a different meaning to the data. If fish that transfer each year do not return, even a low transfer rate will cause a significant portion of the spawning-age fish produced as recruits on one side to be on the other side when they are sexually mature. Figure 3-1 shows the increasing proportion of tagged fish in area 2 that were originally released in area 1 for a hypothetical population with T = 0.03, F = 0.1, and M = 0.14. After 300 weeks (about 6 years), the proportion is up to 0.20 even though the annual transfer rate is only 0.03; that is, 20% of the fish on one side of the Atlantic Ocean will have originated from the other side. This example shows that small annual transfer rates can have a large effect on population dynamics over a relatively short time period.
Because none of the fish tagged in the eastern Atlantic Ocean were from the Mediterranean Sea, caution should be exercised in interpreting the east to west movement rate.
Estimates of Pe and Pw obtained from Tables 3-22 and 3-23 were calculated with the assumption of no type I (initial) tagging mortality. If some, or most, of the differences between the VPA estimates of age 1 fish in the east and west result from type I mortality, the estimates of Pe and Pw would be incorrect. However, it is the ratio Pe/Pw that appears in equation 3, and if the type I mortality was the same for east and west, the ratio would still be the same as the estimates used to estimate Tw.
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An Assessment of Atlantic Bluefin Tuna
This page in the original is blank.
Representative terms from entire chapter:
bluefin tuna