bull bull

Technical Paper no 212004

UY 0 GROWTH RATE OF SOUTHERN SALTWATER TERRAPIN

(Batagllr bask a Gray)

v l)ltlIUYl HmiddotH1ltWH~

~fl1Yl~ 1mnUJlllrJ ( I

ItlTJflU1i 1J1H1J

Chaiwat Ratanadadas

Wannut Hirunchullia

Supawat Komolmarl

Saowakon Rungruang

Inland Fisheries Research and Development Bureau

Department of Fisheries

Ministry of Agriculture and Cooperatives

bullbull

I 9 aJrgtXtJ~))1

A V Technical Paper no 212004101lQl111flfl1Hllllltl lisamplt~

4 v ~ ~lt fl1jfl f)jl ~fI l1fIli~jOJ1lJ t~IejVIl f) j~ i1t4

v

Sl1IDY ON GROWTH RATE OF SOUTHERN SALTWATER TERRAPIN

(Batagur baska Gray)

9 ~~I to 4 IV

~lUu~~~Ul~~~~~1A~a

bull v v t)llIt)a~ ~~~1~~iI 60

1m ~ci~o-ltlI1eI66 leIampciw

1mJ

~Jlillll iVlumVlUl

V Q~

~~flUn I1~ru~III1

v

~fII1ill 1f1~llll oIJ

mnflUfi lilgttH

ChaiVl ai ~atanadadas

Wannut Hirunchulha

Supawat Komolmarl

Saowakon Rungruang

Satun Inland Fisheries Station

Amphoe La-ngu Satun 91110

Tel 0-7478-1299

2004

IV c QltII d

IIItIfllIlllUl)rJllUfl 42-44-2-15-73-01-1-480-049

A-e10 lJJfsvq m2vJVfI t~QLJJLflLH QJt P = If

Ud-ell~ll~l~Mll-eS W~qlnOS mftLQQJb I

rtlb Of

bull bull

bull bull

bullfjU61jJmmYlIlnmh1J~1l1~~Ylll~ l)lijl)~ ~Ylfl~~ 5GlloOO v ~ bull

bull rrf1lihJ~1J~ih~IIltOnrl~iftitlii l)iiJirr ~itltlii GllbGllo

v i f2i1Jl1l11 fIIJ1IlJ111l

thirflYlIllUlH1IclltJfll)IgtUIlflIU 11lufllnh1J~ m1JlhJ1J~ ~IIltom m~I1Vi1 0500

bull rrf1lilth1J~1l1~~~~)rilltT~1I l)11~ Ui~1I I1f)lto

lliJ~1milm)lUm~Ml~t1U 1 iJ ~~HIl1Jt~milll (6111 2 ~6U) O~)lq 61J ~lmU 7 iu 1 y

11 5 ~1 ~1lJi1ulmj6fl6Ufl~IIUU11I 50 1l1~HlJll~ ~~tl1l1 30 11U~lJm i~1i1l111lllfftl11lJi1Ullfl~~ y y

liluml1lnjfl1uluilIlHlmlJ 10 1i6JlUrU6~1l1I1irflr II tTf1liJ1ii1J~1l1~IIltOnrrllrr~1I 1lU1111 36

~ III $I ~ I ~ ~ 1 fi ~l)U IlIml1JtJ11flillmIll1ltlUl)~llllml)lUm~ 2 11Il)1J mmq 9 II 1I~91 851plusmn002 tH 4332plusmn103

lllUVlI1JII~ m11Jfl1Hm~lIe~miilJr~i 854plusmn009 O~ 3980plusmn168 tll1JVlIlJlI~ l1IUll1irf1l~iilJil~I~

9280plusmn228 O~ 1O5240Qplusmn19269 flilJ ilrrlJfllimllJnlJYl1J1liI1h~mllJtIl1mlIe~t1t1mq1lu L bull

598 l - e middot~U38 ( + 02(6) U1lrrlJfllimllJnlJ1I1liwh~lhH1rflft6mllJtll1m1I6~1i111l1J W =

01l46 L30309 lifh R2 = 09936 fhmlllftl1~u6~fl1illllqJllJlll 0138 ~~rrllJlitnnfllf111lJtl11

mItl)~U6~lthmtl1l1~~~lll L = 598 llfllAllJlli 1IluihimlflliIIljf)jIVllJ 111 Iul1i1ll)~flliI~1JUl)~ r J J 4 lid IIIJ 1

fl1lJlJl1fll1t6jfl611U1~f1~l1W6lJ61fl20 1lU1I 1]

bullfhthilru l~lm)lll fllililltlllAlJ 111 flmntl~

u bull

bullbull

I 9 aJrgtXtJ~))1

A V Technical Paper no 212004101lQl111flfl1Hllllltl lisamplt~

4 v ~ ~lt fl1jfl f)jl ~fI l1fIli~jOJ1lJ t~IejVIl f) j~ i1t4

v

Sl1IDY ON GROWTH RATE OF SOUTHERN SALTWATER TERRAPIN

(Batagur baska Gray)

9 ~~I to 4 IV

~lUu~~~Ul~~~~~1A~a

bull v v t)llIt)a~ ~~~1~~iI 60

1m ~ci~o-ltlI1eI66 leIampciw

1mJ

~Jlillll iVlumVlUl

V Q~

~~flUn I1~ru~III1

v

~fII1ill 1f1~llll oIJ

mnflUfi lilgttH

ChaiVl ai ~atanadadas

Wannut Hirunchulha

Supawat Komolmarl

Saowakon Rungruang

Satun Inland Fisheries Station

Amphoe La-ngu Satun 91110

Tel 0-7478-1299

2004

IV c QltII d

IIItIfllIlllUl)rJllUfl 42-44-2-15-73-01-1-480-049

A-e10 lJJfsvq m2vJVfI t~QLJJLflLH QJt P = If

Ud-ell~ll~l~Mll-eS W~qlnOS mftLQQJb I

rtlb Of

bull bull

bull bull

bullfjU61jJmmYlIlnmh1J~1l1~~Ylll~ l)lijl)~ ~Ylfl~~ 5GlloOO v ~ bull

bull rrf1lihJ~1J~ih~IIltOnrl~iftitlii l)iiJirr ~itltlii GllbGllo

v i f2i1Jl1l11 fIIJ1IlJ111l

thirflYlIllUlH1IclltJfll)IgtUIlflIU 11lufllnh1J~ m1JlhJ1J~ ~IIltom m~I1Vi1 0500

bull rrf1lilth1J~1l1~~~~)rilltT~1I l)11~ Ui~1I I1f)lto

lliJ~1milm)lUm~Ml~t1U 1 iJ ~~HIl1Jt~milll (6111 2 ~6U) O~)lq 61J ~lmU 7 iu 1 y

11 5 ~1 ~1lJi1ulmj6fl6Ufl~IIUU11I 50 1l1~HlJll~ ~~tl1l1 30 11U~lJm i~1i1l111lllfftl11lJi1Ullfl~~ y y

liluml1lnjfl1uluilIlHlmlJ 10 1i6JlUrU6~1l1I1irflr II tTf1liJ1ii1J~1l1~IIltOnrrllrr~1I 1lU1111 36

~ III $I ~ I ~ ~ 1 fi ~l)U IlIml1JtJ11flillmIll1ltlUl)~llllml)lUm~ 2 11Il)1J mmq 9 II 1I~91 851plusmn002 tH 4332plusmn103

lllUVlI1JII~ m11Jfl1Hm~lIe~miilJr~i 854plusmn009 O~ 3980plusmn168 tll1JVlIlJlI~ l1IUll1irf1l~iilJil~I~

9280plusmn228 O~ 1O5240Qplusmn19269 flilJ ilrrlJfllimllJnlJYl1J1liI1h~mllJtIl1mlIe~t1t1mq1lu L bull

598 l - e middot~U38 ( + 02(6) U1lrrlJfllimllJnlJ1I1liwh~lhH1rflft6mllJtll1m1I6~1i111l1J W =

01l46 L30309 lifh R2 = 09936 fhmlllftl1~u6~fl1illllqJllJlll 0138 ~~rrllJlitnnfllf111lJtl11

mItl)~U6~lthmtl1l1~~~lll L = 598 llfllAllJlli 1IluihimlflliIIljf)jIVllJ 111 Iul1i1ll)~flliI~1JUl)~ r J J 4 lid IIIJ 1

fl1lJlJl1fll1t6jfl611U1~f1~l1W6lJ61fl20 1lU1I 1]

bullfhthilru l~lm)lll fllililltlllAlJ 111 flmntl~

u bull

A-e10 lJJfsvq m2vJVfI t~QLJJLflLH QJt P = If

Ud-ell~ll~l~Mll-eS W~qlnOS mftLQQJb I

rtlb Of

bull bull

bull bull

bullfjU61jJmmYlIlnmh1J~1l1~~Ylll~ l)lijl)~ ~Ylfl~~ 5GlloOO v ~ bull

bull rrf1lihJ~1J~ih~IIltOnrl~iftitlii l)iiJirr ~itltlii GllbGllo

v i f2i1Jl1l11 fIIJ1IlJ111l

thirflYlIllUlH1IclltJfll)IgtUIlflIU 11lufllnh1J~ m1JlhJ1J~ ~IIltom m~I1Vi1 0500

bull rrf1lilth1J~1l1~~~~)rilltT~1I l)11~ Ui~1I I1f)lto

lliJ~1milm)lUm~Ml~t1U 1 iJ ~~HIl1Jt~milll (6111 2 ~6U) O~)lq 61J ~lmU 7 iu 1 y

11 5 ~1 ~1lJi1ulmj6fl6Ufl~IIUU11I 50 1l1~HlJll~ ~~tl1l1 30 11U~lJm i~1i1l111lllfftl11lJi1Ullfl~~ y y

liluml1lnjfl1uluilIlHlmlJ 10 1i6JlUrU6~1l1I1irflr II tTf1liJ1ii1J~1l1~IIltOnrrllrr~1I 1lU1111 36

~ III $I ~ I ~ ~ 1 fi ~l)U IlIml1JtJ11flillmIll1ltlUl)~llllml)lUm~ 2 11Il)1J mmq 9 II 1I~91 851plusmn002 tH 4332plusmn103

lllUVlI1JII~ m11Jfl1Hm~lIe~miilJr~i 854plusmn009 O~ 3980plusmn168 tll1JVlIlJlI~ l1IUll1irf1l~iilJil~I~

9280plusmn228 O~ 1O5240Qplusmn19269 flilJ ilrrlJfllimllJnlJYl1J1liI1h~mllJtIl1mlIe~t1t1mq1lu L bull

598 l - e middot~U38 ( + 02(6) U1lrrlJfllimllJnlJ1I1liwh~lhH1rflft6mllJtll1m1I6~1i111l1J W =

01l46 L30309 lifh R2 = 09936 fhmlllftl1~u6~fl1illllqJllJlll 0138 ~~rrllJlitnnfllf111lJtl11

mItl)~U6~lthmtl1l1~~~lll L = 598 llfllAllJlli 1IluihimlflliIIljf)jIVllJ 111 Iul1i1ll)~flliI~1JUl)~ r J J 4 lid IIIJ 1

fl1lJlJl1fll1t6jfl611U1~f1~l1W6lJ61fl20 1lU1I 1]

bullfhthilru l~lm)lll fllililltlllAlJ 111 flmntl~

u bull

bull bull

bull bull

bullfjU61jJmmYlIlnmh1J~1l1~~Ylll~ l)lijl)~ ~Ylfl~~ 5GlloOO v ~ bull

bull rrf1lihJ~1J~ih~IIltOnrl~iftitlii l)iiJirr ~itltlii GllbGllo

v i f2i1Jl1l11 fIIJ1IlJ111l

thirflYlIllUlH1IclltJfll)IgtUIlflIU 11lufllnh1J~ m1JlhJ1J~ ~IIltom m~I1Vi1 0500

bull rrf1lilth1J~1l1~~~~)rilltT~1I l)11~ Ui~1I I1f)lto

lliJ~1milm)lUm~Ml~t1U 1 iJ ~~HIl1Jt~milll (6111 2 ~6U) O~)lq 61J ~lmU 7 iu 1 y

11 5 ~1 ~1lJi1ulmj6fl6Ufl~IIUU11I 50 1l1~HlJll~ ~~tl1l1 30 11U~lJm i~1i1l111lllfftl11lJi1Ullfl~~ y y

liluml1lnjfl1uluilIlHlmlJ 10 1i6JlUrU6~1l1I1irflr II tTf1liJ1ii1J~1l1~IIltOnrrllrr~1I 1lU1111 36

~ III $I ~ I ~ ~ 1 fi ~l)U IlIml1JtJ11flillmIll1ltlUl)~llllml)lUm~ 2 11Il)1J mmq 9 II 1I~91 851plusmn002 tH 4332plusmn103

lllUVlI1JII~ m11Jfl1Hm~lIe~miilJr~i 854plusmn009 O~ 3980plusmn168 tll1JVlIlJlI~ l1IUll1irf1l~iilJil~I~

9280plusmn228 O~ 1O5240Qplusmn19269 flilJ ilrrlJfllimllJnlJYl1J1liI1h~mllJtIl1mlIe~t1t1mq1lu L bull

598 l - e middot~U38 ( + 02(6) U1lrrlJfllimllJnlJ1I1liwh~lhH1rflft6mllJtll1m1I6~1i111l1J W =

01l46 L30309 lifh R2 = 09936 fhmlllftl1~u6~fl1illllqJllJlll 0138 ~~rrllJlitnnfllf111lJtl11

mItl)~U6~lthmtl1l1~~~lll L = 598 llfllAllJlli 1IluihimlflliIIljf)jIVllJ 111 Iul1i1ll)~flliI~1JUl)~ r J J 4 lid IIIJ 1

fl1lJlJl1fll1t6jfl611U1~f1~l1W6lJ61fl20 1lU1I 1]

bullfhthilru l~lm)lll fllililltlllAlJ 111 flmntl~

u bull

2

STUDY ON GROWTH RATE OF SOUTHERN SALTWATER TERRAPIN

(Batagur baska Gray)

Chaiwat Ratanadadas

Phattalung Inland Fisheries Research and Development Center Amphoe Muang

Phattalung 93000 Thailand

Wannut Hirunchulha

Chaiyaphum Inland Fisheries Station Amphoe Jatural Chaiyaphum 36130 Thailand

Supawat Komolmarl

Fisheries Development and Technology Transfer Bureau Fisheries Department

Bangkok 10900 Thailand

Saowakon Rungruang

Satun Inland Fisheries Station Amphoe La-ngu Satun 91110 Thailand

ABSTRACT

A study on growth rate of southern saltwater terrapin (Batagur baska Gray) was

carried out from May 1998 to Ma 301 in 6 periods of age (newborn to 6 years) The 35

terrapins were cultured in 50 m2 concrete pond with 30 cm of water depth Terrapin were fed

with chopped trash fish and moming glory at 10 of body weight for 36 months The results

showed the average carapace length of8512002 - 4332plusmn103 em average carapace width of

854plusmnO09 - 3980plusmnL68 em and average weight of 9280plusmn228 - 1052400plusmn19269 g The

equation of age-length relationship was L == 598 (1 - e -0138( + Q216~ and lengthmiddotweight

bull rclationship was W = 01146 L 30309 ~ == 09936) The estimate of growth parameters were 8

0138 per year of growth constant (K) 598 cm of the maximum asymtotic length (L) The

growth rate ofcarapace length was quite stable from 20 years old

Key words southern saltwater terrapin (Batagur baska Gray) growth culture

I)

bull bullbull

bull bull

3

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lY1llffflll1fll1111tlfiIJ11lIJ6~flliliitJ~1~lmmllillmQ~1~ 1 till

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i

4

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1ImlMiJulllJ-U~IIJ~11l 21~JU~lJfI~~ nmtillJltl 0900 U ~il

ju11l1l 11l11~1JIllillJllIill1 1l1htj1~6~11lIllt91Vll

fI11llllum~lllulgth~ (pH) 1Iilllil1Ji pH meter bull v bulltI~lJ1U1J6flJlIIlWl11l1l161Wll1 (DO) fl11lJlll u~h~ (alkalinity) fl11llmIilH

(hardness) 1l11m1uJ1l ilillltlfl illvlUllJ (free carbondioxide) 1~ lliltlfllJ ~1il1gtIjfl

IilllJ1TI~fl~l1eH1ullJll1~1l11~lFmll (2528) mhtl1lij11i1f1flJ~li1J

25 fllSl1lJi1l1-laquom1l 1 v v V 0 r y ltJ V I wf ltlI Sf IIll IllUfl Il16l11ln~fl11l1011l1lmllJf11 HIIt)~f11~6~11il11l flIil111161l11 I m ~ YIJ6l11Hbull

tiru1l~lJ1IUtJ1Hl1iilmii(Jl ~1lfl1U 36 1~6U

11 l-laquollJ II fl11lJ1J1111l ~IJIIl ~m~6 ~I~11W uqj 1lll~f1l1 11 ~lll~ lI11i6Wil W fl1 wlllll~~ii~~1middot I f1l111l~ t)JI~lJ1~ ml111ll)1Jml1JIIilf1wU6lmllJill~ItIJ~ Cslope) l6~lff1lm wn1ll~11ll~fll11lIil11B~

1Ulh~~Ilil61fJIl7 ~IhilW (lcrWf1Jl1lmgBlItiW) 1~6H1TI1IflnMfl11lJlIIh1l111l i1l1 CANeOV A) I

I i

11l1tJHHmmlJ Systat version 50 9i~Olliiifl11lJUlilnlHti1l~~111ll1lf1l11l7t)J~U 1~~~lJ1I~llJ ~611 W lI~~ tJ 1 nW m11tU (l1l1ilu m whltI~~ fl11l1 fflJIl1lnl wh~ fl11l11J11 il~ tJII l lm J 1il6~n1J61QII M~1

bull bull

5

Ll~~H~ t11ihl1J1~lfyampt~a~ftuhmlil1~iu lilt 5 i1111lJ 7 iu ti6Jfl1i161~ 2 1~6U tlilllQ 1234

5 HI161Q 6 iJ $1

~~ Ifl1l~I1lJl1J11f titali 1 i1i~Il~I1l~IImnl~ lilmli~lJ111 tTfl 1~fI11lJ1J11 UI1f111lJfl1N v

IIl~m ~ Iill~l~11)fl9l rilW11lhnm11JflUiuU61l1il11l ~VIlllfihiflMu~ II l~tIfIl11U11ilJ56lJ1flJ5fi a~tiu

jj1UIJjl~

23 lmnultIml11fl1I1n Jf )I v

HI11mJI1111li~ti1J1ll~MUI~UUI~fl 1 i1l1tiUNfl~~ lutY~Hi11l 1 ~l 1 l~1J11mtTfl 1~1l v bull bull

611111 11111lflhm611ln1lll11l111fl11l 1 ltI~ 1 fl~~ l1a1il11 10 tilrl911lvlll6~11lI1tTfl1 i1ll1111

li1lI1U 1500 1l

24 fl1lilf)11

Vilflmhtl1hi1l1j61l~HIl~ 1116l1~~lilfl11l1t1l1Il1UlfflJillMll 3 11l (11li1llli VlliUll11l ~ gt d r 0 l

11m IlnmtiJlIltl 0900 1l) Ull~ti1l61J LI1l1l11lg 1lUlll lIlI1lllllllfl11llllllIl1U11111J1lnmtJJlIltl 3 i11lJ~ ri6u1li16IJ111MhlljJiUI111I1300 U ~uill~Ii1J1hlilnvti~~lIltl 30 llU~llJlilJ

bull v bull

1ImlMiJulllJ-U~IIJ~11l 21~JU~lJfI~~ nmtillJltl 0900 U ~il

ju11l1l 11l11~1JIllillJllIill1 1l1htj1~6~11lIllt91Vll

fI11llllum~lllulgth~ (pH) 1Iilllil1Ji pH meter bull v bulltI~lJ1U1J6flJlIIlWl11l1l161Wll1 (DO) fl11lJlll u~h~ (alkalinity) fl11llmIilH

(hardness) 1l11m1uJ1l ilillltlfl illvlUllJ (free carbondioxide) 1~ lliltlfllJ ~1il1gtIjfl

IilllJ1TI~fl~l1eH1ullJll1~1l11~lFmll (2528) mhtl1lij11i1f1flJ~li1J

25 fllSl1lJi1l1-laquom1l 1 v v V 0 r y ltJ V I wf ltlI Sf IIll IllUfl Il16l11ln~fl11l1011l1lmllJf11 HIIt)~f11~6~11il11l flIil111161l11 I m ~ YIJ6l11Hbull

tiru1l~lJ1IUtJ1Hl1iilmii(Jl ~1lfl1U 36 1~6U

11 l-laquollJ II fl11lJ1J1111l ~IJIIl ~m~6 ~I~11W uqj 1lll~f1l1 11 ~lll~ lI11i6Wil W fl1 wlllll~~ii~~1middot I f1l111l~ t)JI~lJ1~ ml111ll)1Jml1JIIilf1wU6lmllJill~ItIJ~ Cslope) l6~lff1lm wn1ll~11ll~fll11lIil11B~

1Ulh~~Ilil61fJIl7 ~IhilW (lcrWf1Jl1lmgBlItiW) 1~6H1TI1IflnMfl11lJlIIh1l111l i1l1 CANeOV A) I

I i

11l1tJHHmmlJ Systat version 50 9i~Olliiifl11lJUlilnlHti1l~~111ll1lf1l11l7t)J~U 1~~~lJ1I~llJ ~611 W lI~~ tJ 1 nW m11tU (l1l1ilu m whltI~~ fl11l1 fflJIl1lnl wh~ fl11l11J11 il~ tJII l lm J 1il6~n1J61QII M~1

bull bull

6

~ ~ lt 31 mnnmllJlflJIfWll

bull Ullt1iltJlllUWIl1fWl1 mllJf1~l~miW~ UllmllJV11m~ltJ~tvilmJlUlJl11lmllJalJ~U

~lliNUl1l11f1Hllf111lJV11m~H mllJlim~u~lliNUlllUmlllJ111Jf1~Hf1~~ltJ~ UllmllJ

I qlaquol I fflJWUll~1l11~f111lJml~tlllmllJUl1mgt1e~ YlllJ111llN Rounsefell and Everhart (1953) VlUlf~~

tnllJlilJ~tI)lf1m~1ImllMfmjj~HlV (regression analysis) itlllllle~fflJm~vf1lhti~lI11llflJmmitI bull

YlH ~~n

W = a Lh

logW = log a + b log L

bw a w

logW = log a + b log w

tnllJlilJ~tI~llh~tnllJf1~HUlltnllJVl1m~N

w = a+bLbull w = Illllllf11l11tvi 1111Uf1ilJ

L = J 11lJV11myen1wl] tI lliU91lJ Yl ~

w = J 11lJf111~m ~ ltJ~t11 tI tlfU AtlJYl ~ JlJ~1ab

~

32 mnh~lJlilftllmliJIYlltJif1l~Jil~91U1~ 011 J 1 I Itt I I d lt) tf Itl1IJ()lJflfIlt~flVIlltJ~lIlfmelUInll1t1VlI~IlU tlU~llli1t11lJl1f1nlMll1VlllmYle

Il()~mllftlJiflll111llJlJil118~m~ii1JlflllIH von Bertalanffy (1938) flltJftl L Illll K ~U11i Gulland

and Holt plot (1959 ~Hfllll Sparre and Venema 1992) ~~lflJmi J

Q

= KL- K X L61

L+~ + LtlJe L =

2

K ftl ffmhdhliff1 mAllIYl

L bull w

J11lJIJl1Mlf ~IlltJ~lffl1

l~ K = -b

A un~ La =

K

7

~lf)fllll1llll1l~IIi1NI~lm1llW hWTW 7 111 1 ll 5 il1 t](l 13f1l~11hif1lfill()1Q 2 lii(lU 13fl

1~1J1Q 12 345 um 611e1i~~mllltlnmIlf)~m~tI 851plusmn0Q2 1337plusmnO06 1990plusmn119 2360plusmn022 (I JJ =i

2934plusmnO34 3346plusmnO69 till 3706plusmnO68 llfUllllim 1l1llll1ll1J mlllfl1l~mIlf)~IIHW 854plusmn009

1277plusmn026 1902plusmnl04 2 166plusmn023 2664plusmnO82 3004plusmn100 llll 3420plusmn157 lfU911llIl1 1l1llih911J luii

o W s lJ1MlJflll11l1l1l n80plusmn228 35240plusmn477 94000plusmn19494 148000plusmn5701 302000plusmn16047

470800plusmn13609 IIll 683400plusmn4336 dll fllllll1911J 1Iltllli()1Igt1l10UIUtiJfliJlJf1~IllllJlll 50 1l1Jl~

lIlll (mlllMW1U11u 143 i1l11~lIi11~()h) Vhll1Jlhmntl 30 1lfU911lli1 1lJlJ1lll 311 ilHllmlVlllllf)~

dlf)V1llllf)~iII~~1f1l()1lJ11~lliuI1llJl1lll 12 Ull 311 Vl1Jhhm()1ulu~h1l1Jiu11f1

I 4q ~ 0

13f1lllllnq 2 1llW lImlllllnmIlf)~mlll 1418plusmn087 1900plusmn068Illl 2570plusmn049 lIUllllllll 1l1lllllll1J

i1l~I~lI~1f1vilf)lQ III iifl11lllll1mIlf)~m~I 1994plusmn187 2378plusmn298 Ull 28I6plusmn224 1li1J911)J1l1 Illll

h~IJ ilJ~I~)Ji)lmvil()1Q 2 II iifl1llllnmIlf)~I1~I 2534plusmn188 2968plusmn192 1I11 3340plusmn164

1li1l~1)J1l1 m)J~l~1J i1l~1~)Ji)1f1l~1()lQ 3 1J_iiml)Jm1mIlf)~11l~1I 2988plusmn125 3362plusmn1l7 till

37707070 lIll~ltlllllll)J~l~1J i1l~~lIi)1f1~lfJ1~ 41J ~ml)Jm1mIlf)~m~tI 3418plusmn058 3848plusmnOI9

11ft 4O60plusmnO71 1lf1l9illlm 1l1llihJ1J fh1lllJ~I~lI~1f1~1fJ11J 5 iJ iimllltlllmIl(l~m~tI 3700plusmn096

4O42plusmnL09Ull 4202plusmnO86 1llW~llllll 1l1ll~1911J 11lvililJ~~t1l1fmh1l1Q 61J iifl11)Jtll1mIlHm~1J Q 041 d ampd

3971ttO834174plusmnO47 tift 4332plusmnl03 1liWllltllll 1l1tlll1llIJ (llln~Vl I) lf~lJf)WllfJ1ml)JlIl1mIl)~

Utm~il~iiV11JIliUU11il~fJlllrhilm)1lJfJ1~ 2 lii(llJ I 2 3 4 5 6 7 8 Ullit 9 1J ilfl1llJtlllbull ~ ~ ~

mlltHlIlllVlYllf11J 8Si 1378 1961 2461 2927 3367 3756 4027 4188 11ll 4332 1lflJllllli11 bull

7

~lf)fllll1llll1l~IIi1NI~lm1llW hWTW 7 111 1 ll 5 il1 t](l 13f1l~11hif1lfill()1Q 2 lii(lU 13fl

1~1J1Q 12 345 um 611e1i~~mllltlnmIlf)~m~tI 851plusmn0Q2 1337plusmnO06 1990plusmn119 2360plusmn022 (I JJ =i

2934plusmnO34 3346plusmnO69 till 3706plusmnO68 llfUllllim 1l1llll1ll1J mlllfl1l~mIlf)~IIHW 854plusmn009

1277plusmn026 1902plusmnl04 2 166plusmn023 2664plusmnO82 3004plusmn100 llll 3420plusmn157 lfU911llIl1 1l1llih911J luii

o W s lJ1MlJflll11l1l1l n80plusmn228 35240plusmn477 94000plusmn19494 148000plusmn5701 302000plusmn16047

470800plusmn13609 IIll 683400plusmn4336 dll fllllll1911J 1Iltllli()1Igt1l10UIUtiJfliJlJf1~IllllJlll 50 1l1Jl~

lIlll (mlllMW1U11u 143 i1l11~lIi11~()h) Vhll1Jlhmntl 30 1lfU911lli1 1lJlJ1lll 311 ilHllmlVlllllf)~

dlf)V1llllf)~iII~~1f1l()1lJ11~lliuI1llJl1lll 12 Ull 311 Vl1Jhhm()1ulu~h1l1Jiu11f1

I 4q ~ 0

13f1lllllnq 2 1llW lImlllllnmIlf)~mlll 1418plusmn087 1900plusmn068Illl 2570plusmn049 lIUllllllll 1l1lllllll1J

i1l~I~lI~1f1vilf)lQ III iifl11lllll1mIlf)~m~I 1994plusmn187 2378plusmn298 Ull 28I6plusmn224 1li1J911)J1l1 Illll

h~IJ ilJ~I~)Ji)lmvil()1Q 2 II iifl1llllnmIlf)~I1~I 2534plusmn188 2968plusmn192 1I11 3340plusmn164

1li1l~1)J1l1 m)J~l~1J i1l~1~)Ji)1f1l~1()lQ 3 1J_iiml)Jm1mIlf)~11l~1I 2988plusmn125 3362plusmn1l7 till

37707070 lIll~ltlllllll)J~l~1J i1l~~lIi)1f1~lfJ1~ 41J ~ml)Jm1mIlf)~m~tI 3418plusmn058 3848plusmnOI9

11ft 4O60plusmnO71 1lf1l9illlm 1l1llihJ1J fh1lllJ~I~lI~1f1~1fJ11J 5 iJ iimllltlllmIl(l~m~tI 3700plusmn096

4O42plusmnL09Ull 4202plusmnO86 1llW~llllll 1l1ll~1911J 11lvililJ~~t1l1fmh1l1Q 61J iifl11)Jtll1mIlHm~1J Q 041 d ampd

3971ttO834174plusmnO47 tift 4332plusmnl03 1liWllltllll 1l1tlll1llIJ (llln~Vl I) lf~lJf)WllfJ1ml)JlIl1mIl)~

Utm~il~iiV11JIliUU11il~fJlllrhilm)1lJfJ1~ 2 lii(llJ I 2 3 4 5 6 7 8 Ullit 9 1J ilfl1llJtlllbull ~ ~ ~

mlltHlIlllVlYllf11J 8Si 1378 1961 2461 2927 3367 3756 4027 4188 11ll 4332 1lflJllllli11 bull

i

8

IlmNl1 1 1l11lJV11mtl~m~~IItl~t~lmmlJiil1l1lNtnV~hnjlfltlUfl~lltllulJmn 3 II

d ~ ~ ml)Wl1mIIl~liliW (lilJ)

1l1qlll1lJlI U ~ IgtlJIIW III 2lJ 3lJ

lIl1l1~ 21tlU 851plusmnOO2 1418plusmnO87 190QplusmnO68 2570plusmnO49

l~ltll~ IU 1337plusmnOO6 1994plusmn187 2378plusmn298 2816plusmn224

~lm~ 2U 1990plusmn1l9 2534plusmn188 2968plusmn192 3340plusmn164

~lllQ 3U 2360plusmnO22 2988plusmn125 3362plusmn1l7 3770plusmn070

~ltl1Q 4U 2934plusmnO34 3418plusmnO58 3848plusmnO19 4060plusmnO71

dl IlIlml~ 5 33A6plusmnO69 3700plusmnO96 4042plusmn 109 4202plusmn086

~l1l1q 6U 3706plusmnO68 3978plusmnO83 4174plusmn047 4332plusmn103

lrimJ11l11lJfJl11l~ VlJl~mIItl~i1fl1l1WWU~OIl1I fm 11110 l~lJl~VU11lUfl nll 1$I~U

fmllfl1I~1f1l~U 111~ltl~ I 1I1fJihh~~I~Umlyj9itlUf1~llJfiW~~Ii1Il~lfl~lmtl1UUU~11Illlfl1

Vlll 0 tl~lh)lQ~~ri1fim 1lJlttf~11lJll 6 ~ ~~I11l11tlUmllJ1111liiV~IIl~~Wm 1~1 W Ji1~~~tlU fiU11 ~tllJtlW bull

I~m~I1y(uhhjUllfl~ HffUVll~l1ll$i (pgtO 05) nu~n~ihh1l11lJn11ll V~lJtl ~f1H1l ~ii

- ~Yl I ~hiJtllQ~1~ 1 3U ~~mllJft1I1iiWlJtm~umwllJtl~~l~t~lJtiV~~lflm~ 2 ~tlU lin I tl ovhou 0Q25 lin 0023 ~llmh~u

- ~~ 2 ~liJtl1q~~ 2 - 4 iJ ~~m1lJ1l1iiUIItl~tiWm1lilltl~~1~1~1lifJ~i)1f161q 1 lin 2

1I1~tiu 0013 11lt1 0018 ~l1mll~u bull ~Q d amp W v J

- fJn 3 lIllJ1l1q~H 3 - 5 11 lf~1l11lJmll)jUIItl~ttuml1lllNlllnlnln~~~1f1mq 2 110 3

lInilou 00 II 11ft 0015 IIllJih~u t I d Q ~ s centI Y I J J

- till 4 llllJ61q~~~ 4 - 6 11 li~mllJl11~lfillltl4milml1~lJtl~~1l1I~lJIM~~1f1tll~ 3 tin 4 bull 1I11ilu 0032 un 0038 ~llJhfiu

bulld ec d ltvp tdQbull - t]l1 5 ~l)jlJlqn~ 5 - 7 11 lf~fl11)jil~)jUIIl~mIJmllllJtl~I~1l1mlftIH~lfltl1~ 4 Ull~ 5

lI1111tlu 0008 un 0009 ~l)jthru d bull Q 4 4 ltII V I J ~ e1

- ~ll 6 HlnlJm~~H 6 - 8 11 li~mllJ(ll~)jUIItl~ll1Wml1llltl~I~ll1t~lJIM~~lmJ1q 5 11lt1 6

UVlltlU 0005 Ult1~ 0005 IIllJh~u

j )

bullbull

9

50

--tr--- 011J 2 ~Ol

-e-- 01 I iJ

-b- fJ1 lJ2U

~f)llJ)ll

~01j4iI

-e-01QSl

--f----- Oli 6 iI

Omiddot~middot------ bull

o 2 345 6 7 8 9

bull bull bull v

dJllilmlilm81WI~l~1ju1JWmn I 2 Ill 311l~1Yhlilm11WiWvhlJJiWIi1J~)1f1Qfl bull d q jJ 4J ( ltV

llIllJ1Q 2 IlllJW lJmllJfI1NmlllJ41~M 1356plusmnOS3 1846plusmn038 U1I 2416plusmn023 illWllllJlIl lIllJ1IlllU

iW~I~lJ)lmh811J 1 11 ilmllJnlwrj~w~m~(I 1900=1 57 2274plusmn255 Ill 2636plusmn234 IIW~illJlIl

lIllJeh9iU iW~I~lJillmlil)lQ 2 11 ilmllJnlHfllIllWil~(I 2398plusmn154 2870plusmn222 lI1I 3110plusmn139

IIli~llJlll flllHh~U iW~t~lJ)lmillJl~ 311 jjfl11lJfIJmm~m~rtNIl~1J 2672plusmnO67 3130plusmnO87 1111

3460plusmn1lO llIliAlJfll 1l1lJill~U iU~I~lJ)lmlil)lQ 4 11 ilfmlJnlHIJ)~mlllJ~~~1J 3056plusmn043

3498plusmnO48 1111 369OplusmnLl4 llfU~hlJY11 1l1lJ~1~U iW~I~lJillmh811J 5 11 ilmllJnlNmY1)~ll~l

3366plusmnO7S 3710plusmn213 lin 3SS2plusmnO73 llIUAllJlIl mlJ~l~U 1I1iu~l~lJillmilmQ 611llmllJf11H d ~ d kdJoQ

mlllJ4lllM 3666plusmnO07 3840plusmn152 un 3980plusmn168 1IIWllllJIl1 lIllHllllU (llllWYl 2) II~llJ)W1IlJlbulliJ r q$ bull ~ l toS 1 mllJml~lJIHfilIlHIi1l1l1lJal~mlf1UlJ1IIlMlm11tmmllU)llJ 211ll)U 12345678 Ut1 9 lJ

ilmllJf1lHnml)~m~1J IvhoU 854 1317 IS83 2314 2711 3084 3436 3689 3861 U1I 3980

1IIWAllJlll fillJ~~1J (m~1~-Yi 4)

10

IlTmn 2 mllJfr~Hm1 tl~t II tiUltl~tlhm61WVl111ltl tl~ltieJ~1WUtlt1tl U flllillhnUmll 311

= v tl11J11iI111lJliIU

IVll611J 2 l1tlU

= ~ ~lJliIU

854009

~ fI11lJfilHfl11tl~UW (SlJ)

ItJ 211

356plusmn053 1846038

3tJ

2416023

lltl11J IlJ 1277tO26 19OO157 2274255 2636234

lltl11J 211 1902104 2398154 2870222 3 LlOplusmnL39

ll611J 3 tJ 2166023 2672067 3130087 3460plusmnLlO

bull lh61Q 4 tJ

lltl1U 5 iJ

2664tO82

3004100

3056plusmn043

3366plusmn0078

3498plusmn048

3710213

3690plusmn1l4

3 882plusmn0 73

rbull tltJllj 611 3420plusmn157 3666plusmn070 3840plusmnL52 3980 168

~

3 U1t1Uf1IIl11lJ

JI bull JI

lhl1lflIi1mti(lltl~lhfljtl1UiUIIlt1-iW njmtiv~ltlU11lt11 12 1I11 3 il ViuhhmtJlUbull I d V At I ltI bull ltII

~UYltlUvlW1)lf)~nliijltllQ 2 ~tl1J lJW111UflHUnY11fl1J 3390Qplusmn5527 1OlOOOplusmn9247 U~ y

2364QOplusmn15323 flilJ IiIllJih~1J itl1111lJ~1fll61q 1 tJ ihlll11flllU 9520025568 172400plusmn74L13 bull bull JI

11lt1 275000plusmn 75911 flilJ gtlllJib~1J ~tl~it1lJ~lflll611J 2 11 ihl1111flIllilU 208800plusmn418 bull bull II

328600plusmn56478 11lt1 461O0069138 flilJ Vl1lJih~1J 1tllit1lJIl1fl1ltllQ 3 11 ihhlnf1IltilJlvhii1J 316000plusmn15L66 472000plusmn42071 Ult1 65300061090 oilJ 11I1lJihAll itlnI1lJ~1061Q 4 tJ ihlll1lfl

bull bull bull II

Illilt 5020QO22804 7060QO25681 1111 834025LOO oilJ ~l1mi1~ll itliiilIJ~lfltl1~ 5lJ ihhl11tl

Illat 71000024495 8756QO53393 11ft 9820QOI9235 oilJ VlllJli1~1lImitl~I~IJ~lfl611J 611 jj u J 0 d d

11lll11OlllftV 8780QOI4832 9540QO14748 IIn~ 1O52400plusmn19269 flllJ ~llHn~1J (~ln~I1 3) S4

ljtllillltJ1Jllltflltl~Lh~ih~llQnhfilllJllll~Vlfllmiddotht~i1tmtJ1WtJ11J 2 I~tltl lk 234567 8 Ht 9 tJ iJ 0)

mltlOl~llV L111111J 928034570 96733 191400 305400 476450 688100 862533 968000

1m 1052400 niIJ IiIllJthvi1J (RUHn 4)

11

bull ct bull Illt11lf1lIH1V (flnl)

illqllh1lJ~Il

mill~ 2 Iililll

l~hfl1fJ 1iI iIlYllmq 2

IlilillV 3 iIbull Ililill~ 4 iIbull IliltllQ 5V bullllil1mfj 6 1J

bull ~~

l~lJ~Il

928plusmn228

3524j477

9400plusmn19494

14800plusmn5701

30200plusmn16047

47080plusmn13609

68340plusmn4336

Iii 2i1

3390plusmn5527 lOIO0plusmn9247

9520plusmn25568 I 7240plusmn741 13

20880j41118 32860plusmn56478

31600plusmn15166 4nooplusmn42071

50200plusmn22804 70600plusmn25681

7lOooplusmn24495 87560plusmn53393

87800plusmn14832 95400plusmn147AS

3i1

23640plusmn15323

27500i75911

46100plusmn69138

65300plusmn61090

83400plusmn25100

98200plusmn19235

105240plusmn19269

IINtl1Q 9 ti

IV 2V 3V

4i1

bull 5V

6i1

7i1

8i1

9V

851plusmnoo2

1378plusmnO57

1961plusmnO53

2461plusmnl07

2927plusmnO77

3367plusmnO36

3756plusmnO69

4027plusmnO43

4188plusmnO20

4332plusmnI03

854plusmnO09

1317plusmnO56

1883plusmnO32

23 14plusmn1l7

271lplusmnl07

3084plusmnOA3

3436plusmnO56

3689plusmnO22

3861plusmnO30

3980plusmn168

9280plusmn228

34570plusmn948

96733plusmn3743

191400plusmn39040

305400plusmn22995

476450plusmn17732

4688100plusmn26167

862533plusmn24740

968000plusmn19799

1O52400plusmn19269

bull bull

bull bull

12

~1f1mlflf11nii~fl11llnll~u1i111middoth~lilMrf1Utlfl111Jjnmfla~Ua~l~hmtJ1Ugt1~U~alfJ

21~aU~uii~tJ1fJ 91J (ltl~ 2) lifilml1Jnll~Il1~ff)JilIl log W = 30309 log L - 094082

309= 01146 e0

nFlnlltl1a~igt11nlHIfl 09936

~1l11iftmiddothn1Jf111fl111Jnll~u1i~fill1m1fnfilnlltl1n~ ihhM Ufl~~lllf1 unfl~tIf bull

I~U11fflalJtlfl11lJnlJ~Ir1111h~1l1Mrmltlfl11lllnmflNUa~19hmfl1Uilfl11lJNUmhll1llf1U

q~lllmMmiddotb~middotmtl1~If1ar11lll11f11fla~Utlfi1mhfllllM1all~i1rf1911IOhmtJ1U Utl1l1f1f111l1fll

41 P I _ i P Ql Qb llAl t1Iflj~ 3 II~alllllul1I1lf111 1I1111l1mal1lJP1111l1t]1IU flllUU Isometric IIl1J~qlltlf111

11l1t]IiltlU 1Iluuu~iilNf1111~h IIlUN von Bertalanffy (1938)

~ laquoJGJr IV Jl 04$

fl11lJffll)I7iM11~I1llitllmfl111JmHmlfltJ~a~lnmru~Ugt1middotm9lalfJ 2 Illalilll~

811 9 iI (1t1~ 3) ilfilfl111lntl~u1ili~U1Jf111 log W = 32348 log w - 114206

maW = 00721 ~2348

= 09900

ltIf I

fl11lJn1JltU1il1t1Hft111Jf11Hfllfta~IIi1fl111Jjnfl1fta~Ila~lftlmalUIl~UflllfJ 2

l~aUlluii~tJl~ 9iI (ltl~ 4) iifhfll1JuimiuiRunJf111 I-

w = 08781L+15116

= 09859

bull bull bullbull

bull bull

bull bullbull

13

14000

12000

0000 ~ S aOOOs

p

) 6000 000

2003

0

W=01146L303W

R = 09936

10 5

bull

161OIl

14001 i

1 tX)Il J~~ oS 0000 I shy

lImlD ~sect 6006 F

I

000 ZlOOO

bull

lmi 3 trn1Jmnt1iHmiddothnilH1imillIftf1Trnfl~1~flif16~lfhfliztll1nnfJ 21iillu ii~ 911

bull 50-

i w=O8781l + 15116 ~ ~bull

R =09859Iii 30

~ ~

F 10

bull

14

Ao~- J d4-Sf11 I

IlJij U1 ij lJllfl111l tJ l1UUW m Ilij~IIij ~ tllllllillll ll1l1111fllfl111l tJl111ij~m1il f) ~ VIIIllJII ll ll

hmtJt1111 I il i~~fllJllllllllllm~ 5

1l11H~S fllmlf1l~lJ IIlllHIilmfl1ll

fll11lO1111liiuhl fllllh~lJINfl111JUn fllfl11111J1111l ilU

16lJil(SII (SII) (SII)

lUlfl161q 2Ifl)U 1~lIl1le~ 850 568 1134

bull iI~ 1 1418 482 1659

1J~ 2 1900 670 2235

~ ij~ 3 2570

~Ih)lQ 111 111J11~aS~ 1337 657 16655

UdY1 1~i4 384 186

il~) 23]8 438 2597

tl~ 3 2816

iJIli1~ 2 f 11lllfilliH J990 544 2262

r~Yl 2534 434 27SJ

1l~2 2968 372 3154

1J~3 3340

~1qll 11I11I1IHl~ 2360 628 2674

11II 1 2988 374 3175

l~2 3362 408 3566

Iii 3 3770

illlfi181q 4 l r ~1lYI1I1I8~ 2934 484 3176

bull j~ 1 3418 430 3633

iI~2 3848 212 6 3954

bull iI~3 4060

llUrilllq 51 1~IIIIilll6~ 3346 354 3523

1~ 1 3700 342 3871

11~2 4042 160 4122

iJ~ 3 4202

ilmilJ1q 6 iJ lIIIIIIll~ 3706 272 3amp42

iJ~1 3978 196 4076

11~ 2 4174 158 4253

iI~ 3 4332

15

lll~thlJl1IflnllI11Wl11iJLVltlIJtl~fl111~hlIVlnlJuuuhltl~fl1~IRUIIlIJN von Bertalanffy

(1938) flVfh L UIl K i1tl11i Gulland and Holt plot (1959 tyen1~911lJ Sparre and Venema 1992)

~fflJ fl1 ~

~Wuh L 598 lI1J91UJ~11 lllzfh K = 0138 ~vjJ liI~lflfh to = -0216 jJ (fljJnUil1~~WUl1

2545) ihlllfl1Jfl11fl1111l~tyL9ill lVlIJtl~ l~lmZtllUlflll~ lUlllUlJflli fl11lJfflJliutf 1zlh~mj (t) fill

fl11Utll1 (L) fitl

Lt = 598(1 e)--OmiddotI38(I+O216)

- ~Ilti R2 =067bull - +

L--middot~--~ -~ ~ ~-~ o I I ) Lt

o 10 20 30 40 50 60 70

3tl~ 5 urntn411ftltil111Jlfmiutflllfltllfl9Im~~ 51il611iIJtl~ Outland and Holt (1959 ~l~llllJ Spam and Venema 1992)

bull fl111J0l1mflfl~lftlfmf11Uii1flfl1~IIYluthlUff1Jfl1iIJH von Bertalanffy rlvllhfiVlj 1 - 60 tl ulutamph1I1)tlUli1Ufl~ifll11Jilniui~1i~fl11lJ[Jl1flilltl~mlZfllj1iI~UUVUlli~ IIl[JV9111fll ~ ~ 1 Jt ~ -tLIlitylllll 111 uIlIIJamp~fl1iIWUfl11U011fllflfl~flVUlJHfHYlIIJVUV1j 20 1J 1111 111 (l1JYl 6)

bull bull

bull bull

16

80~

i ~

60 G r ro 40 ri ~

rshyro

20

0

o 10 20 30 40 50 60

in~h (ill

amptlllfl~6~iv~t9ilfl1iltllUlL~tI~lU 1Iumll 3 iJ ~itlliW~fl11VlYltltl~ lhlflDhwh Aw _IltldflfI1UTUIJfI1UtI8flSnaflflltJ 100 L1Ja11liU~

t J bull I _4fUIfI11IUn1IHfI1iYliltla~laV~f11fliilalU 1IuSWrltll 3 iJ VlUllflIll11fJlJU1IJfl1

ilImiddoth~ lSG-no 84fru1JtU1io flllJI1ltflSfll1lUh~lJamiddotnhiiti1iHh~ 75-82 1l1IJ1Illtltlfl~I~Ufi

annIl1U1hiitin1t1H 62-7_6 Utliiflntiaiifll ml1l11lt~WJamiddotnhtil11middoth~ 750-1754 ilaaflfmia bull

afll f1lJfl1~HlJtl~1hiltin1Imiddoth4 862-168Ii1aafl111rianfll Uafn1mll 1fltltlflliIi~itl1dltil

bull

bull bull

bull bull

17

f111~~1la~iiu~t~hmttl1ttl1qh~ l fit~1t1t 7 it l 11t 5 ~ 11~tiUhl1j6fl6tflil

IJUl~ 50 9l1il~IIJf1i ~~iifl11~11UlI11lt1mlflv~titlfl1ill~116~iv~lfhmtlmaii1I1qJl (~U~91 2528)

191Vuii11titHIlfll911~iimqllhfilIl1lti1~ftl1l11 III flfl 1fllllhlluimn91mq 2 IflflU uhmq 111211

~Ua~ 611 hJnldjllfftl11lJfltllin~~ nll(hu 1 I 191vll111umlhltlT11i lJfllU l n I fll~ 1Ui9l11 10 $I ~ bull bull

11IflfU~Ufl~I1111unfil dl11itIlmillimfl91illiMln6tlfl~lfljihm~Vl1Ufl~fli~6~I~iiv 654plusmn003 1IIUiium1 flI1IJn11~m91MI~nV 666plusmnO05 lft~IlJili 1I11Ul11Um~nvlvhtitl 526IH267 flllJ 1m

Iflv~fimlv~1U16~UV1Iinrumiltl (2544) IIn11fJ~1tIItJ~~1tlll1n111tlUIl (2545) ~~llfll9hiJ~ii

il11lJVmllltmnhj~UIilVtitl6TI11i flmlllJlliiJ~i1lJrtlli1Yijjlt191111 ltyflhvhl1yeniilltl1111tlltl~UtH 4 ltl q~ bull - YI w~ bull

i11ifliltll111~ Il~ 11lll1fl11Nn m1fl1911flttll111nlIllUfl11~ftifJ~lIJtIllJl~mtlalI11~ trflflUlllU11l1 2

lillatl ii~IlllJl~ilfttl~ 1pl1milhm~ 2 flaUiiIJU1f)mllJm1ni91fl~ ml1Jfl~lmiddotmillfl~ IfII1111Um~nl

1~1JJiU nhtiU 851plusmn0021lfURIlJili 854t0091lfU~1JIi La 9280t228 fill mlJhiiu thullljiIJiU

~ a I dill ~ 41 d ~ i ~I bull1)U 1 fltlllflfllllflllfllmllI~1~lnflmiddotI~II1tIiIJljllffmlltl1lJ~U1~91J~lnffliltl 1J1J1l11l l ~l 1t1lIIlm~

~rI dl 1 Jjl Jq_II1W~ lVlfft11Ul ~l 111)1111ill ~ 1Ifl1i111l~~~tl~~WlllUlJ~~1t)~Ulll

J ~ J bull r i I 4 i I ~ I bull A fmllll1lfl~fI1~U ~II1lirnllflfm~mlflllJi1lJf1t IlHlllltl1 lJIWmI1V~IiJUltl~ I 1IJ lWfl~

~lfl~fl~fl1111yenflmhl~n~IIfl~I911mmUflalvfla~titlnS1IlI1PlJlfl~q~ l~mll4imJ~fl1mltJjlt1U1~

U6~11i1iii Ii~nfln11~9l1ltl~ih1IwlitljfllmVtlliiJtl111fl1fIItHIIi1Viit~Illflli1111n9 ffTl1fttl11liiiili itfl1illf)llti~1~1I n1lilfftlhlltitlfiml~ ffilfflU 11 lil(Ji111Ufl lUtillllflViUhl~i1iU1ill1U1t1lffltlfltl

bull y

61H11111fl11 (ff1aft 2506 thlii 2529) ulil~hilliil~Hua=iifl11flfliHfltl61H11ll11J11flflUilin4II

U1lZId61I111 (Vihnflla1io 2525 ~fllC 2529) li1utl1H11~ihMlfilflitllUlftt1~i~~ qfl~88l1f1nlallfuiu~ ftlhu lllI1i1HUfl iU6~111Wll 10 11I6flo1U~IItl~t1111iiflIgti1

- 1 (tplJllllafllC2533 IRnflIUlUIftfIt2541) ~ vitl p

fll1nflllltnllJtflJlnl1i111~UlHUflll1l1 fl111J I11n itiltlll II fl ~1lllfl1tJ1Ui 11 ff~fl1i fl1111

tI 3030) A f Ai pound (l 4 1d I I GO

alJIItli W = 01146 L lIlIff1Jfl1iTURlJfl1f(lIlJltlYllimfl111W~Ya~lJlfl 9l~il1lmflli

09936 bull

11tf~~ -)1ItlIltUtl Ill fl1111 n lJIiU S 11-l1~t11 HUflllll fl11lJl11 m 11 tl ~ Iii 1il fl111J NU 11111 IIllJ lit tllllJ1 fl

~ i1I-l1~~11111i~Iiitlil111lfJl1m1I611 1I1lilu1l1f111JflmiTI1Ufl 1lJtllJfl1ifl1111IDJiUliiftuUu~

T I d 1 Y I v a tlflfl11UtlJ1lU111tfl9111lflI1lJlltllllJl911lJIItlIlUfl~il11lJl11n191tllllillfl~itll1l 9936 lf~tlllfl11fll1

~mnUH~1tllU1l111tlUI1 (2545) ~ilifilIDJ1IlffYi~ItiWItfIII6~1~1fl1fllt1flfllltiiUi~ilmllJ ~

1l1l1nlill1Il11t1111lliil1lHVifffllil1l1lJ~1l1ii~i~11191fftl1l 1lllllflWhhI1hfitI 08659 UfIIIIflWfJh

fit 05019 I~ilfhiflalfllllfltUtl~l~lmU1l1~111qJl~ltfv~itlUtlfl6t1fll91IJUiV1111122 IfltlU ~~Ii filnlJ1Iliilli9i1t)11t1f1l1htiu 09968 1I1l 09895 ~llJihAtI (~IJ1~ 2528)

I

14

Ao~- J d4-Sf11 I

IlJij U1 ij lJllfl111l tJ l1UUW m Ilij~IIij ~ tllllllillll ll1l1111fllfl111l tJl111ij~m1il f) ~ VIIIllJII ll ll

hmtJt1111 I il i~~fllJllllllllllm~ 5

1l11H~S fllmlf1l~lJ IIlllHIilmfl1ll

fll11lO1111liiuhl fllllh~lJINfl111JUn fllfl11111J1111l ilU

16lJil(SII (SII) (SII)

lUlfl161q 2Ifl)U 1~lIl1le~ 850 568 1134

bull iI~ 1 1418 482 1659

1J~ 2 1900 670 2235

~ ij~ 3 2570

~Ih)lQ 111 111J11~aS~ 1337 657 16655

UdY1 1~i4 384 186

il~) 23]8 438 2597

tl~ 3 2816

iJIli1~ 2 f 11lllfilliH J990 544 2262

r~Yl 2534 434 27SJ

1l~2 2968 372 3154

1J~3 3340

~1qll 11I11I1IHl~ 2360 628 2674

11II 1 2988 374 3175

l~2 3362 408 3566

Iii 3 3770

illlfi181q 4 l r ~1lYI1I1I8~ 2934 484 3176

bull j~ 1 3418 430 3633

iI~2 3848 212 6 3954

bull iI~3 4060

llUrilllq 51 1~IIIIilll6~ 3346 354 3523

1~ 1 3700 342 3871

11~2 4042 160 4122

iJ~ 3 4202

ilmilJ1q 6 iJ lIIIIIIll~ 3706 272 3amp42

iJ~1 3978 196 4076

11~ 2 4174 158 4253

iI~ 3 4332

15

lll~thlJl1IflnllI11Wl11iJLVltlIJtl~fl111~hlIVlnlJuuuhltl~fl1~IRUIIlIJN von Bertalanffy

(1938) flVfh L UIl K i1tl11i Gulland and Holt plot (1959 tyen1~911lJ Sparre and Venema 1992)

~fflJ fl1 ~

~Wuh L 598 lI1J91UJ~11 lllzfh K = 0138 ~vjJ liI~lflfh to = -0216 jJ (fljJnUil1~~WUl1

2545) ihlllfl1Jfl11fl1111l~tyL9ill lVlIJtl~ l~lmZtllUlflll~ lUlllUlJflli fl11lJfflJliutf 1zlh~mj (t) fill

fl11Utll1 (L) fitl

Lt = 598(1 e)--OmiddotI38(I+O216)

- ~Ilti R2 =067bull - +

L--middot~--~ -~ ~ ~-~ o I I ) Lt

o 10 20 30 40 50 60 70

3tl~ 5 urntn411ftltil111Jlfmiutflllfltllfl9Im~~ 51il611iIJtl~ Outland and Holt (1959 ~l~llllJ Spam and Venema 1992)

bull fl111J0l1mflfl~lftlfmf11Uii1flfl1~IIYluthlUff1Jfl1iIJH von Bertalanffy rlvllhfiVlj 1 - 60 tl ulutamph1I1)tlUli1Ufl~ifll11Jilniui~1i~fl11lJ[Jl1flilltl~mlZfllj1iI~UUVUlli~ IIl[JV9111fll ~ ~ 1 Jt ~ -tLIlitylllll 111 uIlIIJamp~fl1iIWUfl11U011fllflfl~flVUlJHfHYlIIJVUV1j 20 1J 1111 111 (l1JYl 6)

bull bull

bull bull

16

80~

i ~

60 G r ro 40 ri ~

rshyro

20

0

o 10 20 30 40 50 60

in~h (ill

amptlllfl~6~iv~t9ilfl1iltllUlL~tI~lU 1Iumll 3 iJ ~itlliW~fl11VlYltltl~ lhlflDhwh Aw _IltldflfI1UTUIJfI1UtI8flSnaflflltJ 100 L1Ja11liU~

t J bull I _4fUIfI11IUn1IHfI1iYliltla~laV~f11fliilalU 1IuSWrltll 3 iJ VlUllflIll11fJlJU1IJfl1

ilImiddoth~ lSG-no 84fru1JtU1io flllJI1ltflSfll1lUh~lJamiddotnhiiti1iHh~ 75-82 1l1IJ1Illtltlfl~I~Ufi

annIl1U1hiitin1t1H 62-7_6 Utliiflntiaiifll ml1l11lt~WJamiddotnhtil11middoth~ 750-1754 ilaaflfmia bull

afll f1lJfl1~HlJtl~1hiltin1Imiddoth4 862-168Ii1aafl111rianfll Uafn1mll 1fltltlflliIi~itl1dltil

bull

bull bull

bull bull

17

f111~~1la~iiu~t~hmttl1ttl1qh~ l fit~1t1t 7 it l 11t 5 ~ 11~tiUhl1j6fl6tflil

IJUl~ 50 9l1il~IIJf1i ~~iifl11~11UlI11lt1mlflv~titlfl1ill~116~iv~lfhmtlmaii1I1qJl (~U~91 2528)

191Vuii11titHIlfll911~iimqllhfilIl1lti1~ftl1l11 III flfl 1fllllhlluimn91mq 2 IflflU uhmq 111211

~Ua~ 611 hJnldjllfftl11lJfltllin~~ nll(hu 1 I 191vll111umlhltlT11i lJfllU l n I fll~ 1Ui9l11 10 $I ~ bull bull

11IflfU~Ufl~I1111unfil dl11itIlmillimfl91illiMln6tlfl~lfljihm~Vl1Ufl~fli~6~I~iiv 654plusmn003 1IIUiium1 flI1IJn11~m91MI~nV 666plusmnO05 lft~IlJili 1I11Ul11Um~nvlvhtitl 526IH267 flllJ 1m

Iflv~fimlv~1U16~UV1Iinrumiltl (2544) IIn11fJ~1tIItJ~~1tlll1n111tlUIl (2545) ~~llfll9hiJ~ii

il11lJVmllltmnhj~UIilVtitl6TI11i flmlllJlliiJ~i1lJrtlli1Yijjlt191111 ltyflhvhl1yeniilltl1111tlltl~UtH 4 ltl q~ bull - YI w~ bull

i11ifliltll111~ Il~ 11lll1fl11Nn m1fl1911flttll111nlIllUfl11~ftifJ~lIJtIllJl~mtlalI11~ trflflUlllU11l1 2

lillatl ii~IlllJl~ilfttl~ 1pl1milhm~ 2 flaUiiIJU1f)mllJm1ni91fl~ ml1Jfl~lmiddotmillfl~ IfII1111Um~nl

1~1JJiU nhtiU 851plusmn0021lfURIlJili 854t0091lfU~1JIi La 9280t228 fill mlJhiiu thullljiIJiU

~ a I dill ~ 41 d ~ i ~I bull1)U 1 fltlllflfllllflllfllmllI~1~lnflmiddotI~II1tIiIJljllffmlltl1lJ~U1~91J~lnffliltl 1J1J1l11l l ~l 1t1lIIlm~

~rI dl 1 Jjl Jq_II1W~ lVlfft11Ul ~l 111)1111ill ~ 1Ifl1i111l~~~tl~~WlllUlJ~~1t)~Ulll

J ~ J bull r i I 4 i I ~ I bull A fmllll1lfl~fI1~U ~II1lirnllflfm~mlflllJi1lJf1t IlHlllltl1 lJIWmI1V~IiJUltl~ I 1IJ lWfl~

~lfl~fl~fl1111yenflmhl~n~IIfl~I911mmUflalvfla~titlnS1IlI1PlJlfl~q~ l~mll4imJ~fl1mltJjlt1U1~

U6~11i1iii Ii~nfln11~9l1ltl~ih1IwlitljfllmVtlliiJtl111fl1fIItHIIi1Viit~Illflli1111n9 ffTl1fttl11liiiili itfl1illf)llti~1~1I n1lilfftlhlltitlfiml~ ffilfflU 11 lil(Ji111Ufl lUtillllflViUhl~i1iU1ill1U1t1lffltlfltl

bull y

61H11111fl11 (ff1aft 2506 thlii 2529) ulil~hilliil~Hua=iifl11flfliHfltl61H11ll11J11flflUilin4II

U1lZId61I111 (Vihnflla1io 2525 ~fllC 2529) li1utl1H11~ihMlfilflitllUlftt1~i~~ qfl~88l1f1nlallfuiu~ ftlhu lllI1i1HUfl iU6~111Wll 10 11I6flo1U~IItl~t1111iiflIgti1

- 1 (tplJllllafllC2533 IRnflIUlUIftfIt2541) ~ vitl p

fll1nflllltnllJtflJlnl1i111~UlHUflll1l1 fl111J I11n itiltlll II fl ~1lllfl1tJ1Ui 11 ff~fl1i fl1111

tI 3030) A f Ai pound (l 4 1d I I GO

alJIItli W = 01146 L lIlIff1Jfl1iTURlJfl1f(lIlJltlYllimfl111W~Ya~lJlfl 9l~il1lmflli

09936 bull

11tf~~ -)1ItlIltUtl Ill fl1111 n lJIiU S 11-l1~t11 HUflllll fl11lJl11 m 11 tl ~ Iii 1il fl111J NU 11111 IIllJ lit tllllJ1 fl

~ i1I-l1~~11111i~Iiitlil111lfJl1m1I611 1I1lilu1l1f111JflmiTI1Ufl 1lJtllJfl1ifl1111IDJiUliiftuUu~

T I d 1 Y I v a tlflfl11UtlJ1lU111tfl9111lflI1lJlltllllJl911lJIItlIlUfl~il11lJl11n191tllllillfl~itll1l 9936 lf~tlllfl11fll1

~mnUH~1tllU1l111tlUI1 (2545) ~ilifilIDJ1IlffYi~ItiWItfIII6~1~1fl1fllt1flfllltiiUi~ilmllJ ~

1l1l1nlill1Il11t1111lliil1lHVifffllil1l1lJ~1l1ii~i~11191fftl1l 1lllllflWhhI1hfitI 08659 UfIIIIflWfJh

fit 05019 I~ilfhiflalfllllfltUtl~l~lmU1l1~111qJl~ltfv~itlUtlfl6t1fll91IJUiV1111122 IfltlU ~~Ii filnlJ1Iliilli9i1t)11t1f1l1htiu 09968 1I1l 09895 ~llJihAtI (~IJ1~ 2528)

I

15

lll~thlJl1IflnllI11Wl11iJLVltlIJtl~fl111~hlIVlnlJuuuhltl~fl1~IRUIIlIJN von Bertalanffy

(1938) flVfh L UIl K i1tl11i Gulland and Holt plot (1959 tyen1~911lJ Sparre and Venema 1992)

~fflJ fl1 ~

~Wuh L 598 lI1J91UJ~11 lllzfh K = 0138 ~vjJ liI~lflfh to = -0216 jJ (fljJnUil1~~WUl1

2545) ihlllfl1Jfl11fl1111l~tyL9ill lVlIJtl~ l~lmZtllUlflll~ lUlllUlJflli fl11lJfflJliutf 1zlh~mj (t) fill

fl11Utll1 (L) fitl

Lt = 598(1 e)--OmiddotI38(I+O216)

- ~Ilti R2 =067bull - +

L--middot~--~ -~ ~ ~-~ o I I ) Lt

o 10 20 30 40 50 60 70

3tl~ 5 urntn411ftltil111Jlfmiutflllfltllfl9Im~~ 51il611iIJtl~ Outland and Holt (1959 ~l~llllJ Spam and Venema 1992)

bull fl111J0l1mflfl~lftlfmf11Uii1flfl1~IIYluthlUff1Jfl1iIJH von Bertalanffy rlvllhfiVlj 1 - 60 tl ulutamph1I1)tlUli1Ufl~ifll11Jilniui~1i~fl11lJ[Jl1flilltl~mlZfllj1iI~UUVUlli~ IIl[JV9111fll ~ ~ 1 Jt ~ -tLIlitylllll 111 uIlIIJamp~fl1iIWUfl11U011fllflfl~flVUlJHfHYlIIJVUV1j 20 1J 1111 111 (l1JYl 6)

bull bull

bull bull

16

80~

i ~

60 G r ro 40 ri ~

rshyro

20

0

o 10 20 30 40 50 60

in~h (ill

amptlllfl~6~iv~t9ilfl1iltllUlL~tI~lU 1Iumll 3 iJ ~itlliW~fl11VlYltltl~ lhlflDhwh Aw _IltldflfI1UTUIJfI1UtI8flSnaflflltJ 100 L1Ja11liU~

t J bull I _4fUIfI11IUn1IHfI1iYliltla~laV~f11fliilalU 1IuSWrltll 3 iJ VlUllflIll11fJlJU1IJfl1

ilImiddoth~ lSG-no 84fru1JtU1io flllJI1ltflSfll1lUh~lJamiddotnhiiti1iHh~ 75-82 1l1IJ1Illtltlfl~I~Ufi

annIl1U1hiitin1t1H 62-7_6 Utliiflntiaiifll ml1l11lt~WJamiddotnhtil11middoth~ 750-1754 ilaaflfmia bull

afll f1lJfl1~HlJtl~1hiltin1Imiddoth4 862-168Ii1aafl111rianfll Uafn1mll 1fltltlflliIi~itl1dltil

bull

bull bull

bull bull

17

f111~~1la~iiu~t~hmttl1ttl1qh~ l fit~1t1t 7 it l 11t 5 ~ 11~tiUhl1j6fl6tflil

IJUl~ 50 9l1il~IIJf1i ~~iifl11~11UlI11lt1mlflv~titlfl1ill~116~iv~lfhmtlmaii1I1qJl (~U~91 2528)

191Vuii11titHIlfll911~iimqllhfilIl1lti1~ftl1l11 III flfl 1fllllhlluimn91mq 2 IflflU uhmq 111211

~Ua~ 611 hJnldjllfftl11lJfltllin~~ nll(hu 1 I 191vll111umlhltlT11i lJfllU l n I fll~ 1Ui9l11 10 $I ~ bull bull

11IflfU~Ufl~I1111unfil dl11itIlmillimfl91illiMln6tlfl~lfljihm~Vl1Ufl~fli~6~I~iiv 654plusmn003 1IIUiium1 flI1IJn11~m91MI~nV 666plusmnO05 lft~IlJili 1I11Ul11Um~nvlvhtitl 526IH267 flllJ 1m

Iflv~fimlv~1U16~UV1Iinrumiltl (2544) IIn11fJ~1tIItJ~~1tlll1n111tlUIl (2545) ~~llfll9hiJ~ii

il11lJVmllltmnhj~UIilVtitl6TI11i flmlllJlliiJ~i1lJrtlli1Yijjlt191111 ltyflhvhl1yeniilltl1111tlltl~UtH 4 ltl q~ bull - YI w~ bull

i11ifliltll111~ Il~ 11lll1fl11Nn m1fl1911flttll111nlIllUfl11~ftifJ~lIJtIllJl~mtlalI11~ trflflUlllU11l1 2

lillatl ii~IlllJl~ilfttl~ 1pl1milhm~ 2 flaUiiIJU1f)mllJm1ni91fl~ ml1Jfl~lmiddotmillfl~ IfII1111Um~nl

1~1JJiU nhtiU 851plusmn0021lfURIlJili 854t0091lfU~1JIi La 9280t228 fill mlJhiiu thullljiIJiU

~ a I dill ~ 41 d ~ i ~I bull1)U 1 fltlllflfllllflllfllmllI~1~lnflmiddotI~II1tIiIJljllffmlltl1lJ~U1~91J~lnffliltl 1J1J1l11l l ~l 1t1lIIlm~

~rI dl 1 Jjl Jq_II1W~ lVlfft11Ul ~l 111)1111ill ~ 1Ifl1i111l~~~tl~~WlllUlJ~~1t)~Ulll

J ~ J bull r i I 4 i I ~ I bull A fmllll1lfl~fI1~U ~II1lirnllflfm~mlflllJi1lJf1t IlHlllltl1 lJIWmI1V~IiJUltl~ I 1IJ lWfl~

~lfl~fl~fl1111yenflmhl~n~IIfl~I911mmUflalvfla~titlnS1IlI1PlJlfl~q~ l~mll4imJ~fl1mltJjlt1U1~

U6~11i1iii Ii~nfln11~9l1ltl~ih1IwlitljfllmVtlliiJtl111fl1fIItHIIi1Viit~Illflli1111n9 ffTl1fttl11liiiili itfl1illf)llti~1~1I n1lilfftlhlltitlfiml~ ffilfflU 11 lil(Ji111Ufl lUtillllflViUhl~i1iU1ill1U1t1lffltlfltl

bull y

61H11111fl11 (ff1aft 2506 thlii 2529) ulil~hilliil~Hua=iifl11flfliHfltl61H11ll11J11flflUilin4II

U1lZId61I111 (Vihnflla1io 2525 ~fllC 2529) li1utl1H11~ihMlfilflitllUlftt1~i~~ qfl~88l1f1nlallfuiu~ ftlhu lllI1i1HUfl iU6~111Wll 10 11I6flo1U~IItl~t1111iiflIgti1

- 1 (tplJllllafllC2533 IRnflIUlUIftfIt2541) ~ vitl p

fll1nflllltnllJtflJlnl1i111~UlHUflll1l1 fl111J I11n itiltlll II fl ~1lllfl1tJ1Ui 11 ff~fl1i fl1111

tI 3030) A f Ai pound (l 4 1d I I GO

alJIItli W = 01146 L lIlIff1Jfl1iTURlJfl1f(lIlJltlYllimfl111W~Ya~lJlfl 9l~il1lmflli

09936 bull

11tf~~ -)1ItlIltUtl Ill fl1111 n lJIiU S 11-l1~t11 HUflllll fl11lJl11 m 11 tl ~ Iii 1il fl111J NU 11111 IIllJ lit tllllJ1 fl

~ i1I-l1~~11111i~Iiitlil111lfJl1m1I611 1I1lilu1l1f111JflmiTI1Ufl 1lJtllJfl1ifl1111IDJiUliiftuUu~

T I d 1 Y I v a tlflfl11UtlJ1lU111tfl9111lflI1lJlltllllJl911lJIItlIlUfl~il11lJl11n191tllllillfl~itll1l 9936 lf~tlllfl11fll1

~mnUH~1tllU1l111tlUI1 (2545) ~ilifilIDJ1IlffYi~ItiWItfIII6~1~1fl1fllt1flfllltiiUi~ilmllJ ~

1l1l1nlill1Il11t1111lliil1lHVifffllil1l1lJ~1l1ii~i~11191fftl1l 1lllllflWhhI1hfitI 08659 UfIIIIflWfJh

fit 05019 I~ilfhiflalfllllfltUtl~l~lmU1l1~111qJl~ltfv~itlUtlfl6t1fll91IJUiV1111122 IfltlU ~~Ii filnlJ1Iliilli9i1t)11t1f1l1htiu 09968 1I1l 09895 ~llJihAtI (~IJ1~ 2528)

I

bull bull

bull bull

16

80~

i ~

60 G r ro 40 ri ~

rshyro

20

0

o 10 20 30 40 50 60

in~h (ill

amptlllfl~6~iv~t9ilfl1iltllUlL~tI~lU 1Iumll 3 iJ ~itlliW~fl11VlYltltl~ lhlflDhwh Aw _IltldflfI1UTUIJfI1UtI8flSnaflflltJ 100 L1Ja11liU~

t J bull I _4fUIfI11IUn1IHfI1iYliltla~laV~f11fliilalU 1IuSWrltll 3 iJ VlUllflIll11fJlJU1IJfl1

ilImiddoth~ lSG-no 84fru1JtU1io flllJI1ltflSfll1lUh~lJamiddotnhiiti1iHh~ 75-82 1l1IJ1Illtltlfl~I~Ufi

annIl1U1hiitin1t1H 62-7_6 Utliiflntiaiifll ml1l11lt~WJamiddotnhtil11middoth~ 750-1754 ilaaflfmia bull

afll f1lJfl1~HlJtl~1hiltin1Imiddoth4 862-168Ii1aafl111rianfll Uafn1mll 1fltltlflliIi~itl1dltil

bull

bull bull

bull bull

17

f111~~1la~iiu~t~hmttl1ttl1qh~ l fit~1t1t 7 it l 11t 5 ~ 11~tiUhl1j6fl6tflil

IJUl~ 50 9l1il~IIJf1i ~~iifl11~11UlI11lt1mlflv~titlfl1ill~116~iv~lfhmtlmaii1I1qJl (~U~91 2528)

191Vuii11titHIlfll911~iimqllhfilIl1lti1~ftl1l11 III flfl 1fllllhlluimn91mq 2 IflflU uhmq 111211

~Ua~ 611 hJnldjllfftl11lJfltllin~~ nll(hu 1 I 191vll111umlhltlT11i lJfllU l n I fll~ 1Ui9l11 10 $I ~ bull bull

11IflfU~Ufl~I1111unfil dl11itIlmillimfl91illiMln6tlfl~lfljihm~Vl1Ufl~fli~6~I~iiv 654plusmn003 1IIUiium1 flI1IJn11~m91MI~nV 666plusmnO05 lft~IlJili 1I11Ul11Um~nvlvhtitl 526IH267 flllJ 1m

Iflv~fimlv~1U16~UV1Iinrumiltl (2544) IIn11fJ~1tIItJ~~1tlll1n111tlUIl (2545) ~~llfll9hiJ~ii

il11lJVmllltmnhj~UIilVtitl6TI11i flmlllJlliiJ~i1lJrtlli1Yijjlt191111 ltyflhvhl1yeniilltl1111tlltl~UtH 4 ltl q~ bull - YI w~ bull

i11ifliltll111~ Il~ 11lll1fl11Nn m1fl1911flttll111nlIllUfl11~ftifJ~lIJtIllJl~mtlalI11~ trflflUlllU11l1 2

lillatl ii~IlllJl~ilfttl~ 1pl1milhm~ 2 flaUiiIJU1f)mllJm1ni91fl~ ml1Jfl~lmiddotmillfl~ IfII1111Um~nl

1~1JJiU nhtiU 851plusmn0021lfURIlJili 854t0091lfU~1JIi La 9280t228 fill mlJhiiu thullljiIJiU

~ a I dill ~ 41 d ~ i ~I bull1)U 1 fltlllflfllllflllfllmllI~1~lnflmiddotI~II1tIiIJljllffmlltl1lJ~U1~91J~lnffliltl 1J1J1l11l l ~l 1t1lIIlm~

~rI dl 1 Jjl Jq_II1W~ lVlfft11Ul ~l 111)1111ill ~ 1Ifl1i111l~~~tl~~WlllUlJ~~1t)~Ulll

J ~ J bull r i I 4 i I ~ I bull A fmllll1lfl~fI1~U ~II1lirnllflfm~mlflllJi1lJf1t IlHlllltl1 lJIWmI1V~IiJUltl~ I 1IJ lWfl~

~lfl~fl~fl1111yenflmhl~n~IIfl~I911mmUflalvfla~titlnS1IlI1PlJlfl~q~ l~mll4imJ~fl1mltJjlt1U1~

U6~11i1iii Ii~nfln11~9l1ltl~ih1IwlitljfllmVtlliiJtl111fl1fIItHIIi1Viit~Illflli1111n9 ffTl1fttl11liiiili itfl1illf)llti~1~1I n1lilfftlhlltitlfiml~ ffilfflU 11 lil(Ji111Ufl lUtillllflViUhl~i1iU1ill1U1t1lffltlfltl

bull y

61H11111fl11 (ff1aft 2506 thlii 2529) ulil~hilliil~Hua=iifl11flfliHfltl61H11ll11J11flflUilin4II

U1lZId61I111 (Vihnflla1io 2525 ~fllC 2529) li1utl1H11~ihMlfilflitllUlftt1~i~~ qfl~88l1f1nlallfuiu~ ftlhu lllI1i1HUfl iU6~111Wll 10 11I6flo1U~IItl~t1111iiflIgti1

- 1 (tplJllllafllC2533 IRnflIUlUIftfIt2541) ~ vitl p

fll1nflllltnllJtflJlnl1i111~UlHUflll1l1 fl111J I11n itiltlll II fl ~1lllfl1tJ1Ui 11 ff~fl1i fl1111

tI 3030) A f Ai pound (l 4 1d I I GO

alJIItli W = 01146 L lIlIff1Jfl1iTURlJfl1f(lIlJltlYllimfl111W~Ya~lJlfl 9l~il1lmflli

09936 bull

11tf~~ -)1ItlIltUtl Ill fl1111 n lJIiU S 11-l1~t11 HUflllll fl11lJl11 m 11 tl ~ Iii 1il fl111J NU 11111 IIllJ lit tllllJ1 fl

~ i1I-l1~~11111i~Iiitlil111lfJl1m1I611 1I1lilu1l1f111JflmiTI1Ufl 1lJtllJfl1ifl1111IDJiUliiftuUu~

T I d 1 Y I v a tlflfl11UtlJ1lU111tfl9111lflI1lJlltllllJl911lJIItlIlUfl~il11lJl11n191tllllillfl~itll1l 9936 lf~tlllfl11fll1

~mnUH~1tllU1l111tlUI1 (2545) ~ilifilIDJ1IlffYi~ItiWItfIII6~1~1fl1fllt1flfllltiiUi~ilmllJ ~

1l1l1nlill1Il11t1111lliil1lHVifffllil1l1lJ~1l1ii~i~11191fftl1l 1lllllflWhhI1hfitI 08659 UfIIIIflWfJh

fit 05019 I~ilfhiflalfllllfltUtl~l~lmU1l1~111qJl~ltfv~itlUtlfl6t1fll91IJUiV1111122 IfltlU ~~Ii filnlJ1Iliilli9i1t)11t1f1l1htiu 09968 1I1l 09895 ~llJihAtI (~IJ1~ 2528)

I

bull bull

bull bull

17

f111~~1la~iiu~t~hmttl1ttl1qh~ l fit~1t1t 7 it l 11t 5 ~ 11~tiUhl1j6fl6tflil

IJUl~ 50 9l1il~IIJf1i ~~iifl11~11UlI11lt1mlflv~titlfl1ill~116~iv~lfhmtlmaii1I1qJl (~U~91 2528)

191Vuii11titHIlfll911~iimqllhfilIl1lti1~ftl1l11 III flfl 1fllllhlluimn91mq 2 IflflU uhmq 111211

~Ua~ 611 hJnldjllfftl11lJfltllin~~ nll(hu 1 I 191vll111umlhltlT11i lJfllU l n I fll~ 1Ui9l11 10 $I ~ bull bull

11IflfU~Ufl~I1111unfil dl11itIlmillimfl91illiMln6tlfl~lfljihm~Vl1Ufl~fli~6~I~iiv 654plusmn003 1IIUiium1 flI1IJn11~m91MI~nV 666plusmnO05 lft~IlJili 1I11Ul11Um~nvlvhtitl 526IH267 flllJ 1m

Iflv~fimlv~1U16~UV1Iinrumiltl (2544) IIn11fJ~1tIItJ~~1tlll1n111tlUIl (2545) ~~llfll9hiJ~ii

il11lJVmllltmnhj~UIilVtitl6TI11i flmlllJlliiJ~i1lJrtlli1Yijjlt191111 ltyflhvhl1yeniilltl1111tlltl~UtH 4 ltl q~ bull - YI w~ bull

i11ifliltll111~ Il~ 11lll1fl11Nn m1fl1911flttll111nlIllUfl11~ftifJ~lIJtIllJl~mtlalI11~ trflflUlllU11l1 2

lillatl ii~IlllJl~ilfttl~ 1pl1milhm~ 2 flaUiiIJU1f)mllJm1ni91fl~ ml1Jfl~lmiddotmillfl~ IfII1111Um~nl

1~1JJiU nhtiU 851plusmn0021lfURIlJili 854t0091lfU~1JIi La 9280t228 fill mlJhiiu thullljiIJiU

~ a I dill ~ 41 d ~ i ~I bull1)U 1 fltlllflfllllflllfllmllI~1~lnflmiddotI~II1tIiIJljllffmlltl1lJ~U1~91J~lnffliltl 1J1J1l11l l ~l 1t1lIIlm~

~rI dl 1 Jjl Jq_II1W~ lVlfft11Ul ~l 111)1111ill ~ 1Ifl1i111l~~~tl~~WlllUlJ~~1t)~Ulll

J ~ J bull r i I 4 i I ~ I bull A fmllll1lfl~fI1~U ~II1lirnllflfm~mlflllJi1lJf1t IlHlllltl1 lJIWmI1V~IiJUltl~ I 1IJ lWfl~

~lfl~fl~fl1111yenflmhl~n~IIfl~I911mmUflalvfla~titlnS1IlI1PlJlfl~q~ l~mll4imJ~fl1mltJjlt1U1~

U6~11i1iii Ii~nfln11~9l1ltl~ih1IwlitljfllmVtlliiJtl111fl1fIItHIIi1Viit~Illflli1111n9 ffTl1fttl11liiiili itfl1illf)llti~1~1I n1lilfftlhlltitlfiml~ ffilfflU 11 lil(Ji111Ufl lUtillllflViUhl~i1iU1ill1U1t1lffltlfltl

bull y

61H11111fl11 (ff1aft 2506 thlii 2529) ulil~hilliil~Hua=iifl11flfliHfltl61H11ll11J11flflUilin4II

U1lZId61I111 (Vihnflla1io 2525 ~fllC 2529) li1utl1H11~ihMlfilflitllUlftt1~i~~ qfl~88l1f1nlallfuiu~ ftlhu lllI1i1HUfl iU6~111Wll 10 11I6flo1U~IItl~t1111iiflIgti1

- 1 (tplJllllafllC2533 IRnflIUlUIftfIt2541) ~ vitl p

fll1nflllltnllJtflJlnl1i111~UlHUflll1l1 fl111J I11n itiltlll II fl ~1lllfl1tJ1Ui 11 ff~fl1i fl1111

tI 3030) A f Ai pound (l 4 1d I I GO

alJIItli W = 01146 L lIlIff1Jfl1iTURlJfl1f(lIlJltlYllimfl111W~Ya~lJlfl 9l~il1lmflli

09936 bull

11tf~~ -)1ItlIltUtl Ill fl1111 n lJIiU S 11-l1~t11 HUflllll fl11lJl11 m 11 tl ~ Iii 1il fl111J NU 11111 IIllJ lit tllllJ1 fl

~ i1I-l1~~11111i~Iiitlil111lfJl1m1I611 1I1lilu1l1f111JflmiTI1Ufl 1lJtllJfl1ifl1111IDJiUliiftuUu~

T I d 1 Y I v a tlflfl11UtlJ1lU111tfl9111lflI1lJlltllllJl911lJIItlIlUfl~il11lJl11n191tllllillfl~itll1l 9936 lf~tlllfl11fll1

~mnUH~1tllU1l111tlUI1 (2545) ~ilifilIDJ1IlffYi~ItiWItfIII6~1~1fl1fllt1flfllltiiUi~ilmllJ ~

1l1l1nlill1Il11t1111lliil1lHVifffllil1l1lJ~1l1ii~i~11191fftl1l 1lllllflWhhI1hfitI 08659 UfIIIIflWfJh

fit 05019 I~ilfhiflalfllllfltUtl~l~lmU1l1~111qJl~ltfv~itlUtlfl6t1fll91IJUiV1111122 IfltlU ~~Ii filnlJ1Iliilli9i1t)11t1f1l1htiu 09968 1I1l 09895 ~llJihAtI (~IJ1~ 2528)

I

bull bull bull

bull bull

18

bull fi 11U nmiUil d1~ 11 nn1flU llm1Ufl1Hm llll4IJll~l~1mll1UiilUJfllS fi11unmlUsect

lJU W = 00721 2348 ~~17mhnnt11ti111U1l 09900 n4flhmlfffl1J1IJll4n11l1l1l1SStlItTl1

(2545) ~~fh R liltl4vJllliUinilillJ 08659 ImuliliUill11ilJ 05019 ff111SmmunuliUsectlJfNm1U

liillnWf(llmSYIIlllll~~lmll1Wli~ 7 ~tii1i)9111m1n)ilfll~ 0O lIllIJ1W~ UniI~hfm bull

~ 1llll~fll1~11~l1llIJ Nl~11111lliill(ululi 1 1 Ull1~ i ll~1l1~1 YI Illlmiddot1yen1flIJW1 ll fIlfftimnUljll~ ~1f(WtrU bull shy

1~1mlllUnlml1 iil191lYi J ri1WfJIlml1IJll~U11W UllYIllllll~ III flHll fll11lm 1nillll llllh~ flllYIIlllll~

iliiii1fl111~6fllm1fl19ilJli1IJfHI~lm61WI~mh~1

lflHllfll1fff11J1fli~i1 ~lmlU~11IJ6~ml~1J1ml1U~illW11l l~illl1Q~H 1 ~~(h

1 f1il flV~f(lJfl11IJ IlJW~IllllJUfh~l1n~lIJilJ1~ lmiddotjll~ il1f1iJ~lliiimSl lV~lU1i~ fll1ffmJ1m ~Ity

~lJ1i1UClf11111fl1 111111fohilmlf1111~tyI9ilJ11U lllIJll~f111 1~lJfl )11JJ11f11~W~lJll41~ lmlllW~ J JIfIY tJ tI~l~I_~cl q A Y ltl

61QilN 1 914HClYI 1llflf111ngtlCl6~fiHWffllJl1ilW1 lJl1J1~1JlnVlJUCl1I~IIJWII1l111n1 llJ1~ll1QIJiH

lJlfll lllW ~IilJ 1~1l1f1lin 1JlI191~~IJU li1l~ 1 nU timIllW111 iVlIU~llf111ftf1ln1W mmtlJ VIlIJll~

l~ lmll1UI141WJ1Wjh1Y1J1 mmnl~v~ UlIf111tll-jsmjlWfWlfill

IJWlllmlIJVTlt)~f(iI (La) uClnu1l1irYItmmtyI91lJ111 (K) 9l~rllWI1miJgt)1JIJl~f11l

lillliliWffumlIJtN von Bertalanffy (1938) Yl1ml1Ifll1Y1~~1 L nhilJ 598 19IU~IlJlIl 1lIri1 K

1l11nlJ 0138 ~6tJ 1hUfhll1~lrlllmllJ(J111lhnlJflU6 (to) 11l1hll1l1tl1Ul1 (2545) l~ftmnHilri11l~V

L = 598 (1 _ e-llmiddot 138 (t + 0216raquo)

bull Y bull bull v

til La il1ftil ilrilmfl61iifJ~nlJmlIJVl1m~ll~lJll~hmnl1UvJllllj~Ui111ii(J~ll~i1n11ihhlNU1i~

~~11gt1 ff~ Cl ill1m1lJ~~lfl1inlJJn91ll1Hitl]UvillUj~Ui J1li~lyeni1J 25 26 u~ lliVJ~ 1lJ1l1~~1llr~ 1 ~lJt~

1~1 li1vt~~mlllUiliiJInil1mliliq~ilm1IJV11fmm)~ fl11IJf)1Nmre~ UCl1l111Uf1lhnlJ 590

580 19IU91llJil1UI1 27800 f11U 1l1IJlh~lJ (fi1Ull~1nllUl1 2545) ll1lIIU1~IIe~IGilmmUl1q 9 tJ

~i~~H~n11ill 9l~hi91l1lJllf1Uf1mi~llll~flf~rl~m1UV11m~ il~lil~V 4250 19lU~lJm ~e1mi Q IV dlq 1 A tflO~f)jJfl1Y1Ii1~lf)lIlJf11lfn~II1lJ ll9l~~IYIlfllJ 4304I91UllllJiI~

ri1nlJlIliV1tIJll~f11mJI9i1J111 (fil K) i~iimtll10~fl11IJ11IJt1~f11ll~~tlJl~lJ1911rlilll1

bull ltV t 4 0 dt t 1

~ q I Q i lt$ i amplt9

31 La) lJil~1YmUl 91~ffmlJ1lllJlJ1~ffiJ ~Jff11J11ty)IillJ 11I1lU1H La) W1llIWN111Hl1llffil~lJlll~

ilfh K ilt)~ 1i1Un1I1111~ilril K ilhl1e~i~nllTI1llfJiI~~ilIJW1ll L (Sparre and Venema 1992)

d11111JfhnlJlI~nl1tf11~ I~~tyI~lJ1fl lJll~fl11IJ(J11 milll~lhmlllW (fil K) 1~~1 f)mllJ1V V411~HflnYl IIs lt1 I d~rltit bull d ls

ff U 91 ~um1Y~~fl111JfflJ Vi U li1111HmlU tn1miI e~lll Im IJ W f1lJfi 1[l1lIJIJ ll~fl11lWl1ml ~ lill11Jetl1lJ

19

~1~ 1 011 iitimhOIJ 0138 lfffl~1lin11111ItilmtJl11Ii111tY9Ii~iill1~Vll tIt1th~iill1~Vllfl-htill1~l~ iltil K lIi1OIJ 01915 (1tjJll1ff 2535)

I tJ fl111 fl ihlltJ1I111 ru1l91l1mil1ill191IJ19111 tJ~tilflj t tJ1 II illfltl11lJtYlJ~II t1al11~tJltJli1 II

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1

Gulland J A and S J Holt 1959 Estimation ofgrowth parameters for data at unequal time

intervals J ConsCons Int Explor Mer 25(1)47-49

Rounsefell GA and WH Everhart 1953 Fishery Science John Wiley and Sons Inc New

York 444 pp

Sparre P and S C Venema 1992 Introduction to Tropical Fish Stock Assessment Part Imiddot

Manual FAO Fish Tech Paper 3061 Rev I FAO Rome 376 pp

Uchida I 1967 Growth Qfloggerhead turtle Bull Sap Soc Sci Fish 33(6)497-506

von BertaIanffy L 1938 A quantitative theory of organic growth Human biology 10(2)81shy

213

19

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1

Gulland J A and S J Holt 1959 Estimation ofgrowth parameters for data at unequal time

intervals J ConsCons Int Explor Mer 25(1)47-49

Rounsefell GA and WH Everhart 1953 Fishery Science John Wiley and Sons Inc New

York 444 pp

Sparre P and S C Venema 1992 Introduction to Tropical Fish Stock Assessment Part Imiddot

Manual FAO Fish Tech Paper 3061 Rev I FAO Rome 376 pp

Uchida I 1967 Growth Qfloggerhead turtle Bull Sap Soc Sci Fish 33(6)497-506

von BertaIanffy L 1938 A quantitative theory of organic growth Human biology 10(2)81shy

213

bull bull bull

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20

bull 9 I laquoII q bull AI4 flU Ufl~lllU 2529 fllHlflEll11J~~WUflllH UUtl~fllmgttnU 1Hniflli1JilJ~ 39(5)535-539

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Eschscholtz 1WlifliitJ~laquor)1l1 lu ntJ~TWflliifIJlJW11Jilfllnh~lil 2528 flilJUiZlJ1 11J11

1

Gulland J A and S J Holt 1959 Estimation ofgrowth parameters for data at unequal time

intervals J ConsCons Int Explor Mer 25(1)47-49

Rounsefell GA and WH Everhart 1953 Fishery Science John Wiley and Sons Inc New

York 444 pp

Sparre P and S C Venema 1992 Introduction to Tropical Fish Stock Assessment Part Imiddot

Manual FAO Fish Tech Paper 3061 Rev I FAO Rome 376 pp

Uchida I 1967 Growth Qfloggerhead turtle Bull Sap Soc Sci Fish 33(6)497-506

von BertaIanffy L 1938 A quantitative theory of organic growth Human biology 10(2)81shy

213

1

Gulland J A and S J Holt 1959 Estimation ofgrowth parameters for data at unequal time

intervals J ConsCons Int Explor Mer 25(1)47-49

Rounsefell GA and WH Everhart 1953 Fishery Science John Wiley and Sons Inc New

York 444 pp

Sparre P and S C Venema 1992 Introduction to Tropical Fish Stock Assessment Part Imiddot

Manual FAO Fish Tech Paper 3061 Rev I FAO Rome 376 pp

Uchida I 1967 Growth Qfloggerhead turtle Bull Sap Soc Sci Fish 33(6)497-506

von BertaIanffy L 1938 A quantitative theory of organic growth Human biology 10(2)81shy

213