Age structure and growth of the rough scad, Trachurus lathami (Teleostei: Carangidae), in the Southeastern Brazilian Bight

Lygia C. Ruas André M. Vaz-dos-Santos About the authors

ABSTRACT

The rough scad, Trachurus lathami Nichols, 1920, is a small pelagic species distributed along the West Atlantic coast. It is most abundant in the Southern Brazil (28°30’-34°S) and in the Southeastern Brazilian Bight (SEBB, 22°-28°30’S). The rough scad is fished by purse seines, which main target is the Brazilian sardine, Sardinella brasiliensis (Steindachner, 1879). Age and growth are vital to understand the life cycle of a species, to fishery management and ecosystem modeling. This study aimed to assess the age and growth of T. lathami, to identify its age structure in the SEBB, and to evaluate what causes the wide differences among Trachurus species in terms of body size and growth parameters. Data available on T. lathami was attained between 2008 and 2010 from surveys at SEBB. A total of 278 whole otoliths of T. lathami, total length between 27 mm and 208 mm, were analyzed and compared with the only other source of otolith data, from 1975. Three blind readings were performed and assessed using traditional methods to study fish age and growth. Zero up to eight rings were found, each ring corresponding to one year in the life of an individual of this species. The von Bertalanffy growth model parameters were L = 211.90 mm and K = 0.319 year-1. The results of the analyses have shown similarities between 1975 and 2008-2010, indicating that the otolith development, the growth pattern and the age structure remained stable. T. lathami is the smallest species of Trachurus and it has the highest growth rates among them. This is probably related to the different temperatures where larvae/juvenile and adult grow, to the absence of a strong fishing pressure and to decadal population variability.

KEY WORDS:
ECOSAR; otolith; sclerochronology; von Bertalanffy

INTRODUCTION

In the Atlantic Ocean, the rough scad, Trachurus lathami Nichols, 1920, is a pelagic species distributed between the United States and North of Argentina, mainly on the continental shelf. It occurs between 50 and 100 m in depth, where it forms schools (Smith-Vaniz 2002Smith-Vaniz WF (2002) Carangidae. In: Carpenter KE (Ed.) The living marine resources of the Western Central Atlantic. FAO, vol. 3, Rome, 1426-1468.). In Brazil, their largest concentrations have been recorded both in the Southeastern Brazilian Bight (SEBB, between 22° and 28°30’S) and in the Southern region (between 28°30’ and 34°S) (Saccardo and Haimovici 2007Saccardo SA, Haimovici M (2007) Síntese sobre o chicharro (Trachurus lathami). In: Haimovici M (Ed.) A prospecção pesqueira e abundância de estoques marinhos no Brasil nas décadas de 1960 a 1990: levantamento de dados e avaliação crítica. Ministério do Meio Ambiente, Secretaria de Mudanças Climáticas e Qualidade Ambiental, Brasília, 233-237.). In these areas, the rough scad is an important fishery resource in purse seines, along with other important pelagic species such as the Sardinella brasiliensis (Steindachner, 1879) and Opisthonema oglinum (Lesueur, 1818) (Saccardo and Haimovici 2007Saccardo SA, Haimovici M (2007) Síntese sobre o chicharro (Trachurus lathami). In: Haimovici M (Ed.) A prospecção pesqueira e abundância de estoques marinhos no Brasil nas décadas de 1960 a 1990: levantamento de dados e avaliação crítica. Ministério do Meio Ambiente, Secretaria de Mudanças Climáticas e Qualidade Ambiental, Brasília, 233-237., UNIVALI 2011UNIVALI (2011) Boletim estatístico da pesca industrial de Santa Catarina - Ano 2010. Universidade do Vale do Itajaí, Centro de Ciências Tecnológicas da Terra e do Mar, Itajaí, 59 pp.).

In the SEBB, rough scad landings varied a lot until the end of the 1990’s (Valentini and Pezzutto 2006Valentini H, Pezzutto PR (2006) Análise das principais pescarias comerciais da região Sudeste/Sul do Brasil com base na produção controlada do período 1986-2004. Instituto Oceanográfico, USP, São Paulo, 56 pp.), a period when this fish was caught in association with the Brazilian sardine (S. brasiliensis) (Fig. 1). When there were fewer sardines, purse seine fleets would capture and land the rough scad (Saccardo et al. 2005Saccardo SA, Cergole MC, Masumoto C (2005) Trachurus lathami. In: Cergole MC, Ávila-da-Silva AA, Rossi-Wongtschowski CLDB (Eds) Análise das principais pescarias comerciais da região Sudeste-Sul: dinâmica das principais espécies em explotação. Instituto Oceanográfico, USP, Série REVIZEE, São Paulo, 156-161.). After this period, sardine landings increased and rough scad landings dropped, without a clear association between them. Between 2005 and 2008, rough scad landings were around 700 t/year, with a later reduction to an average of 88 t (2009-2010) (MMA 2007aMMA (2007a) Estatística da Pesca Brasil - 2005: grandes regiões e unidades da federação. Ministério do Meio Ambiente, Brasília, 105 pp., bMMA (2007b) Estatística da Pesca Brasil - 2007: grandes regiões e unidades da federação. Brasília, Ministério do Meio Ambiente, 113 pp., 2008MMA (2008) Estatística da Pesca Brasil - 2006: grandes regiões e unidades da federação. Ministério do Meio Ambiente, Brasília , 180 pp., UNIVALI 2009UNIVALI (2009) Boletim estatístico da pesca industrial de Santa Catarina - ano 2008: programa de apoio técnico e científico ao desenvolvimento da pesca no Sudeste e Sul do Brasil. Universidade do Vale do Itajaí, Centro de Ciências Tecnológicas da Terra e do Mar, Itajaí, 73 pp., 2011UNIVALI (2011) Boletim estatístico da pesca industrial de Santa Catarina - Ano 2010. Universidade do Vale do Itajaí, Centro de Ciências Tecnológicas da Terra e do Mar, Itajaí, 59 pp., Instituto de Pesca 2013Instituto de Pesca (2013) Estatística Pesqueira. Sistema Gerenciador de Banco de Dados de Controle Estatístico de Produção Pesqueira Marítima, Instituto de Pesca/APTA/SAA/SP. Available online at: Available online at: http://www.pesca.sp.gov.br/estatistica/index.php [Accessed: 20/04/2013]
http://www.pesca.sp.gov.br/estatistica/i...
). Historically, this species has presented intense population fluctuations (Katsuragawa and Ekau 2003Katsuragawa M, Ekau W (2003) Distribution, growth and mortality of young rough scad, Trachurus lathami, in the South-eastern Brazilian Bight. Journal of Applied Ichthyology 19: 21-28. https://doi.org/10.1046/j.1439-0426.2003.00335.x
https://doi.org/10.1046/j.1439-0426.2003...
), oscillations that are also common to the other species of Trachurus Rafinesque, 1810, related to climate and environmental conditions, fishing pressure, biological elements (feeding, reproduction, growth, recruitment) and the combination of those (Arancibia and Neira 2002Arancibia H, Neira S (2002) Does ENSO induce changes in recruitment of horse mackerel (Trachurus symmetricus) and in the long-term trend of the trophic level of fishery landings in central Chile? Investigaciones Marinas 30. https://doi.org/10.4067/S0717-71782002030100072
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, Espino 2013Espino M (2013) El jurel Trachurus murphyi y las variables ambientales de macroescala. Revista Peruana de Biología 20: 9-20. https://doi.org/10.15381/rpb.v20i1.2614
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, Geist et al. 2015Geist SJ, Kunzmann A, Verheye HM, Eggert A, Schukat A, Ekau W (2015) Distribution, feeding behaviour, and condition of Cape horse mackerel early life stages, Trachurus capensis, under different environmental conditions in the northern Benguela upwelling ecosystem. ICES Journal of Marine Science 72: 543-557. https://doi.org/10.1093/icesjms/fsu087
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).

Figure 1
Commercial landings of Trachurus lathami and Sardinella brasiliensis in the Southeastern Brazilian Bight (SEBB). Data of 2008, 2009 and 2010 are restricted to states of Paraná and São Paulo that representing 95% of the total landings. Data source: Valentini and Pezzutto 2006Valentini H, Pezzutto PR (2006) Análise das principais pescarias comerciais da região Sudeste/Sul do Brasil com base na produção controlada do período 1986-2004. Instituto Oceanográfico, USP, São Paulo, 56 pp., MMA 2007aMMA (2007a) Estatística da Pesca Brasil - 2005: grandes regiões e unidades da federação. Ministério do Meio Ambiente, Brasília, 105 pp., bMMA (2007b) Estatística da Pesca Brasil - 2007: grandes regiões e unidades da federação. Brasília, Ministério do Meio Ambiente, 113 pp., 2008MMA (2008) Estatística da Pesca Brasil - 2006: grandes regiões e unidades da federação. Ministério do Meio Ambiente, Brasília , 180 pp. UNIVALI 2009UNIVALI (2009) Boletim estatístico da pesca industrial de Santa Catarina - ano 2008: programa de apoio técnico e científico ao desenvolvimento da pesca no Sudeste e Sul do Brasil. Universidade do Vale do Itajaí, Centro de Ciências Tecnológicas da Terra e do Mar, Itajaí, 73 pp., 2010UNIVALI (2010) Boletim estatístico da pesca industrial de Santa Catarina - ano 2009 e panorama 2000-2009: programa de monitoramento e avaliação da atividade pesqueira industrial no Sudeste e Sul do Brasil. Universidade do Vale do Itajaí, Centro de Ciências Tecnológicas da Terra e do Mar, Itajaí, 97 pp., 2011UNIVALI (2011) Boletim estatístico da pesca industrial de Santa Catarina - Ano 2010. Universidade do Vale do Itajaí, Centro de Ciências Tecnológicas da Terra e do Mar, Itajaí, 59 pp., Instituto de Pesca 2013Instituto de Pesca (2013) Estatística Pesqueira. Sistema Gerenciador de Banco de Dados de Controle Estatístico de Produção Pesqueira Marítima, Instituto de Pesca/APTA/SAA/SP. Available online at: Available online at: http://www.pesca.sp.gov.br/estatistica/index.php [Accessed: 20/04/2013]
http://www.pesca.sp.gov.br/estatistica/i...
.

There is a single stock of the rough scad at the SEBB (Saccardo and Haimovici 2007Saccardo SA, Haimovici M (2007) Síntese sobre o chicharro (Trachurus lathami). In: Haimovici M (Ed.) A prospecção pesqueira e abundância de estoques marinhos no Brasil nas décadas de 1960 a 1990: levantamento de dados e avaliação crítica. Ministério do Meio Ambiente, Secretaria de Mudanças Climáticas e Qualidade Ambiental, Brasília, 233-237.). Its life cycle and fishery (Saccardo 1987Saccardo SA (1987) Morfologia, distribuição e abundância de Trachurus lathami Nichols, 1920 (Teleostei: Carangidae) na região sudeste-sul do Brasil. Boletim do Instituto Oceanográfico 35: 65-95. https://doi.org/10.1590/S0373-55241987000100008
https://doi.org/10.1590/S0373-5524198700...
, Saccardo and Kasturagawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277., Saccardo et al. 2005Saccardo SA, Cergole MC, Masumoto C (2005) Trachurus lathami. In: Cergole MC, Ávila-da-Silva AA, Rossi-Wongtschowski CLDB (Eds) Análise das principais pescarias comerciais da região Sudeste-Sul: dinâmica das principais espécies em explotação. Instituto Oceanográfico, USP, Série REVIZEE, São Paulo, 156-161.); growth (Saccardo and Katsuragawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277.); diet (Meneghetti and Alves 1971Meneghetti JO, Alves CC (1971) Nota preliminar sobre o hábito alimentar de chicharro (Trachurus lathami, Nichols) e seu significado ecológico. Ciência e Cultura 23: 388-389., Carvalho and Soares 2006Carvalho MR, Soares LSH (2006) Diel feeding pattern and diet of rough scad Trachurus lathami Nichols, 1920 (Carangidae) from the Southwestern Atlantic. Neotropical Ichthyology 4: 419-426. https://doi.org/10.1590/S1679-62252006000400005
https://doi.org/10.1590/S1679-6225200600...
); parasites (Braicovich et al. 2012Braicovich PE, Luque JL, Timi J (2012) Geographical patterns of parasite infracommunities in the rough scad, Trachurus lathami Nichols, in the Southwestern Atlantic ocean. The Journal of Parasitology 98: 768-777. https://doi.org/10.1645/GE-2950.1
https://doi.org/10.1645/GE-2950.1...
); and larvae and juveniles (Katsuragawa and Matsuura 1992Katsuragawa M, Matsuura Y (1992) Distribution and abundance of carangid larvae in the South-eastern Brazilian Bight during 1975-81. Boletim do Instituto Oceanográfico 40: 55-78. https://doi.org/10.1590/S0373-55241992000100005
https://doi.org/10.1590/S0373-5524199200...
, Katsuragawa and Ekau 2003Katsuragawa M, Ekau W (2003) Distribution, growth and mortality of young rough scad, Trachurus lathami, in the South-eastern Brazilian Bight. Journal of Applied Ichthyology 19: 21-28. https://doi.org/10.1046/j.1439-0426.2003.00335.x
https://doi.org/10.1046/j.1439-0426.2003...
, Campos et al. 2010Campos PN, Castro MS, Bonecker ACT (2010) Occurrence and distribution of Carangidae larvae (Teleostei, Perciformes) from the Southwest Atlantic Ocean, Brazil (12-23°S). Journal of Applied Ichthyology 26: 920-924. https://doi.org/10.1111/j.1439-0426.2010.01511.x
https://doi.org/10.1111/j.1439-0426.2010...
) have been investigated. Some biological aspects can be summarized from those studies, as follow. The rough scad breeds throughout the year, with a peak between October and December (springtime). The larvae are distributed all over the continental shelf. Individuals grow up to a total length of 200-250 mm (TL), with the first maturation at 115-132 mm TL, living up to 8-9 years. The rough scad is mainly a zooplankton feeder, despite some records of predatory behavior. The otoliths were studied in terms of age and growth (Saccardo and Katsuragawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277.) and morphology (Siliprandi et al. 2014Siliprandi CC, Rossi-Wongtschowski CLDB, Brenha MR, Gonsales SA, Santificetur C, Vaz-dos-Santos AM (2014) Atlas of marine bony fish otoliths (Sagittae) of Southeasthern-Southern Brazil Part II: Perciformes (Carangidae, Sciaenidae, Scombridae and Serranidae). Brazilian Journal of Oceanography 62: 28-100.).

Studies on age and growth provide an essential tool to understand the biology and ecology of fish, providing a foundation for population dynamics assessments (King 2007King M (2007) Fisheries biology, assessment and management. Blackwell Publishing, 2nd ed., Oxford, 382 pp. https://doi.org/10.1002/9781118688038
https://doi.org/10.1002/9781118688038...
). Growth parameters have multiple applications in fishery management (Beverton and Holt 1993Beverton RJH, Holt SJ (1993) On the dynamics of exploited fish populations. Chapman and Hall, London, 783 pp., Froese and Binohlan 2000Froese R, Binohlan C (2000) Empirical relationships to estimate asymptotic length, length at first maturity and length at maximum yield per recruit in fishes, with a simple method to evaluate length frequency data. Journal of Fish Biology 56: 758-773. https://doi.org/10.1006/jfbi.1999.1194
https://doi.org/10.1006/jfbi.1999.1194...
, Sponaugle 2010Sponaugle S (2010) Otolith microstructure reveals ecological and oceanographic processes important to ecosystem-based management. Environmental Biology of Fishes 89: 221-238. https://doi.org/10.1007/s10641-010-9676-z
https://doi.org/10.1007/s10641-010-9676-...
), and their estimates through the analysis of otoliths provide precise and accurate results (Green et al. 2009Green BS, Mapstone BD, Arlos G, Begg GA (2009) Tropical fish otoliths: information for assessment, management and ecology. Springer, New York, 313 pp. https://doi.org/10.1007/978-1-4020-5775-5
https://doi.org/10.1007/978-1-4020-5775-...
). Several studies on age and growth, based on otoliths, have been conducted on different species of Trachurus (Webb and Grant 1979Webb BF, Grant CJ (1979) Age and growth of jack mackerel, Trachurus declivis (Jenyns), from Southeastern Australian Waters. Australian Journal of Marine and Freshwater Research 30: 1-9. https://doi.org/10.1071/MF9790001
https://doi.org/10.1071/MF9790001...
, Horn 1993Horn PL (1993) Growth, age structure, and productivity of jack mackerels (Trachurus spp.) in New Zealand waters. New Zealand Journal of Marine and Freshwater Research 27: 145-155. https://doi.org/10.1080/00288330.1993.9516553
https://doi.org/10.1080/00288330.1993.95...
, Karlou-Riga and Sinis 1997Karlou-Riga C, Sinis A (1997) Age and growth of horse mackerel, Trachurus trachurus (L.) in the Gulf of Saronikos (Greece). Fisheries Research 32: 157-171. https://doi.org/10.1016/S0165-7836(97)00044-1
https://doi.org/10.1016/S0165-7836(97)00...
, Karlou-Riga 2000Karlou-Riga C (2000) Otolith morphology and age and growth of Trachurus mediterraneus (Steindachner) in the Eastern Mediterranean. Fisheries Research 46: 69-82. https://doi.org/10.1016/S0165-7836(00)00134-X
https://doi.org/10.1016/S0165-7836(00)00...
, Araya et al. 2001Araya M, Cubillos LA, Guzman M, Peñailillo J, Sepúlveda A (2001) Evidence of a relationship between age and otolith weight in the Chilean jack mackerel, Trachurus symmetricus murphyi (Nichols). Fisheries Research 51: 17-26. https://doi.org/10.1023/A:1026334532390
https://doi.org/10.1023/A:1026334532390...
, Waldron and Kesrtan 2001Waldron ME, Kerstan M (2001) Age validation in horse mackerel (Trachurus trachurus) otoliths. ICES Journal of Marine Science 58: 806-813. https://doi.org/10.1006/jmsc.2001.1071
https://doi.org/10.1006/jmsc.2001.1071...
, Kasapoglu and Duzgunes 2013Kasapoglu N, Duzgunes E (2013) The relationship between somatic growth and otolith dimensions of Mediterranean horse mackerel (Trachurus mediterraneus) from the Black Sea. Journal of Applied Ichthyology 29: 230-233. https://doi.org/10.1111/jai.12019
https://doi.org/10.1111/jai.12019...
, among others), mainly in view of their commercial relevance (Checkley et al. 2009Checkley D, Alheit J, Oozeki Y, Roy C (2009) Climate change and small pelagic fish. Cambridge University Press, Cambridge , 372 pp. https://doi.org/10.1017/CBO9780511596681
https://doi.org/10.1017/CBO9780511596681...
). From these studies, it is possible to conclude that age and growth are very different among Trachurus species, and that otolith analysis is useful, allowing the comprehension of their biological patterns (Karlou-Riga 2000Karlou-Riga C (2000) Otolith morphology and age and growth of Trachurus mediterraneus (Steindachner) in the Eastern Mediterranean. Fisheries Research 46: 69-82. https://doi.org/10.1016/S0165-7836(00)00134-X
https://doi.org/10.1016/S0165-7836(00)00...
, Abaunza et al. 2003Abaunza P, Gordo L, Murta A, Eltink ATGW, Garc MT (2003) Growth and reproduction of horse mackerel, Trachurus trachurus (Carangidae). Reviews in Fish Biology and Fisheries 13: 27-61. https://doi.org/10.1023/A:1026334532390
https://doi.org/10.1023/A:1026334532390...
).

The significance of the small pelagic fisheries at SEBB led the Brazilian government to promote evaluation and monitoring programs of these resources. Although they were not continuous, the most recent initiative was the ECOSAR Program (Prospection and assessment of the sardine stock biomass in the Southeastern coast by the use of hydro-acoustic methods) carried out between 2008 and 2010 (Cergole and Dias Neto 2011Cergole MC, Dias-Neto J (2011) Plano de gestão de uso sustentável de Sardinha verdadeira (Sardinella brasiliensis) no Brasil. Edições Ibama, Ministério do Meio Ambiente, Brasília, 242 pp.). Even though the target species was S. brasiliensis, other representative species captured during the surveys, among which is T. lathami, were also accounted for. This study aimed to assess the age and growth of T. lathami, to identify its age structure in the Southeastern Brazilian Bight, and to evaluate what causes the wide differences among Trachurus species in terms of body size and growth parameters.

MATERIAL AND METHODS

Four survey cruises were carried out between 20 and 100 m deep with the OV Atlântico Sul by FURG during January-February 2008 (Summer), November 2008 (Spring), September-October 2009 (Spring) and February-March 2010 (Summer) in the Southeastern Brazilian Bight (22°-28°30’S). Transects in perpendicular profiles, oblique to the coast, were followed. The echo sounder (Simrad EK500) was employed and when shoals were detected, both pelagic trawling and purse seine were carried out. Details on the methodology are available in Rossi-Wongtschowski et al. (2014Rossi-Wongtschowski CLDB, Vaz-dos-Santos AM, Siliprandi CC (2014) Checklist of the marine fishes collected during hydroacoustic surveys in the Southeastern Brazilian Bight from 1995 to 2010. Arquivos de Zoologia 45: 73-88. https://doi.org/10.11606/issn.2176-7793.v45iespp73-88
https://doi.org/10.11606/issn.2176-7793....
). Trachurus lathami were caught in 17 fishing operations (Fig. 2). On board, samples were frozen. In the laboratory, the total length (TL, mm) of individuals was measured and their sagittae otoliths were removed, washed, dried and stored in microtubes (FAO 1981FAO (1981) Methods of collecting and analyzing size and age data for fish stock assessment. FAO Fisheries Circular 736: 1-100.).

Figure 2
Map of the Southeastern Brazilian Bight (SEBB) showing the locations of fishing hauls with Trachurus lathami catches by period (n = 17, some points are overlapped in the map).

Whenever possible, ten otoliths of T. lathami were selected from each survey by total length class (10 mm) (Araya et al. 2001Araya M, Cubillos LA, Guzman M, Peñailillo J, Sepúlveda A (2001) Evidence of a relationship between age and otolith weight in the Chilean jack mackerel, Trachurus symmetricus murphyi (Nichols). Fisheries Research 51: 17-26. https://doi.org/10.1023/A:1026334532390
https://doi.org/10.1023/A:1026334532390...
). Images of the entire left otolith under water (Fig. 3) were obtained using a stereomicroscope coupled with an image analyzer, and the length (measurement of the horizontal projection of its ends in relation to the longer axis - OL, mm) and height (measurement of the vertical projection in relation to the higher axis of the structure - OH, mm) of the otolith were recorded. The weight of the otolith (OW, g) was obtained with an analytical balance. In order to describe otolith growth in relation to body growth, the allometric (potential) model (Huxley 1993Huxley JS (1993) Problems of relative growth. The John Hopkins University Press, 2nd ed., Baltimore, 276 pp.) was used to fit regressions among the total length and the measurements of the otoliths (Vaz-dos-Santos 2015aVaz-dos-Santos AM (2015a) Métodos quantitativos aplicados ao estudo de otólitos. In: Volpedo AV, Vaz-dos-Santos AM (Eds) Métodos de estudos com otólitos: princípios e aplicações. CAFP-BA-PIESCI, Buenos Aires, 377-395.). The adequacy of the regressions was checked by the values of the coefficient of determination (r2) and the residual analysis (Bervian et al. 2006Bervian G, Fontoura NF, Haimovici M (2006) Statistical model of variable allometric growth: otolith growth in Micropogonias furnieri (Actinopterygii, Scianidae). Journal of Fish Biology 68: 196-208. https://doi.org/10.1111/j.1095-8649.2005.00890.x
https://doi.org/10.1111/j.1095-8649.2005...
, Vaz-dos-Santos and Rossi-Wongtschowski 2013Vaz-dos-Santos AM, Rossi-Wongtschowski CLDB (2013) Length-weight relationships of the ichthyofauna associated with the Brazilian sardine, Sardinella brasiliensis, on the Southeastern Brazilian Bight (22°S-29°S) between 2008 and 2010. Biota Neotropica 13: 326-330. http://www.biotaneotropica.org.br/v13n2/en/abstract?short-communication+bn01613022013
http://www.biotaneotropica.org.br/v13n2/...
).

Figure 3
Trachurus lathami: external surface of a left otolith showing measurements (OL = otolith length and OH = otolith height) and ring analysis (M and L are rings formed before the first annual one; more details in the text). Scale bar: 1 mm.

In order to study age and growth, the annual growth zones of the otoliths (macrostructural analysis) were analyzed (Brothers 1987Brothers EB (1987) Methodological approaches to the examination of otoliths in aging studies. In: Summerfelt RC, Hall GH (Eds) Age and growth of fish. Iowa State University Press, Iowa, 319-330.). Whole otolith images were used to identify the central opaque nucleus and, from this, in the posterior axis to the posterior edge (otolith radius, Ro, mm), the translucent zones. The complete and continuous translucent zones were counted and measured from the beginning of their formation (Fig. 3) (Saccardo and Katsuragawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277., Karlou-Riga and Sinis 1997Karlou-Riga C, Sinis A (1997) Age and growth of horse mackerel, Trachurus trachurus (L.) in the Gulf of Saronikos (Greece). Fisheries Research 32: 157-171. https://doi.org/10.1016/S0165-7836(97)00044-1
https://doi.org/10.1016/S0165-7836(97)00...
, Waldron and Kerstan 2001Waldron ME, Kerstan M (2001) Age validation in horse mackerel (Trachurus trachurus) otoliths. ICES Journal of Marine Science 58: 806-813. https://doi.org/10.1006/jmsc.2001.1071
https://doi.org/10.1006/jmsc.2001.1071...
). Three blind readings were carried out by the same reader, whose consistency was checked by the average percentage error (APE) (Beamish and Fournier 1981Beamish RJ, Fournier DA (1981) A method for comparing the precision of a set of age determinations. Canadian Journal of Fisheries and Aquatic Sciences 38: 982-983. https://doi.org/10.1139/f81-132
https://doi.org/10.1139/f81-132...
) and the value of the coefficient of variation (CV) (Campana 2001Campana SE (2001) Accuracy, precision and quality control in age determination, including a review of the use and abuse of age validation methods. Journal of Fish Biology 59: 197-242. https://doi.org/10.1006/jfbi.2001.1668
https://doi.org/10.1006/jfbi.2001.1668...
). In order to check the consistency of the ring analysis, box plot and constancy analysis were applied (Vaz-dos-Santos 2015bVaz-dos-Santos AM (2015b) Otólitos em estudos de idade e crescimento em peixes. In: Volpedo AV, Vaz-dos-Santos AM (Eds) Métodos de estudos com otólitos: princípios e aplicações. CAFP-BA-PIESCI, Buenos Aires, 303-333.). A box plot of the ring radius by the ring (category) was built in order to check whether there is overlap among measurements. After checking the assumptions, ANOVA followed by post-hoc Tukey test were performed to compare the averages of the radius. A constancy analysis (scatter plot between total length and the rings radius) was applied, and the linearity of each ring group was tested using regression analysis (except for rings 7 and 8, with a narrow range of total length). All statistical procedures followed Zar (2010Zar JH (2010) Biostatistical analysis. Pearson, 5th ed., New Jersey, 944 pp.) with 5% of significance. The ring counts and edge patterns allowed age attribution, since the formation of an annual ring had been previous validated, occurring between October and December (Saccardo and Katsuragawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277.). Two rings are formed during the first year of life, before the first annual ring (Saccardo and Katsuragawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277.). Whenever possible, these rings were recorded, but they were not used in any analysis.

The von Bertalanffy growth model parameters (VBGM) (L, K, t0) were estimated from observed lengths per age and by the average lengths per age using the least-squares iterative method (Aubone and Wöhler 2000Aubone A, Wöhler OC (2000) Aplicación del método de máxima verosimilitud a la estimación y comparación de curvas de creci miento de von Bertalanffy. INIDEP Informe Técnico 37: 1-21.). From the growth parameters, the inverse VBGM was used to estimate the age of the entire sample of T. lathami. Although the value of the theoretical age at zero length has been estimated, it was not used to estimate the age with the inverse VBGM. Next, an age length key (frequencies of individuals by age and TL class) was built and the space-time distributions by ages were analyzed in the surveyed periods using maps.

RESULTS

The total length of 1,312 T. lathami individuals sampled varied between 27 mm and 208 mm, resulting in bimodal distributions in each period surveyed, one of juveniles (20 to 60 mm) and a second mode composed of adults (>115 mm TL), which predominated in the seasons analyzed (Fig. 4). The few numbers of fish sampled between 100-140 mm TL resulted from the selectivity of the fishing gear employed.

Figure 4
Trachurus lathami: length frequency distribution by period (dashed line indicates the average length of first maturity, L50 = 115 mm).

Following the selection criteria, 282 otoliths of T. lathami were analyzed (it was not always possible to attain ten otoliths from each survey by TL class). The regressions among the otolith measurements (Table 1) represented properly the development of the structure (p < 0.001 in the three regressions, 0.970 < r2 < 0.979), thus enabling the use of the otoliths in the growth study. In the adjusted models (TL vs. OL, TL vs. OH, TL vs. OW), residuals did not show any bias. The best fit was presented by the TL vs. OL model (r2 = 0.979), showing that the otolith length provides the best representation of fish growth, following the anterior-posterior axis.

Table 1
Trachurus lathami: coefficients of potential regressions between total length (TL, mm) and otolith length (OL, mm), height (OH, mm) and weight (OW, g). (r2) Coefficient of determination, n = 282.

After three readings, 278 otoliths were considered legible (98.6%) and four non legible (1.4%), which were not considered in the following analysis. The average percentage error among the three readings was 4.1%, and the coefficient of variation was 5.4%. Otoliths with up to eight rings were observed with high precision and consistency, revealed by the box plot and constancy analysis (Figs 5, 6). In relation to the box plot analysis, the average radius of rings was significantly different (ANOVA F = 1527.76, p < 0.001), except for rings 7 and 8 (p > 0.05). Linearity tested in the constancy analysis was not significant in all cases (p > 0.05). Fish with only two rings were not found. The age determination resulted in fishes with age zero up to eight years old.

Figures 5-6
Trachurus lathami: (5) Box plot of ring radius (whiskers = minimum and maximum, box = interquartile range, bar = mean). (6) Constancy graph showing the position of rings radius (R) against total length (TL) (n = 278).

The von Bertalanffy growth model parameters were L = 211.90 mm, K = 0.319 year-1 and t0 = -0.576 years for observed lengths per age, and L = 206.31 mm, K = 0.336 year-1 and t0 = -0.578 years for average lengths per age. When considering the maximum length obtained in the sampling (208 mm), the parameters estimated through the average lengths were disregarded, and the parameters fitted from observed lengths per age were adopted to estimate fish age (Fig. 7). The sample had fish from 0 to 8 years old or above (Table 2).

Figure 7
Trachurus lathami: von Bertalanffy growth curve fitted (line) to observed total lengths (points) by age (n = 278).

Table 2
Trachurus lathami: age-length key in the Southeastern Brazilian Bight during 2008-2010: percentage (from the total) of individuals by length class and age; number of ages estimated through otolith readings and with the inverse von Bertalanffy Growth Model (VBGM).

In relation to the space-time distributions, in the summer of 2008, the rough scad schools were concentrated closer to the coast at 27°-28°S (89%), with all ages represented, and a predominance of one-year old fish (Fig. 8), associated with the Cabo Frio upwelling (22°-23°S), around 100 m isobath. Another school was found at the continental shelf 24°-25°S (Fig. 9), similar to the summer of 2009 (Fig. 10). In the summer of 2010, the rough scad was more concentrated between 23°-24°S, and its school was composed mainly of 4-year old fish, or older (Fig. 11).

Figures 8-11
Trachurus lathami: space-time distribution of age groups in the Southeastern Brazilian Bight during (8) summer of 2008 (January-February), (9) spring of 2008 (November), (10) spring of 2009 (September-October) and (11) summer of 2010 (February-March).

DISCUSSION

The growth parameters estimated for T. lathami in this study are the most recent after 1975 (Saccardo and Katsuragawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277.). They originate from the only biological data available for this species in the Southwestern Atlantic (Cergole and Dias Neto 2011Cergole MC, Dias-Neto J (2011) Plano de gestão de uso sustentável de Sardinha verdadeira (Sardinella brasiliensis) no Brasil. Edições Ibama, Ministério do Meio Ambiente, Brasília, 242 pp.), and represent the population and the sampling adequately, due to the total length range analyzed. Previous information on the presence of up to 400 mm TL rough scads at the SEBB (Menezes and Figueiredo 1980Menezes NA, Figueiredo JL (1980) Manual de peixes marinhos do sudeste do Brasil. IV. Teleostei (3). Museu de Zoologia da Universidade de São Paulo, São Paulo, 96 pp.) denotes very rare individuals, never recorded again. Historically, the longer total lengths recorded for the rough scad in the SEBB were 260 mm (Saccardo 1987Saccardo SA (1987) Morfologia, distribuição e abundância de Trachurus lathami Nichols, 1920 (Teleostei: Carangidae) na região sudeste-sul do Brasil. Boletim do Instituto Oceanográfico 35: 65-95. https://doi.org/10.1590/S0373-55241987000100008
https://doi.org/10.1590/S0373-5524198700...
), 207 mm (Figueiredo et al. 2002Figueiredo JL, Santos AP, Yamaguti N, Bernardes RA, Rossi-Wongtschowski CLDB (2002) Peixes da Zona Econômica Exclusiva do Sudeste e Sul do Brasil. Levantamento com rede de meia água. Edusp, São Paulo , 242 pp.), 240 mm (Saccardo et al. 2005Saccardo SA, Cergole MC, Masumoto C (2005) Trachurus lathami. In: Cergole MC, Ávila-da-Silva AA, Rossi-Wongtschowski CLDB (Eds) Análise das principais pescarias comerciais da região Sudeste-Sul: dinâmica das principais espécies em explotação. Instituto Oceanográfico, USP, Série REVIZEE, São Paulo, 156-161.) and 261 mm (Bernardes et al. 2005Bernardes RA, Rossi-Wongtschowski CLDB, Wahrlich R, Vieira RC, Santos AP, Rodrigues AR (2005) Prospecção pesqueira de recursos demersais com armadilhas e pargueiras na Zona Econômica Exclusiva da região Sudeste-Sul do Brasil. Instituto Oceanográfico, USP, São Paulo, 112 pp.). In all these studies, as well as in our results, individuals larger than 200 mm were seldom recorded. This indicates that T. lathami in the study area have smaller size structure (usually up to 180 mm SL) than in the Caribbean Sea (330 mm SL) (Smith-Vaniz 2002Smith-Vaniz WF (2002) Carangidae. In: Carpenter KE (Ed.) The living marine resources of the Western Central Atlantic. FAO, vol. 3, Rome, 1426-1468.), but similar size structure to that found for the Uruguayan and Argentinean coasts (between 80-230 mm TL) (Cousseau and Perrotta 2004Cousseau MB, Perrotta RG (2004) Peces marinos de Argentina. Instituto Nacional de Investigaión y Desarrollo Pesquero, Mar del Plata, 167 pp.). Thus, the size structure analyzed in this work was representative of the stock, even in the absence of larger individuals.

The development of the otoliths of T. lathami had only been analyzed by Saccardo and Katsuragawa (1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277.), whose details are available only in the unpublished thesis of Saccardo from 1980. Using these sources and Saccardo (1987Saccardo SA (1987) Morfologia, distribuição e abundância de Trachurus lathami Nichols, 1920 (Teleostei: Carangidae) na região sudeste-sul do Brasil. Boletim do Instituto Oceanográfico 35: 65-95. https://doi.org/10.1590/S0373-55241987000100008
https://doi.org/10.1590/S0373-5524198700...
), the visual inspection of scatter plots of the total length of individuals and otolith measurements (regression models were not adjusted in these former studies), it was possible to verify that otolith development has kept a similar pattern between the 1975 and 2008-2010. Namely, OL, OH and OW in relation to TL, respectively, showed similar values. In the present results, the allometric models and residual analysis did not show noticeable changes in the pattern of otolith development, indicating a single-phase growth (Bervian et al. 2006Bervian G, Fontoura NF, Haimovici M (2006) Statistical model of variable allometric growth: otolith growth in Micropogonias furnieri (Actinopterygii, Scianidae). Journal of Fish Biology 68: 196-208. https://doi.org/10.1111/j.1095-8649.2005.00890.x
https://doi.org/10.1111/j.1095-8649.2005...
). Previously, it was evidenced a change in otolith thickness (two phases of otolith development) close to 130 mm TL (Saccardo and Katsuragawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277.), due to the first gonadal maturation (Saccardo et al. 2005Saccardo SA, Cergole MC, Masumoto C (2005) Trachurus lathami. In: Cergole MC, Ávila-da-Silva AA, Rossi-Wongtschowski CLDB (Eds) Análise das principais pescarias comerciais da região Sudeste-Sul: dinâmica das principais espécies em explotação. Instituto Oceanográfico, USP, Série REVIZEE, São Paulo, 156-161.), which was not evidenced here probably due to the few numbers of individuals between 100-140 mm TL.

The quantitative elements (APE, CV, box plot, constancy analysis) used to evaluate otolith readings indicated that the ring radius presented high precision and does not vary much (Campana 2001Campana SE (2001) Accuracy, precision and quality control in age determination, including a review of the use and abuse of age validation methods. Journal of Fish Biology 59: 197-242. https://doi.org/10.1006/jfbi.2001.1668
https://doi.org/10.1006/jfbi.2001.1668...
). Accuracy was also high, i.e., the position of the rings in the otoliths was similar to values reported previously (Saccardo and Katsuragawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277.). These authors described the formation of two rings adjacent to the nucleus in 13% and 28% of the 1,908 otoliths analyzed, respectively. The material in this study hardly presented those rings and no clear patterns (at least in the macrostructural analysis); therefore, they were not taken into consideration. To investigate them, a microstructural analysis of the otoliths of T. lathami needs to be carried out.

The use of whole otoliths, sectioned otoliths or both, to count rings and to estimate age has been broadly discussed (Webb and Grant 1979Webb BF, Grant CJ (1979) Age and growth of jack mackerel, Trachurus declivis (Jenyns), from Southeastern Australian Waters. Australian Journal of Marine and Freshwater Research 30: 1-9. https://doi.org/10.1071/MF9790001
https://doi.org/10.1071/MF9790001...
, Karlou-Riga and Sinis 1997Karlou-Riga C, Sinis A (1997) Age and growth of horse mackerel, Trachurus trachurus (L.) in the Gulf of Saronikos (Greece). Fisheries Research 32: 157-171. https://doi.org/10.1016/S0165-7836(97)00044-1
https://doi.org/10.1016/S0165-7836(97)00...
, Stewart and Ferrell 2001Stewart J, Ferrell DJ (2001) Age, growth, and commercial landings of yellowtail scad (Trachurus novaezelandiae) and blue mackerel (Scomber australasicus) off the coast of New South Wales, Australia. New Zealand Journal of Marine and Freshwater Research 35: 541-551. https://doi.org/10.1080/00288330.2001.9517021
https://doi.org/10.1080/00288330.2001.95...
, Waldron and Kerstan 2001Waldron ME, Kerstan M (2001) Age validation in horse mackerel (Trachurus trachurus) otoliths. ICES Journal of Marine Science 58: 806-813. https://doi.org/10.1006/jmsc.2001.1071
https://doi.org/10.1006/jmsc.2001.1071...
, Kerkich et al. 2013Kerkich M, Aksissou M, Casal JAE (2013) Age and growth of the horse mackerel Trachurus trachurus (Linnaeus,1758) catches in the bay of M’diq (Mediterraneen coast of Morocco). IRACST - Engineering Science and Technology: An International Journal 3: 708-714., Costa 2004Costa AM (2004) Idade e crescimento do carapau (Trachurus trachurus L.) da costa portuguesa no período de 1992 a 1998. IPIMAR, Lisboa, 25 pp., Dioses 2013Dioses T (2013) Edad y crecimiento del jurel Trachurus murphyi, (Nichols 1920) en el Perú. Revista Peruana de Biología 20: 45-52., Goicochea et al. 2013Goicochea C, Mostacero J, Moquillaza P, Dioses T, Topiño Y, Guevara-Carrasco R (2013) Validación del ritmo de formación de los anillos de crecimiento en otolitos del jurel Trachurus murphyi Nichols 1920. Revista Peruana de Biología 20: 053-060. https://doi.org/10.15381/rpb.v20i1.2619
https://doi.org/10.15381/rpb.v20i1.2619...
, among others). Previous studies on Trachurus postulated that, when using whole otoliths, the following considerations need to be taken into account: (i) whole otoliths are suitable for estimating the age of fish smaller than 250 mm TL (Costa 2004Costa AM (2004) Idade e crescimento do carapau (Trachurus trachurus L.) da costa portuguesa no período de 1992 a 1998. IPIMAR, Lisboa, 25 pp.); (ii) only complete, continuous and thicker rings should be counted (Dioses 2013Dioses T (2013) Edad y crecimiento del jurel Trachurus murphyi, (Nichols 1920) en el Perú. Revista Peruana de Biología 20: 45-52.); (iii) the precision (sensu Campana 2001Campana SE (2001) Accuracy, precision and quality control in age determination, including a review of the use and abuse of age validation methods. Journal of Fish Biology 59: 197-242. https://doi.org/10.1006/jfbi.2001.1668
https://doi.org/10.1006/jfbi.2001.1668...
) must be acceptable (Lyle et al. 2000Lyle JM, Krusic-Golub K, Morison AK (2000) Age and growth of jack mackerel and the age structure of the jack mackerel purse seine catch. Tasmanian Aquaculture and Fisheries Institute, Taroona, 49 pp.); (iv) otoliths of adults must not show multi-ring formation (Karlou-Riga and Sinis 1997Karlou-Riga C, Sinis A (1997) Age and growth of horse mackerel, Trachurus trachurus (L.) in the Gulf of Saronikos (Greece). Fisheries Research 32: 157-171. https://doi.org/10.1016/S0165-7836(97)00044-1
https://doi.org/10.1016/S0165-7836(97)00...
). These four considerations can be safely applied to our data and results. Actually, the analysis of whole otoliths in Trachurus is not recommended in older fish with thicker otoliths (Horn 1993Horn PL (1993) Growth, age structure, and productivity of jack mackerels (Trachurus spp.) in New Zealand waters. New Zealand Journal of Marine and Freshwater Research 27: 145-155. https://doi.org/10.1080/00288330.1993.9516553
https://doi.org/10.1080/00288330.1993.95...
, Karlou-Riga and Sinis 1997Karlou-Riga C, Sinis A (1997) Age and growth of horse mackerel, Trachurus trachurus (L.) in the Gulf of Saronikos (Greece). Fisheries Research 32: 157-171. https://doi.org/10.1016/S0165-7836(97)00044-1
https://doi.org/10.1016/S0165-7836(97)00...
, Karlou-Riga 2000Karlou-Riga C (2000) Otolith morphology and age and growth of Trachurus mediterraneus (Steindachner) in the Eastern Mediterranean. Fisheries Research 46: 69-82. https://doi.org/10.1016/S0165-7836(00)00134-X
https://doi.org/10.1016/S0165-7836(00)00...
, Lyle et al. 2000Lyle JM, Krusic-Golub K, Morison AK (2000) Age and growth of jack mackerel and the age structure of the jack mackerel purse seine catch. Tasmanian Aquaculture and Fisheries Institute, Taroona, 49 pp.). In the case of T. lathami, a smaller species than its congenerics, this was not an issue. Added to this, the present study was also looking for comparisons with the previously one (Saccardo and Katsuragawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277.).

The lack of self-validation, which was not possible in view of our sample design, does not compromise our results. The previously adopted validation was correctly performed, based on 399 otoliths (21% of the total) with coincident ring analysis by Saccardo and Katsuragawa (1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277.). It was based on the percentage of the edge pattern (Panfili and Morales Nin 2002Panfili J, Morales Nin B (2002) Semi-direct validation. In: Panfili J, Pontual H, Troadec H, Wright PJ (Eds) Manual of fish sclerochronology. Éditions Ifremer, Brest, 463 pp.) and on the average lengths by age (Campana 2001Campana SE (2001) Accuracy, precision and quality control in age determination, including a review of the use and abuse of age validation methods. Journal of Fish Biology 59: 197-242. https://doi.org/10.1006/jfbi.2001.1668
https://doi.org/10.1006/jfbi.2001.1668...
), methods that are accepted for validation. Moreover, the resemblance found in the regressions and in the ring analysis between 1975 (Saccardo and Katsuragawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277.) and 2008-2010 (present study) suggests that the development pattern of the otolith has remained stable. This could also be considered in the case of growth ring formation.

The growth parameters (L, K) estimated with observed lengths per age were most suitable to describe the growth of T. lathami, since that obtained from the average lengths per age underestimated the maximum length values. The estimated theoretical age at zero length, is inconsistent in biological terms, since the rough scad hatches around 1.5 mm long (Katsuragawa and Ekau 2003Katsuragawa M, Ekau W (2003) Distribution, growth and mortality of young rough scad, Trachurus lathami, in the South-eastern Brazilian Bight. Journal of Applied Ichthyology 19: 21-28. https://doi.org/10.1046/j.1439-0426.2003.00335.x
https://doi.org/10.1046/j.1439-0426.2003...
), corresponding to seven days of age using the VBGM adjusted in this study. This value seldom has a biological meaning, since it would represent the zero length age whenever the growth continues following the same pattern described by the equation (Pauly 1984Pauly D (1984) Fish population dynamics in tropical waters: a manual for use with programmable calculators. ICLARM, Manila, 325 pp.), which is not true. Several authors (Abaunza et al. 2003Abaunza P, Gordo L, Murta A, Eltink ATGW, Garc MT (2003) Growth and reproduction of horse mackerel, Trachurus trachurus (Carangidae). Reviews in Fish Biology and Fisheries 13: 27-61. https://doi.org/10.1023/A:1026334532390
https://doi.org/10.1023/A:1026334532390...
, Yankova et al. 2010Yankova MH, Raykov VS, Gerdzhikov DB, Frateva PB (2010) Growth and length-weight relationships of the horse mackerel, Trachurus mediterraneus ponticus (Aleev, 1956), off the Bulgarian Black Sea coast. Turkish Journal of Zoology 34: 85-92. https://doi.org/10.3906/zoo-0811-10
https://doi.org/10.3906/zoo-0811-10...
, Yoda et al. 2014Yoda M, Shiraishi T, Yukami R, Ohshimo S (2014) Age and maturation of jack mackerel Trachurus japonicus in the East China Sea. Fisheries Science 80: 61-68. https://doi.org/10.1007/s12562-013-0687-5
https://doi.org/10.1007/s12562-013-0687-...
), working on Trachurus, also found inconsistent t0 values due to the lack of small fish in their samples. In the present study, the t0 value led to overestimated ages, and it should not be used.

Comparison of the growth parameters obtained here with previous estimates in Brazil (Table 3) showed that the growth pattern and age structure of T. lathami have been stable (Saccardo and Katsuragawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277., Saccardo et al. 2005Saccardo SA, Cergole MC, Masumoto C (2005) Trachurus lathami. In: Cergole MC, Ávila-da-Silva AA, Rossi-Wongtschowski CLDB (Eds) Análise das principais pescarias comerciais da região Sudeste-Sul: dinâmica das principais espécies em explotação. Instituto Oceanográfico, USP, Série REVIZEE, São Paulo, 156-161.). However, the rough scad has historically presented seasonal displacements within the SEBB, resulting in variations in the space-time distribution of the individuals in each age (Saccardo 1987Saccardo SA (1987) Morfologia, distribuição e abundância de Trachurus lathami Nichols, 1920 (Teleostei: Carangidae) na região sudeste-sul do Brasil. Boletim do Instituto Oceanográfico 35: 65-95. https://doi.org/10.1590/S0373-55241987000100008
https://doi.org/10.1590/S0373-5524198700...
, Saccardo and Haimovici 2007Saccardo SA, Haimovici M (2007) Síntese sobre o chicharro (Trachurus lathami). In: Haimovici M (Ed.) A prospecção pesqueira e abundância de estoques marinhos no Brasil nas décadas de 1960 a 1990: levantamento de dados e avaliação crítica. Ministério do Meio Ambiente, Secretaria de Mudanças Climáticas e Qualidade Ambiental, Brasília, 233-237.). Except for the summer of 2008, the typical spring-summer pattern (September to March), when the rough scad schools usually concentrated between 23° and 26°S for spawning, was observed. Such movement coincides with the most productive period in terms of water column eutrophication (Castro et al. 2006Castro BM, Lorenzetti JA, Silveira ICA, Miranda LB (2006) Estrutura termohalina e circulação na região entre o Cabo de São Tomé (RJ) e o Chuí (RS). In: Rossi-Wongtschowski CLDB, Madureira LSP (Eds) O ambiente oceanográfico da plataforma continental e do talude na região Sudeste-Sul do Brasil. Edusp, São Paulo , 11-120.) and biological productivity (Pires-Vanin et al. 1993Pires-Vanin AMS, Rossi-Wongtschowski CLDB, Aidar E, Mesquita HSL, Soares LSH, Katsuragawa M, Matsuura Y (1993) Estrutura e função do ecossistema de plataforma continental do Atlântico Sul brasileiro: síntese dos resultados. Publicação especial do Instituto Oceanográfico 10: 217-231.), which favors the rough scad in terms of their feeding habits (Carvalho and Soares 2006Carvalho MR, Soares LSH (2006) Diel feeding pattern and diet of rough scad Trachurus lathami Nichols, 1920 (Carangidae) from the Southwestern Atlantic. Neotropical Ichthyology 4: 419-426. https://doi.org/10.1590/S1679-62252006000400005
https://doi.org/10.1590/S1679-6225200600...
) and conditions for larval growth (Katsuragawa and Matsuura 1992Katsuragawa M, Matsuura Y (1992) Distribution and abundance of carangid larvae in the South-eastern Brazilian Bight during 1975-81. Boletim do Instituto Oceanográfico 40: 55-78. https://doi.org/10.1590/S0373-55241992000100005
https://doi.org/10.1590/S0373-5524199200...
).

Table 3
Trachurus lathami: growth parameters estimated and average total length at age for the species in the Southeastern Brazilian Bight.

Comparison with the nine congeneric species, considering the parameters of VBGM for sex pooled and based on the total length available at the FishBase (Froese and Pauly 2015Froese R, Pauly D (2015) FishBase. Available online at: Available online at: http://www.fishbase.org , version 04/2015 [Accessed 24/06/2015].
http://www.fishbase.org...
), confirmed that the age and growth of species of the genus are broadly different (Webb and Grant 1979Webb BF, Grant CJ (1979) Age and growth of jack mackerel, Trachurus declivis (Jenyns), from Southeastern Australian Waters. Australian Journal of Marine and Freshwater Research 30: 1-9. https://doi.org/10.1071/MF9790001
https://doi.org/10.1071/MF9790001...
, Araya et al. 2001Araya M, Cubillos LA, Guzman M, Peñailillo J, Sepúlveda A (2001) Evidence of a relationship between age and otolith weight in the Chilean jack mackerel, Trachurus symmetricus murphyi (Nichols). Fisheries Research 51: 17-26. https://doi.org/10.1023/A:1026334532390
https://doi.org/10.1023/A:1026334532390...
). Trachurus lathami presented the lowest value of the maximum theoretical length (L = 211.90 mm), together with other lower estimates of this parameter for the species in Brazil (previously mentioned in the text). Although the inverse relationship between L and K is well known, some considerations about this can be done, as follows.

Worldwide, environmental conditions play an important role in the growth of Trachurus species (Geist et al. 2013Geist SJ, Ekau W, Kunzmann A (2013) Energy demand of larval and juvenile Cape horse mackerels, Trachurus capensis, and indications of hypoxia tolerance as benefit in a changing environment. Marine Biology 160: 3221-3232. https://doi.org/10.1007/s00227-013-2309-2
https://doi.org/10.1007/s00227-013-2309-...
, Sassa et al. 2014Sassa C, Takahashi M, Nishiuchi K, Tsukamoto Y (2014) Distribution, growth and mortality of larval jack mackerel Trachurus japonicus in the southern East China Sea in relation to oceanographic conditions. Journal of Plankton Research 36: 542-556. https://doi.org/10.1093/plankt/fbt134
https://doi.org/10.1093/plankt/fbt134...
). In the FishBase (Froese and Pauly 2015Froese R, Pauly D (2015) FishBase. Available online at: Available online at: http://www.fishbase.org , version 04/2015 [Accessed 24/06/2015].
http://www.fishbase.org...
), the growth parameters of other Trachurus species were associated with low water temperature (average = 14.4 °C, n = 50). This indicates that most of these species grow in cold waters, which also applies to most T. lathami. The temperature of the SEBB continental shelf is usually higher than 20 °C (Castro et al. 2006Castro BM, Lorenzetti JA, Silveira ICA, Miranda LB (2006) Estrutura termohalina e circulação na região entre o Cabo de São Tomé (RJ) e o Chuí (RS). In: Rossi-Wongtschowski CLDB, Madureira LSP (Eds) O ambiente oceanográfico da plataforma continental e do talude na região Sudeste-Sul do Brasil. Edusp, São Paulo , 11-120.) and values lower than 20 °C are restricted to the upwelling areas, mainly in the spring and summer (October to March) (Braga and Niencheski 2006Braga ES, Niencheski LFH (2006) Composição de massas de água e seus respectivos potenciais produtivos na área entre o Cabo de São Tomé (RJ) e o Chuí (RS). In: Rossi-Wongtschowski CLDB, Madureira LSP (Eds) O ambiente oceanográfico da plataforma continental e do talude na região Sudeste-Sul do Brasil. Edusp, São Paulo, 161-218.), when the spawning and initial growth of T. lathami larvae take place (Saccardo and Katsuragawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277.). On the other hand, part of the adult stock can be found in deeper and colder waters (11.5-18.5 °C) (Haimovici et al. 2008Haimovici M, Rossi-Wongtschowski CLDB, Bernardes RA, Fischer LG, Vooren CM, Santos RA, Rodrigues AR, Santos S (2008) Prospecção pesqueira de espécies demersais com rede de arrasto-de-fundo na Região Sudeste-Sul do Brasil. Instituto Oceanográfico, USP, São Paulo, 183 pp.). Larvae and juveniles grow in different environmental conditions of the adults, fact that may cause variations in the length and age structure, which could not be evaluated here.

Another factor that may explain the differences in body size and growth parameters (between 1975 and 2008-2010, and in relation to the congeneric species) is fishing. In Brazil, during the 1970s, the fishing pressure on T. lathami was quite strong (Saccardo and Katsuragawa 1995Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277., Saccardo et al. 2005Saccardo SA, Cergole MC, Masumoto C (2005) Trachurus lathami. In: Cergole MC, Ávila-da-Silva AA, Rossi-Wongtschowski CLDB (Eds) Análise das principais pescarias comerciais da região Sudeste-Sul: dinâmica das principais espécies em explotação. Instituto Oceanográfico, USP, Série REVIZEE, São Paulo, 156-161.), but this has changed: since 1997, the species has been fished accidentally, as accessory fauna (Saccardo and Haimovici 2007Saccardo SA, Katsuragawa M (1995) Biology of the rough scad Trachurus lathami, on the southeastern coast of Brazil. Scientia Marina 59: 265-277.). All other Trachurus species, on the other hand, are target species of fisheries, and are under an intense pressure (Araya et al. 2001Araya M, Cubillos LA, Guzman M, Peñailillo J, Sepúlveda A (2001) Evidence of a relationship between age and otolith weight in the Chilean jack mackerel, Trachurus symmetricus murphyi (Nichols). Fisheries Research 51: 17-26. https://doi.org/10.1023/A:1026334532390
https://doi.org/10.1023/A:1026334532390...
). Usually, growth parameters are influenced by density dependent and independent causes (Cardoso and Haimovici 2011Cardoso LG, Haimovici M (2011) Age and changes in growth of the king weakfish Macrodon atricauda (Günther, 1880) between 1977 and 2009 in southern Brazil. Fisheries Research 111: 177-187. https://doi.org/10.1016/j.fishres.2011.06.017
https://doi.org/10.1016/j.fishres.2011.0...
). In the present case, the reduction in body size and the increase in growth rates could be due to an increase in the density of the T. lathami population, which apparently has not been overfished in the period analyzed.

Decadal shifts in the populations of pelagic fish (Alheit et al. 2009Alheit J, Roy C, Kifani S (2009) Decadal-scale variability in popu lations. In: Checkley D, Alheit J, Oozeki Y, Roy C (Eds) Climate change and small pelagic fish. Cambridge University Press, Cambridge, 64-87. https://doi.org/10.1017/CBO9780511596681.007
https://doi.org/10.1017/CBO9780511596681...
), similar to the idea suggested by the visual aspect in Fig. 1 (although it only expresses the landings instead of abundance, density or biomass), deserve further inspection. The growth parameters estimated here are essential elements for the management of small pelagic species at SEBB, and this should not be ignored in future assessments. Besides, the fishing statistics in Brazil are limited and there is no tradition of sampling biological landings. The lack of biological data series is notorious. As long as such deplorable historic situation persists, basic and fundamental studies of auto-ecology like this will remain scanty. Undoubtedly, the next study on the age and growth of the T. lathami cannot wait 40 years.

ACKNOWLEDGMENTS

We thank IBAMA/ICMBio for funding ECOSAR surveys, Lauro S.P. Madureira (FURG) and their staff during surveys and all students who processed the biological samples. We express our gratitude to Maria Cristina Cergole, Suzana Anita Saccardo, Carmen Lúcia Del Bianco Rossi Wongtschowski, Antônio Olinto Ávila da Silva, Marcus Rodrigues da Costa and Teodoro Vaske Júnior. To CAPES for the scholarship granted to the first author. For referees and their valuable recommendations. The first author expresses his gratitude to CNPq due to the research grant 305403/2015-0.

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  • Zar JH (2010) Biostatistical analysis. Pearson, 5th ed., New Jersey, 944 pp.

  • Editorial responsibility:

    Cassiano Monteiro Neto

  • Zoobank:

    http://zoobank.org/87858997-8113-4717-A67A-673CD99E3D52

Publication Dates

  • Publication in this collection
    2017

History

  • Received
    16 Aug 2016
  • Reviewed
    16 Dec 2016
  • Accepted
    20 Mar 2017
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