Growth Pattern of the Young of the Year Argentine Hake Merluccius Hubbsi Marini, 1933 (Gadiformes Merluccidae) Along the Brazilian and Uruguayan Coasts
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Environ Biol Fish (2011) 91:155–164 DOI 10.1007/s10641-011-9768-4 Growth pattern of the young of the year Argentine hake Merluccius hubbsi Marini, 1933 (Gadiformes Merluccidae) along the Brazilian and Uruguayan coasts María Inés Lorenzo & André M. Vaz-dos-Santos & Carmen Lúcia D. B. Rossi-Wongtschowski Received: 23 November 2009 /Accepted: 3 February 2011 /Published online: 22 February 2011 # Springer Science+Business Media B.V. 2011 Abstract Morphology and morphometry of the sag- northern and southern area. As regards the Uruguayan ittae otolith were studied in young of the year Argentine coast and Southern Brazil, age groups zero shared hake, Merluccius hubbsi in the Southeastern Atlantic. some similarity as well as differences in growth; Geographical variation in the growth pattern of the environmental features explain these differences but young of the year Merluccius hubbsi was correlated their effect on the adult fishes growth is still unknown. with the differences found in otolith morphometry, relative growth, ring positions, as well as the formation Keywords Otoliths . Young of the year . of a new ring. The otolith development of M. hubbsi Morphometry . Merluccius hubbsi . throughout its area of distribution accompanies an Southeastern Atlantic increasing northward temperature gradient, resulting in bigger otoliths and greater variability of ring position from northern to southern area. The canonical discrim- Introduction inate analysis showed that the otolith length and height explained most of the variation of the first discriminant Geographical variations of marine fish populations function. We found significant differences in the are common and constitute a tool for stock identifi- discriminant scores between those samples from cation (Cadrin et al. 2005) in fishery science. The understanding of the degree of homogeneity of fish M. I. Lorenzo (*) stocks is necessary for effective fisheries manage- Dirección Nacional de Recursos Acuáticos—DINARA, ment, such as the prevention of overfishing, the Constituyente, 1497, preservation of their reproductive and genetic diver- CP 11200 Montevideo, Uruguay sity and the achievement of optimal strategies for their e-mail: [email protected] replenishment (Lockwood 1988; Begg et al. 1999). A. M. Vaz-dos-Santos Growth pattern studies are very useful to ascertain Universidade São Judas Tadeu—USJT. geographical variations in a dispersed population Núcleo de Estudos Biológicos, within a given area. In many marine fish, the otoliths Rua Taquari, 546 Mooca, CEP 03166-000 São Paulo, SP, Brazil are the structures used to differentiate a species geographical growth pattern (Campana and Casselman C. L. D. B. Rossi-Wongtschowski 1993). Morphometric methods based on the character- — Instituto Oceanográfico da Universidade de São Paulo istics of the otoliths and the deposition pattern of their IOUSP. Laboratório de Ictiofauna e Crescimento (LABIC), Praça do Oceanográfico, 191 sala 103ª, rings have been used to approach this subject. These CEP 05508-900 São Paulo, SP, Brazil studies reveal quantitative measures of the otoliths 156 Environ Biol Fish (2011) 91:155–164 shape thus making possible the differentiation of stocks and Brazilian coast respectively. As regards spawning and populations (De Vries et al. 2002;Cardinaleetal. and nursery areas of M. hubbsi on Uruguayan and 2004; Bergenius et al. 2006). These different morpho- Brazilian coasts, this study has focused on the growth logic characters have been associated with environ- of the young of the year in order to identify mental factors such as temperature, salinity and depth geographical variations and thus give support to the (Castonguay et al. 1991;Dawson1991; Torres et al. knowledge of the species dynamics in the area. 1996; Gauldie and Crampton 2002). It is not the origin of biological differences that is important, but their maintenance over time (Cadrin et al. 2005). Material and methods The genus Merluccius constitutes one of the most important fishery resources of the world (Alheit and Trawl surveys and samples Pitcher 1995). This is also de case of the Argentine hake (Merluccius hubbsi Marini, 1933), a species found Trawl surveys were conducted on the RV Aldebarán from 21°S to 55°S in the Southwestern Atlantic Ocean, off Uruguayan and Argentinian coasts and off the which occurs throughout Brazilian, Uruguayan and Brazilian coast on the RV Altântico Sul, covering the Argentinian waters. It is the main target species of the area from 22oSto37oS, from depths between 50 and Uruguayan and Argentinean trawler fleets and since 300 m, during autumn (April to June) and spring 2001 has also been a target of Brazilian trawlers (Perez (October to December), 1997. The whole area was et al. 2003; Vaz-dos-Santos and Rossi-Wongtschowski divided into three sections for analysis purposes: A 2005). Recently M. hubbsi was considered a threatened (34°–37°S), B (29°–34°S) and C (22°–29°S) (Fig. 1). species due to excess of juveniles fishery, reduction of Fish sampling focused on juveniles of M. hubbsi (LT, biomass and an intense decrease of catches in the three total length < 250 mm). During surveys hydrological countries which it occurs (Vaz-dos-Santos et al. 2010). data was also collected, being the local depth and The species is most abundant between 35ºS (off the La bottom temperature of each fishing site recorded. Plata River estuary, Uruguay) and 45ºS (the Patagonian The Argentine hake specimens caught were mea- shelf, Argentina). In southern Brazil, its abundance is sured (LT, mm), and the sagittae otoliths removed. In related to cold waters, mainly between 33oS and 35oS the absence of difference between left and right (Haimovici et al. 2004, 2008). Throughout its distribu- otoliths (Vaz-dos-Santos and Rossi-Wongtschowski tion area, M. hubbsi presents displacements relating to 2007), when the right one was unavailable the left oceanographic features (Podestá 1989; Vaz-dos-Santos otolith, was measured for length (OL, mm, the and Rossi-Wongtschowski 2005) and its life history maximum distance between the anterior and posterior (Vaz-dos-Santos et al. 2009). thus it is possible to edges) and for height (OH, mm, the maximum recognize some spawning and nursery areas at Isla distance between upper and lower edge) (FAO Escondida (Argentina) during summer (December to 1981). The otoliths were also weighted (OW, g). February), off the Rio de la Plata Estuary during autumn The otoliths were cut transversely along the nucleus, (main peak) and spring (second peak), off the Patos grinded and burned according to the technique Lagoon and on the continental shelf (29oS–34oS, proposed by ICSEAF (1983), and examined under Southern Brazil) in autumn-winter and off Southeastern reflected light with magnification of 10×. The otolith Brazil (21oS–29oS) during spring-summer (Honji et al. radius (OR, mm) were measured from the nucleus to 2006;Vaz-dos-Santosetal.2009). the ventral margin. , the position of the two rings In specific relation to otoliths and growth pattern, only formed during the first year was located, and named Torres et al. (1996, 2000) focused on the otolith of M. demersal rings De I and De II according to ICSEAF hubbsi, in search for intraspecific geographic differ- (1983), Renzi and Pérez (1992) and Vaz-dos-Santos ences. Regarding the Argentine hake, there are many and Rossi-Wongtschowski (2007). other studies focused both on larvae/juveniles and adult growth, but without geographic comparisons. Most Data analysis recent contributions related to adult growth have been presented by Lorenzo (2003) and Vaz-dos-Santos and Depth and water temperature were tested by a two Rossi-Wongtschowski (2007)inregardtoUruguayan way analysis of variance (ANOVA) with both factors Environ Biol Fish (2011) 91:155–164 157 Fig. 1 Survey area (depths – S between 50 300 m) and 22 distribution of bottom trawl Autumn 1997 Brazil Cape Frio fishing surveys carried out Spring 1997 in autumn and spring 1997 24 in the Southwestern Atlantic Ocean. Area A: 34°–37°S; 26 Area B 29°–34°S and Area AREA C C 22°–29°S 28 Cape of Santa Marta Grande 30 Latitude AREA B 32 UruguayUruguay 34 Southwestern La Plata River 36 Atlantic Ocean 38 AREA A 100 m 40 500 m 58 56 54 52 50 48 46 44 42 40 Longitude W (area and season) fixed. Total length samples were provided for each significant variable represented the described based on exploratory analysis, taking area contribution of the respective morphometric variables and season into consideration (Zar 2010). for the discrimination of Argentine hake samples Regressions between LT and OL, OH, OW were between areas (Manly 2008). calculated for each season and area, fitting allometric As regards the otolith sections, differences in models (Huxley 1993) by means of the least square ring measurements were analyzed in the light of method (Zar 2010). The equation used was Y=a+Xb, length frequency distributions and compared by with logarithmic data to homogenize the residuals. means of a Kolmogorov-Smirnov test in accordance ANCOVA was applied to determine the effect of fish withthesequenceofareas(Zar2010). The lengths on the magnitude of each otolith’s morpho- comparisons between averages demersal I and II metric variable, first by season and then by area. The rings were made with a Scheirer-Ray-Hare test, after variables significantly correlated with fish length were normality and homocedasticity tests (Sokal and Rolf corrected by using the common intra-group slope (b) 1995). (Sokal and Rolf 1995). Relative growth was also analyzed by the application of a t-test, taking the reference values for isometry β=1 for relationships Results involving length and height and β=2, for weight (Huxley 1993; Quinn and Deriso 1999). Environmental data (depth and bottom temperature) Forward stepwise canonical discriminant analysis varied between areas and seasons (Table 1). Variations (CDA) with standardized data was used to detect on depth were significant due to areas (F=7.85, df=2, morphometric differences in otolith samples from the P=0.05).