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Growth of the Pacific jack Caranx caninus
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1. Cuba Instituto de Oceanolog a Acad Cien Cuba La Habana Cuba Sudekum A E J D Parrish R L amp S Ralston 1991 Life history and ecology of large jacks in undisturbed shallow oceanic communities Fish Bull U S 89 493 513 Taylor C C 1958 Cod growth and temperature J Conseil 23 366 370 Taylor C C 1960 Temperature growth and mortality the Pacific cockle J Conseil 26 177 124 Torres Aguilar M 2002 Estudio preliminar de edad y crecimiento del jurel Caranx caninus G nther 1867 con base en v rtebras y espinas en las costas de Guerrero y Michoacan M Sc Thesis UNAM M xico van der Heiden A M 1985 Taxonomia Biologia y eva luaci n de la ictiofauna demersal del Golfo de California 149 200 In A Yanez Arancibia ed Recursos pesqueros potenciales de M xico La Pesca acompa ante del camar n Progr Univ de Alimentos ICMyL UNAM M xico D F M xico von Bertalanffy L 1938 A quantitative theory of organic growth inquiries on growth laws Human Biology 10 181 213 Walford L A 1946 A new graphic method of describing the growth of animals Biol Bull 90 141 147 Zar 1996 Biostatistical analysis Prentice Hall New Jersey USA Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 56 1 171 179 March 2008 179
2. due to the very few organisms under one year of age in the com mercial catch because of gear size selectivity The use of the ELEFAN program Gayanilo et al 1994 distinguished similar age group Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 56 1 171 179 March 2008 175 comparable with those found by Cruz Romero et al 1993 who also used an indirect method for their analysis It is important to note that the values of this paper and those of Torres A guilar 2002 in relation to the ones found a decade ago by Cruz Romero et al 1993 suggest that the population is stable in spite of the fluc tuations in the physical chemical and fishery variables a reduction in its length which is an over fishing symptom has not been observed Espino Barr 2000 Allometric growth index shows intrannual variations which depend in great measure on the relation of the organism with its environ ment and is inversely proportional to the condi tion factor Safran 1992 Cruz Romero et al 1993 found a negative allometric value 1 b 2 28 while in the present work the indexes are isometric b 2 957 with total weight and b 2 954 with eviscerated weight Theoretical value of longevity Taylor 1958 is based in von Bertalanffy s formula as 95 of the asymptotic length Taylor 1960 As with the growth parameters longevity is very sensitive to the sample that is if the organisms are large the asymptotic length increases and t
3. 12 12 661 12 12 6661 19 12 1661 1010 1661 122 9861 S861 Ag an M y sono OJAWOY ZNIJ OIDWOY ZN IQ OJAWOY ZNIJ winyapng UOSUYOf W UIMPOOL 29 8 al snuas xue ayy fo saizads quasaffip asp yova fo syiBua 8 2 dfjunjojiag ayy fo YIMOLL 177 171 179 March 2008 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 56 1 a massive spawning period is also observed during May RESUMEN El jurel Caranx caninus es una especie com n que se captura en la pesca artesanal frente a las costas de Colima M xico Mensualmente durante 2002 se tomaron datos morfom tricos y muestras de otolitos para determinar la edad y el crecimiento Se encontraron siete grupos de edad durante el primer a o de vida se da el mayor crecimiento hasta 14 4 cm Durante el segundo a o C caninus crece 11 76 cm el tercero 9 61 cm el cuarto 7 85 cm el quinto 6 41 cm y el ltimo a o crece 5 24 cm Las constantes de la ecuaci n de crecimiento de von Bertalanffy fueron L 83 26 cm W 18 138 g 0 202 t 0 283 y la longevidad A ys 15 a os Las curvas de crecimiento de otras especies del mismo g nero se compararon con las obtenidas en el presente trabajo El ndice gonadosom tico present valores m s altos durante noviembre y mayo Los periodos de tiempo de alimenta
4. Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 56 1 171 179 March 2008 TABLE 2 Average length for each age caninus according to different authors Standard length cm Age Cruz Romero et al 1993 1 20 64 2 32 95 3 42 44 4 49 76 5 55 40 6 59 75 7 8 9 indirect method Gayanilo et al 1994 direct method with vertebrae Standard length cm Torres Aguilar 2002 Standard length cm Present paper direct method with otoliths adjusted with Solver Microsoft 1992 0 041 0 039 0 037 0 035 0 033 0 031 0 029 0 027 0 025 Condition factor a 4 62 22 35 19 02 29 86 30 78 36 80 40 39 43 20 48 24 49 11 54 65 54 57 59 89 59 60 64 25 68 54 5 4 5 4 3 5 3 10 25 2 0 15 O Fig 5 Monthly values of the condition factor a tot total weight and a evi eviscerated weight gonadosomatic index in females GSI and gastric repletion index are deposited in the otoliths validating the use of this structure to estimate age based in the number of bands in Caranx caninus In this paper the values obtained with the length frequency analysis were very similar to those found by the identification of growth rings in the sagittae and asterisci otoliths which renders age determination in caninus valid with both methods The otolith analysis allowed the calculation of the average length for age group cero which was not identified by the polimodal curves
5. MATERIALS AND METHODS From January to December 2002 fifty specimens of the Pacific jack C caninus were obtained monthly from the commercial catch in Manzanillo Colima M xico and taken to the lab where standard Ls cm and total length Lt cm total Wt g and eviscerated weight We g gonad Wg g and stomach weight Ws g were recorded for each individual The total weight Wt standard Ls and total length Lt of over 200 individuals per month were taken in situ The length frequency distribution was ana lyzed with the ELEFAN program of the FISAT package Gayanilo et al 1994 to obtain the average length of the polimodal curves corre sponding to each age group The average length of the age groups determined by the analysis of the otoliths sagittae and asterisci by Espino Barr et al 2006 were used to obtain the constants of von Bertalanffy s 1938 growth equation The observed values were for age 0 6 33 cm age 1 15 09 cm age 2 32 19 cm age 3 41 50 cm age 4 48 17 cm age 5 55 58 cm and age 6 58 75 cm The time of the growth ring formation was determined observing whether the borders had slow or fast growth rings In every case otoliths were observed by transpar ency with transmitted light the hyaline zone corresponds to the slow growth band and the opaque zone to the fast growth band which is in contrast with reflected light Blacker 1974 There were no growth differences between sexes Espino Barr
6. Trop 54 195 207 Mendenhall W 1987 Introduction to probability and sta tistics PWS Kent Publishing Boston Massachussets USA Microsoft 1992 Excel Manual de usuario Referencia de funciones Microsoft Redmond Washington USA Ramirez Granados R 1977 Estudio de la factibilidad de una terminal pesquera en Manzanillo Col Depto Pesca M xico 2 268 338 Rodr guez Guti rrez M 1992 T cnicas de evaluaci n cuantitativa de la madurez gon dica en peces AGT M xico DF Rojas M J R E Maravilla amp Chicas 2004 H bitos alimentarios del pargo mancha Lutjanus guttatus Pisces Lutjanidae en Los C banos y Puerto La Libertad El Salvador Rev Biol Trop 52 163 170 Rojas Herrera A A M Mascar amp X Chiappa Carrara 2004 H bitos alimentarios de los peces Lutjanus peru y Lutjanus guttatus Pisces Lutjanidae en Guerrero M xico Rev Biol Trop 52 959 972 Safran P 1992 Theoretical analysis of the weight length relationship in fish juveniles Mar Biol 112 545 551 Saucedo Lozano M G G nzalez Sans n amp X Chiappa Carrara 1999 Alimentaci n Natural de j venes de Lutjanus peru Nichols y Murphy 1922 Lutjanidae Perciformes en la costa de Jalisco y Colima M xico Ciencias Marinas 25 381 400 Sierra 8 Popota 1988 Velocidad de digesti n ritmo diario de la alimentaci n y raciones alimen ticias de algunas especies de peces del Golfo de Bataban
7. Growth of the Pacific jack Caranx caninus Pisces Carangidae from the coast of Colima M xico Elaine Espino Barr Manuel Gallardo Cabello Esther Guadalupe Cabral Solis Arturo Garcia Boa amp Marcos Puente G mez 1 CRIP Manzanillo INP Playa Ventanas s n Manzanillo Colima 28200 M xico Tel 01 314 33 32 37 50 Fax 01 314 33 32 37 51 escama webtelmex net mx elespino gmail com 2 Instituto de Ciencias del Limnologia UNAM Apartado Postal 70 305 C P 04510 M xico D F M xico gallardo mar icmyl unam mx Received 02 IV 2006 Corrected 08 V 2007 Accepted 22 2007 Abstract The Pacific jack Caranx caninus is a common species fished by artisanal fishermen off the coast of Colima M xico During 2002 monthly samples of morphometric data and otoliths were taken to determine age and growth Seven age groups were identified The highest growth 14 4 cm takes place during the first year of life During the second year C caninus grows 11 76 cm the third year 9 61 cm the fourth 7 85 cm the fifth 6 41 cm and sixth year 5 24 cm The constants of von Bertalanffy s growth equation were L 83 26 cm W 18 138 g K 0 202 t 0 283 and Ay ys 15 years Growth curves of other species of the same genus were calculated in order to compare them with the one obtained in the present work The gonadosomatic index presented higher values during November and May The per
8. Zool 1 81 88 Cruz Romero M E Espino Barr amp A Garcia Boa 1996 Potencial de la pesca ribere a en el estado de Colima M xico en 1989 INP SEMARNAP Ciencia Pesquera 12 30 35 Espino Barr E M Cruz Romero amp A Garcia Boa 2003 Peces marinos con valor comercial de la costa de Colima M xico CONABIO INP CRIP Manzanillo ISBN 970 9000 23 3 M xico D F M xico Espino Barr E E G Cabral Sol s A Garcia Boa amp M Puente G mez 2004 Especies marinas con valor comercial de la costa de Jalisco M xico SAGARPA INP ISBN 968 800 570 3 M xico D F M xico Espino Barr E 2000 Criterios Biol gicos para la admi nistraci n de la pesca multiespec fica artesanal en la costa de Colima M xico Tesis de Doctorado Colima M xico Espino Barr E M Gallardo Cabello A Garcia Boa E G Cabral Sol s amp M Puente G mez 2006 Morphologic and morphometric analysis and growth rings identification of otoliths sagitta asteriscus and lapillus of Caranx caninus Pisces Carangidae in the coast of Colima Mexico J Fish Aquat Sci 1 157 170 Ford E 1933 An account of the herring investigations conducted at Plymouth during the years from 1924 to 1933 J Mar Biol Assoc U K 19 305 384 Gallardo Cabello M E Espino Barr A Garcia Boa E G Cabral Sol s amp M Puente G mez 2006 Morphologic and morphometric analysis and growth rings identification of otoliths sagitta asteriscus a
9. ci n m s intensa son de agosto a febrero Palabras clave Caranx caninus edad y crecimiento distribuci n de frecuencia de talla sagittae asterisci ecua ci n von Bertalanffy longevidad REFERENCES Aguirre Le n A 8 S D az Ruiz 2006 Estructura de tallas madurez gon dica y alimentaci n del pez Diapterus rhombeus Gerreidae en el sistema fluvio deltaico Pom Atasta Campeche M xico Rev Biol Trop 54 599 611 Blacker R W 1974 Recent advances in otolith studies p 67 90 In J Harden Jones ed Sea Fisheries Research London England Bussing amp M I Lopez 2004 Peces de Isla del Coco y peces arrecifales de la costa pacifica de Centro Am rica Meridional Rev Biol Trop 52 1 191 Castro Aguirre J L 1978 Cat logo sistem tico de los peces marinos que penetran a las aguas continentales de M xico con aspectos zoogeogr ficos y ecol gi cos Depto Pesca INP Serie Cient fica 19 M xico D F M xico Chirichigno N W Fischer amp C W Nawen 1982 Cat logo de especies marinas de inter s econ mico actual o potencial para Am rica Latina Infopesca Pac fico Centro y Suroriental Roma fao pnud Italy Cruz Romero M E Espino Barr amp A Garcia Boa 1989 Lista de peces del litoral Colimense INP SEPESCA Serie Doc Trab 1 21 p Cruz Romero M E Espino Barr amp A Garcia Boa 1993 Car ngidos aspectos biol gico pesqueros en el lito ral colimense Cuad Mex
10. e and October This helps to avoid interspecific competition and achieve a successful spawning Ib ez Aguirre 1995 Rojas Herrera et al 2004 These two species have similar characteristics and behavior Morphologically the Pacific jack reaches larger sizes but from the fishery point of view they are both coastal pelagic and are fished in the same way The three indexes condition factor a gonadosomatic IGS and gastric repletion IRG help to assume that the main spawn ing periods occur in November and May because during this period the corporal waste of the fish is higher and looses weight The most active feeding periods coincide with the period of the fast growth rings in the oto liths that is during the months of August to February period that also coincides with the rainy season Observatorio de Manzanillo pers com Espino Barr 2000 responsible for the wash of abundant nutriments to the rivers and to the coastal zones the same is true for the hurricane season Observatorio de Manzanillo pers com Espino Barr 2000 which remove the marine bottoms of Colima s narrow continental shelf and sends nutrients to the surface of the ocean and increases food availability During this time the individual weight increases and a recovery of the repro ductive population takes place larvae and juveniles also the recruitment to the fishing zone occurs Conversely during the spring summer period March August the ava
11. et al 2006 The constants L K y t of von Bertalanffy s 1938 equation were obtained with Ford s 1933 Walford s 1946 and Gulland s method 1964 and were adjusted by convergent iterations with Newton s algo rithm with the Solver program in Excel soft ware Microsoft 1992 Weight length relationship was calculated with the function W a L gt and a t student test was used to prove allometry Mendenhall 1987 Zar 1996 The weight for every age was obtained with the growth data in length and the weight length function Weight growth was obtained substituting L and for W and W in von Bertalanffy s 1938 equation Age limit or longevity 95 of L was determined with Taylor s equation 1958 1960 Ao In 1 0 95 K Monthly values of the condition factor equivalent to the a parameter of weight length equation were obtained for total and eviscerated weight Safran 1992 and com pared with the confidence interval to explain changes in the gonad liver and stomach throughout the year The gonadosomatic index GSI was determined with the equation described by Rodr guez Guti rrez 1992 GSI Wg Wt 100 where Wg is gonad weight and Wt is the total weight of the individual 172 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 56 1 171 179 March 2008 The gastric repletion index accord ing to Sierra and Popota 1988 is GRI Ws Wt 100 where Ws is the stomach weig
12. fi 50004 Wt a 2454 dence intervals of the indexes y 4000 4 0 033 x 157 Wt calc 30004 82 0 991 showed that there are sig 20004 551 1000 0 10 20 30 40 50 60 70 Standard length cm Fig 4 Relation between standard length Ls and total Wt and eviscerated weight We of C caninus Theoretical growth in weight table 1 shows the values of theoretical growth for total and eviscerated weight at each age The first year the organism reaches 230 g of total weight and 198 g of eviscerated weight For the second year the values increase potential ly 956 g of total weight and 822 g of eviscer ated weight The asymptotic total weight was Wt 18 138 g and the eviscerated asymp totic weight We 15 544 g Longevity Age caninus reached 95 of its infinite length L at 15 years nificant changes throughout the year Figure 5 shows the trends of the monthly average data of the indexes of condi tion factor a gonadosomatic GSI and the gastric repletion GRI The massive spawning periods occur in November and May GSI The most active feeding sea son represented as condition factor a tot and a evi is from August to February The gastric repletion index GRI seems stable during this period reaching its higher value in December January and May DISCUSSION According to the results showed in Table 2 each year a band of fast and of slow growth 174 Rev
13. ges three to four 6 41 from ages four to five and 5 24 cm from ages five to six Growth in weight The allometric growth index value of the weight length equation was isometric b 2 957 with total weight data t 0 561 lt ty 05 2 426 and b 2 954 with eviscer ated specimens t 0 875 lt t 52 they were statistically equal to 3 Fig 4 N N N N N N N N N N F 22 9 9 gt gt o 5 5 5 Y lt 7 2835 O Zz Months O hyaline MH opaque Fig 2 Monthly frequency of the slow growth and fast growth borders in the C caninus sagittae Months Standard length cm Fig 3 Monthly length frequency distribution of C caninus 173 1 Average of the standard length and total and eviscerated weight for each age group years caninus A Standard length cm Standard length Total weight Eviscerated se frequency distribution cm otoliths g weight g 0 4 62 4 3 1 16 34 19 02 230 198 2 29 99 30 78 956 822 3 40 53 40 39 2 136 1 834 4 48 65 48 24 3611 3 099 5 54 91 54 65 5 222 4 481 6 59 74 59 89 6 845 5 872 7 63 46 64 17 8 395 7200 8 66 33 67 66 9 821 8 421 9 68 55 70 52 11 098 9515 indirect method with FISAT Gayanilo et al 1994 direct method with otoliths adjusted with Solver Microsoft 1992 8000 Bioecologic consider 7000 wt 0 038 x 152957 ations during the growth of C gt B 60004 R 5 caninus analysis of the con
14. he growth coefficient decreases whereas if the organisms present a large K value its asymptotic length is reached in a shorter period of time In the pres ent work the value of longevity for the different species of carangids is from eight and beyond Calculations for the Pacific jack in Colima are 12 and 15 years Cruz Romero et al 1993 and this study respectively while the values for Guerrero and Michoacan Torres Aguilar 2002 reach up to 37 years Results on growth analysis of other spe cies of Caranx genus were used because of the lack of specific data Garcia Arteaga and Reshetnikov 1985 Goodwin and Johnson 1986 Sudekum et al 1991 Cruz Romero ef al 1993 Torres Aguilar 2002 and Gallardo Cabello et al 2006 For the different species the value of the K index result in curves of steeper trends for larger specimens 0 26 to 0 35 and the length reaches the asymptotic length L faster or in a shorter time Table 3 This occurs with C crysos a species with the highest growth rate which reaches 21 92 cm in its first year Together with C caninus Torres Aguilar 2002 C ruber C ignobilis and C melampygus these are the carangids with the highest longevity The GSI is an indicator of the gonad devel opment and therefore of the reproductive cycle of the fish Rodr guez Guti rrez 1992 The results indicate higher reproductive activity in May and November different from C caballus whose active months are Jun
15. ht and Wt is the total weight of the individual RESULTS Growth rings time of formation of the slow and fast growth bands C caninus shows that a higher percentage of otoliths with fast growth bor ders occurs during August to February autumn winter while the highest percentage of otoliths with slow growth bands in the border were observed during March to July spring summer Fig 2 Analysis of length fre quency the values of standard length for each age are shown in Table 1 and Figure 3 The first year the organism grows 16 34 cm during the second year it has reached 30 cm Growth parameters for the standard length are L 76 00 K 0 26 years and t 0 07 The highest increment occurs during the first year of life during the next years growth rate decreases nota bly 13 66 cm from age one to age two 10 53 from age two to age three 8 12 from age three to age four 6 26 from age four to age five and 4 82 cm from age five to age six Analysis of otoliths anal ysis of the sagittae and asterisci otoliths permitted to identify seven age groups Growth parameters for the standard length were L 83 26 cm K 0 202 years t 0 283 Instantaneous growth rate was 14 4 cm from age cero to Frecuency Jan 02 Frequency Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 56 1 171 179 March 2008 Feb 02 age one 11 76 from age one to age two 9 61 from ages two to three 7 85 from a
16. ilability of the food diminishes the values of the condition factor index decreases and the slow growth rings are formed However during this season 176 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 56 1 171 179 March 2008 8128 6 Tl SI 0L6L 0 8 l 8 by rl ZOLL 158 TEELT 96 8 06101 18 Tp 160 8v78 WTI 5 9619 79 08 051 L9 8 01 189 6 IL LT8L 801 0 0p 589 6 1865 7 IE SL 17 001 ESHE 8 ross 8289 6S 1 916 09 L 68 65 8 05 8759 06 99 6618 6816 Plog 9 so pS 91 LLTS 9L Tp or 19 10 19 ETIL TY LT STOP peor tror 68 LE soss 685 665 r 60 STE 10 8 16 9Y7E 59 Lp ST pp 1 9p ENTE oriz 8L 0 1817 0717 PST 68 b7 6 9 OSZE 6816 TILT T 20 61 18 07 srel 8891 6T l LS 07 ILI 181 2612 LEEI I 0 5323 078 68 8 8 6051 96 67 95 8 96 67 682 0 TS I OSTO 680 0 0 0 0 690 0 br0 0 L60 0 OLTI 8ZL I 4 2020 080 0 090 Z9 0 seco 6 0 DETO ooro 050 ooro y 9068 06 TIT Ive TOTS 1616 851 01768 08 81 Ov Ir 00 95 uo 7 SNUJUDO 5 SNULUDO 9 9 0 9 snypiaspfxas 5 snS dunjau J 511 0 81 J 505412 5 5 ds OxI EENI vna yna YSN XPW Jo ODIXIJA OxI ODIXAJA PUNO eqno ewop euog BIND JO ON sida 2007 6661 10 19 9007
17. ined the average length for each age with the analysis of the sagittae and the asterisci Age and growth analysis are necessary to understand the population s dynamics and Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 56 1 171 179 March 2008 171 the fluctuations of its biomass in time and space Lucano Ramirez et al 2006 The use of the ELEFAN method Gayanilo al 1994 Jim nez Badillo 2006 and the analysis of the otoliths are good instruments to estimate age groups and obtain growth parameters The weight length relation helps to cal culate the condition factor that is necessary to understand weight variations through the year Safran 1992 Likewise the calculations of the gonadosomatic index and the gastric reple tion index help to understand the growth phe nomena the gonadic maturity and the trophic dynamics of the populations Saucedo Lozano et al 1999 Rojas Herrera et al 2004 Rojas et al 2004 Aguirre Le n and Diaz Ruiz 2006 In this study the average length for each age was estimated by means of the monthly length frequency distributions The time of the slow and fast growth bands formation in the otoliths sagittae was determined The growth constants of von Bertalanffy s 1938 equation were calculated from the polimodal analysis and from the analysis of sagittae and asterisci The longevity and the values of the condition factor gonadosomatic and gastric repletion were also calculated
18. iods of more intensive feeding are from August to February Rev Biol Trop 56 1 171 179 Epub 2008 March 31 Key words Caranx caninus age and growth length frequency distribution sagittae asterisci von Bertalanffy s equation longevity Caranx caninus G nther 1868 Fig 1 is a pelagic and coastal species of the continental shelf from California USA to the north of Peru Chirichigno ef 1982 It is fished during the whole year in the coast of Colima M xico and its capture volumes summing about 15 of total catch annual average make it important Cruz Romero et al 1993 The Pacific jack is considered a second and third class commercial product its price does not exceed five pesos on the beach or eight pesos per kg in the market 0 50 to 0 80 USD In the State of Guerrero it is considered a second class fish but with a better price P rez Vel squez pers com Analysis on its taxonomic descrip tion have been made by Castro Aguirre 1978 Chirichigno et al 1982 van der Heiden 1985 Bussing and L pez 2004 on general aspects of its fishery by Ramirez Granados 1977 Fig 1 Caranx caninus G nther 1868 Cruz Romero ef al 1989 1996 Espino Barr et al 2003 2004 Cruz Romero et al 1993 analyzed general aspects of its biology Recently Torres A guilar 2002 studied its age and growth determination in Guerrero and Michoacan with vertebrae and spines Espino Barr et al 2006 determ
19. nd lapillus of Caranx caballus Pisces Carangidae in the coast of Colima Mexico Internat J Zool Research 2 34 47 Garc a Arteaga J P amp Y S Reshetnikov 1985 Age and growth of the barjack Caranx ruber off the coast of Cuba J Ichthyol 25 120 131 Gayanilo F C Sparre amp D Pauly 1994 The FAO ICLARM Stock Assessment Tools FiSAT User s Guide FAO Computarized Information Series Fisheries Rome Italy 178 Rev Biol Trop Int J Trop Biol ISSN 0034 7744 Vol 56 1 171 179 March 2008 Goodwin J M amp Johnson 1986 Age growth and mortality of blue runner Caranx crysos form the northern Gulf of Mexico Northeast Gulf Sci 8 107 114 Gulland J A 1964 Manual of methods of fish population analysis FAO Fish Tech Paper Rome Italy Jim nez Badillo L 2006 Age growth models for tilapia Oreochromis aureus Perciformes Cichlidae of the Iniernillo reservoir Mexico and reproductive behav iour Rev Biol Trop 54 577 588 Ib ez Aguirre A L 1995 Algunos aspectos de la din mi ca de poblaciones de Mugil cephalus Linneo 1758 y M curema 1836 Pisces Mugilidae en la laguna de Tamiahua Veracruz Tesis doctoral Fac de Ciencias UNAM M xico D F M xico Lucano Ram rez G S Ruiz Ram rez 8 J A Rojo V zquez 2006 Composici n por tallas y ciclo repro ductivo de Pseudupeneus grandisquamis Pisces Mullidae en el Pac fico central Mexicano Rev Biol
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