Alonzo Et Al. 2008 Incorporating Sex Change Into Stock Assessments Sheephead

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BULLEIN OF MARINE SCIENCE, 83(1): 163–179, 2008

163 Bulletin of Marine Science© 2008 R S M A S Uv M

MOE SYMPOSIUM INVIED PAPER

THE IMPORTANCE OF INCORPORATING PROTOGYNOUS

SEX CHANGE INTO STOCK ASSESSMENTS

Suzanne H. Alonzo, eresa Ish, Meisha Key, Alec D. MacCall, and Marc Mangel

ABSTRACT

A z - v, b v v - . P (--) v - . W v - -

- [C , Semicossyphus pulcher (A, 1854)] - . F, v b v - (SPR). F, ( SPR 0.08 0.6) b v v z . O , -

, q .

A v v b (B R, 1966; T, 1984; H ., 1997), b b -v . Rv b v ,

v z . Y, -v (P, 1993) v v - (C ., 1999; Lz Eb, 2002; G ., 2003; Az M, 2004, 2005). I v v - (, .., Cv M, 2002; O ., 2004; R ., 2006). A , v - v v b j

.O v

b (--) , C [Semicossyphus pulcher (A, 1854)], b b[Centropristis striata (L, 1758)], [ Mycteroperca microlepis (G B, 1879)], [ Epinephelus morio (V, 1828)].Y - v (b Az ., 2004; H ., 2006). U C v , v -



BULLEIN OF MARINE SCIENCE, VOL. 83, NO. 1, 2008164

- , - b . W - - , .

A b q - b b v . T b b b v - , b- (Q D, 1999). S- vb , b ,

b b b v . I , v, z v, b b (Az M, 2005).

I , z-v v b . F , , v v v b v b. T b v -

b , b b ( bv ) v . W b b v b , v b . C b v v v b - (Az M, 2004, 2005). W , z-v

(Az M, 2004, 2005).P , v,

. I , b , b z b b , z b, (, .., W S, 1991; L, 1994). T b b ( ) b b b . I ,

. A , - , - .

Ev , bv-b b - . F , - (SPR) . SPR

v b




, v .

D v , - - z , z , z (C, 1990). T b b , b

z , v z z-v (C, 1990). P vv v z , (C, 1990). S - z b C, v v (W, 1975; C, 1990; DM ., 1994). R v , v (C H, 2006;

Y L, 2006).

Figure 1. Nearshore populations o Caliornia sheephead (Semicossyphus pulcher ) or which liehistory data were available (rom Warner, 1975 and Cowen, 1990).



ALONZO E AL.: SEX CHANGE AND SOCK ASSESSMEN 167

A , b z C . A , v (Az M,2004), b C , (A ., 2004). S b (B Mø, 1998), .L (A ., 2004). N vb b v.

C v 1800 (S, 2001) 88% -

v- (P-Z ., 1993). A - b (P-Z ., 1993). C Bj C, M, v 25% (R-C Gzz-C, 2003). A , C N F R C D F G. L C b 1996, . I , z (12–13 ) v b b -

.

Modeling Methods and Parameter Estimation

W z- - v S S ( S, M, 1990, 1998, 2000) - C . S - z- j vv, , v . I : (1) A j-

z . (2) A bv ( , b , ) v ( b -) . (3) A b- . S- v b b . M , (1947–2003), , -

-- , - v b (CCOFI).T C b Az .(2004), v b vb . A - - v -. T - - b . F S M (2000).




b - b bv.W b, - b, - b .

W b- ( k z L

inf , F. 3) F (F, 1933; Q D, 1999)

z z q v b bv z (Hb M, 1997). B z q v b bv b

vb , (k ) z ( L

50) (Hb M,

1997). T (F. 2). F b , b vb , C I (W, 1975). I v , , , vb (Cb T S B I, M, S

N I, C; W, 1975; C, 1990) v (v b 1, F. 2) . A vb G I (F. 1), , v - bb . B - v, - “” b b . v v , v z-- C I

b W (1975: . 3.2) v z

Figure 3. Size at age predicted rom the best-ft growth parameters (squares) compared with theobserved mean size-at-age data rom Warner (1975) (curve). Error bars show the 95% confdenceinterval around the mean sizes at age; numbers above the points are sample sizes.




b v (CV = 0.11). W v b vb - v v b. W b vb (W, 1975; DM ., 1994) z-

[v = ( ) (15,000 b ) (86 b )]. N - v vb C .

W (M) - . E (Z) C (M) (F) 0.29 0.55 (b 1, C, 1990). S, bv 9 21 bb b (Lb, 1955;W, 1975; C, 1990). W b

H (1983) b M v C. A b 53 b (F,1974) , b v , b 15 30 (Lb, 1955; W, 1975; C,1990). B b- bv 21 (C, 1990), H (H, 1983) bM = 0.20, b b M (M = 0.15, - 30 ) bv b b . W v (M = 0.3, 15 ) .

Gv b- , S b, (.., - ) b, qb v . T b z - (F. 2) b ( ) b. O b ’ , SPR b, b, b. T SPRb b .

Table 1. Total mortality (natural and shing), growth, maturity, and sex-change parameter estimates for four nearshore populations from Warner (1975) (Catalina Island) and Cowen (1990)(other locations). All lengths have been converted to fork lengths (in cm) for consistency with theavailable sheries data. The data from Catalina Island were the best available for estimation of these parameters and were therefore used in the baseline stock-assessment model.

Parameter Catalina Island San Benitos Island Cabo Thurloe San Nicolas Island

Total mortality 0.35 0.43 0.41 0.29

Maximum age in sample 20 10 12 21

Estimated Linf

83.86 130.00 145.86 85.19

k growth 0.068 0.026 0.028 0.064

L1 (age 1) 12.92 7.35 10.16 16.40

L2 (age 13) 52.49 40.22 49.38 53.30

L50

maturity 25.24 26.90 21.80 33.30

L50 sex change 36.77 33.40 30.90 54.50k maturity (slope) 1.060 0.880 0.520 0.780

k sex change (slope) 0.32 0.46 0.21 0.54




S b v C b - .T b v b j. B v , v z . S Az . (2004) -

, , .

Results

he Effect of Including Sex Change on Stock Dynamics.—U - , v C I M = 0.2 (b 1; W, 1975). I ,b - b b 1950 , b “” b b

1960–1990 (F. 4). T - b b . I b (F. 4), b b ( -) v - (F. 4A,B). T v - v z v, (F. 5). A v- v -- , v ( v b b

Figure 4. Historical total biomass and egg production estimated rom the baseline Caliornia

sheephead stock assessment model with (A) and without (B) inclusion o sex change (or M = 0.2and the Catalina Island parameters rom Table 1).




v ). T, b -, v b (F. 5).

he Effect of Female and Male Biomass on arget Measures.—O b N F M P C D F G (CDFG, 2002), SPR ( v ) SPR

50%. A v SPR

50% b b,

v , SPR b, , b, v .

SPR b v v (.., b ) ; b - v v ( ), v b . A b , v, ( ) b-

(F. 6). W - C 2003 b S (0.23) , SPR b 55% v (F. 7A), b SPR b b ( SPR = 0.17, SPR = 0.07, F. 7A). I (.., ), b v .

Figure 5. Size selectivities or the recreational and commercial (live trap, setnet, and hook-and-line) fsheries estimated with the baseline Synthesis model. The vertical solid lines represent the L

50size at maturity and size at sex change or the baseline model. Results are shown or the case

where M = 0.2 and or the Catalina Island lie-history parameters but are essentially the same or

other values rom Table 1.




F v (M = 0.2), SPR b b b ( 0.23) - 15% SPR. L M v SPR ( SPR = 0.09, F. 7B), b M, v ( SPR = 0.5, F. 7C). C, b M b SPR . (F. 7). A , M (.., SPR M = 0.15 0.30 SPR = 0.45 0.78, F. 7A–C),b b b b

(.., SPR = 0.16 v 0.55, F. 7A) b q.

he Effect of Site-specific Life History Parameters.—A C b , z v b bv. O v v , b , , - . I v SPR

b b - b v b .

O C v Cb T q qv C I(F. 8A,D). I , - S N I S B I, v SPR (F. 8B,C). M S N I z ( b 1),

Figure 6. Spawning potential ratio (SPR) as a unction o exploitation, estimated rom emale ortotal mature biomass, or versions o the stock assessment model that include or ignore sex change(with Catalina Island lie-history parameters rom Table 1 and M = 0.2).




Figure 7. Spawning potential ratio (SPR) as a unction o exploitation (estimated rom emale,male, or total mature biomass) or three dierent natural mortality values. Estimated current ex-ploitation rates and target SPR

50%are also shown (baseline model with Catalina Island lie-history

parameters rom Table 1 otherwise).




v v (F. 5). F b S B I, v - b b . T ,, - bv v - SPR b , b v b SPR b b (.., b).

Discussion

A v , v v v . O - z-v v b

. W b v- z-v b (F. 4, 5). Hv, , z - v v (, .., W S, 1991), b, , b . O , v - b,

b , b

Figure 8. SPR based on total mature and emale biomass estimated rom lie-history parameters(rom Table 1) rom the our populations at the estimated current level o exploitation (0.23 orM = 0.2).




b . S , , b .

O b b z-v (F. 7), v b - . A b b v , b b . T b b v b b. I, b b , v v b

b z (Az M, 2004, 2005). W v SPR SPR b - . F b v .T v b .

O - v v

, b - - . F, b b . S v - bv , b , b .A b bv, z (.., v , b b z) -

, b b b z. B b C , v, , z b, .C, b b v - b b b v, v , v

v .T - b , b . F, b v . A - , b v . T bb - b v ,

v bv b b




b . I , b v ( b ). A, b ; - v b . Dv - , , v , b b b .

Acknowledgments

W E. J. D, M Lb, S. R, A. S . W v R. M S S - S. W R.C, B. E, M. Lv, W. V B, D. V , W. D, S. Wz, H. K, B.L, . L, E. K, D. W-Vb -. F v b C S A R (CSAR), b S Cz Lb N M F Sv UCSC; C D F G; NOAA F; N SF IOB 0450807 SHA. W F. C. C, v, A. B. T , v .

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Addresses: (S.H.A.) Department of Ecology and Evolutionary Biology, Yale University, P.O. Box 208106, New Haven, Connecticut 06520. (.I.) Environmental Defense, 257 Park AvenueSouth, New York, New York 10010. (M.K.) California Department of Fish and Game, c/o Na-tional Marine Fisheries Service, 110 Shaffer Rd., Santa Cruz, California 95060. (A.D.M.) Na-

tional Marine Fisheries Service Southwest Fisheries Science Center, Fisheries Ecology Division,110 Shaffer Road, Santa Cruz, California 95060. (M.M.) Center for Stock Assessment Research

and Department of Applied Mathematics and Statistics, University of California, Santa Cruz,1156 High Street, Santa Cruz, California 95064. Corresponding Author: (S.H.A.) E-mail:

<[email protected]>.

Alonzo Et Al. 2008 Incorporating Sex Change Into Stock Assessments Sheephead

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