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Conversion factor for size structure reconstruction and growth of Lophius gastrophysus Miranda Ribeiro, 1915 catches in the South Atlantic Ocean

Abstract

Lophius gastrophysus is an exploited deep-sea demersal fishing resource in the Southeast-South region of Brazil, a capture target for the twin trawl multi-specific bottom trawling fleet. Considered a prized fish, it is commonly processed on board, retaining commercial cuts of the tail called colas. Thus it is difficult to recognize the original length of the specimens and, consequently, to assess the dynamics of sources and the impacts of catches. Individual length and weight data were used to generate body predictive models correlated to the monkfish cola. The parameters of growth, mortality and exploitation rate were also estimated from the size structure of Lophius gastrophysus captured by the industrial bottom trawling fleet in Southeast Brazil between May 2013 and April 2015, assessing monkfish stock based on indirect estimates of life history information. All predictive models of body size showed coefficients of determination above 0.9 (P <0.001), indicating good data adjustments. The growth parameters were determined by the minimum residual variance methodology, being: L = 92.4 cm and k = 0.158/year. The size structure tended to increase with depth in the spring. Individuals under the age of sexual maturity contributed significantly to catches. The length of first sexual maturation was estimated at 48.6 cm and the recruitment age was between 6 and 7 years. The coefficients of total, natural, fishing and exploitation rates were estimated at: Z = 0.676; M = 0.241; F = 0.436 and E = 0.644. The fishing mortality rate was higher than the reference value (E > 0.50) indicating a state of overexploitation of the southeast most part of stock. A review of the current Brazilian legislation established for the species on the southeast coast is recommended.

Descriptors:
Monkfish; Age; Growth; Mortality; Fishing

INTRODUCTION

Baseline information on the size relationships involving length and weight variables are important predictive indices i. e., weight-length ratio, length-length, fertility-length, and fertility-weight (Silva et al., 2016SILVA, J. P., COSTA, M. R. D., GOMES, I. D. & ARAÚJO, F. G. 2016. Gonadal development and fecundity of the smooth weakfish Cynoscion leiarchus (Teleostei: Perciformes: Sciaenidae) in a tropical Brazilian bay. Zoologia, 33(6), 1-8.), with several applications in the science of fishing. Length measurements of a fish population are an important factor to investigate the ecological processes of a species, particularly due to the ease of obtaining this variable. From the size structure, it is possible to estimate the parameters of population growth and mortality and exploitation to better understand the dynamics of fish populations (Froese et al., 2018FROESE, R., WINKER, H., CORO, G., DEMIREL, N., TSIKLIRAS, A. C., DIMARCHOPOULOU, D., SCARCELLA, G., PROBST, W. N., DUREUIL, M. & PAULY, D. 2018. A new approach for estimating stock status from length frequency data. ICES Journal of Marine Science, 76(1), 350-351.).

The monkfish, Lophius gastrophysus is a marine demersal teleost, distributed along the continental shelf and slope of the Atlantic Ocean between the United States and Argentina (Valentim et al., 2007VALENTIM, M. F. M., VIANNA, M. & CARAMASCHI, E. P. 2007. Length structure of monkfish, Lophius gastrophysus (Lophiiformes, Lophiidae), landed in Rio de Janeiro. Brazilian Journal of Aquatic Science and Technology, 11(1), 31-36.). The species is found at depths down to 1000 meters and occurs with greater abundance in bottoms with unconsolidated substrates and cold waters (Vianna and Almeida, 2005VIANNA, M. & ALMEIDA, T. 2005. Bony fish bycatch in the southern Brazil pink shrimp (Farfantepenaeus brasiliensis and F. paulensis) fishery. Brazilian Archives of Biology and Technology, 48(4), 611-623.; Botelho et al., 2009BOTELHO, M. L. L. A., Á VILA-DA-SILVA, A. O. & BASTOS, G. C. C. 2009. A evolução da pesca paulista de Lophius gastrophysus (Miranda-Ribeiro, 1915) na margem continental da costa brasileira entre 22° e 28°S. Boletim do Instituto de Pesca, 35(3), 413-427.). In the Southeast-South regions of Brazil, it is a relatively common component of the deep demersal ichthyofauna (Brasil, 2006BRASIL. Ministério do Meio Ambiente. 2006. Relatório Executivo. Programa ReviZEE: avaliação do potencial sustentável de recursos vivos na Zona Econômica Exclusiva. Brasília: Ministério do Meio Ambiente.), being exploited throughout the year (Perez et al., 2002aPEREZ, J. A. A., PEZZUTO, P. R., ANDRADE, H. A., SCHWINGEL, P. R., RODRIGUES-RIBEIRO, M. & WAHRLICH, R. 2002. O ordenamento de uma nova pescaria direcionada ao peixe-sapo (Lophius gastrophysus) no Sudeste e Sul do Brasil. Notas Técnicas da FACIMAR, 6, 65-83.), with significant catches made by bottom trawl fleets. However, until 2002, it was also targeted for capture by deep gillnet fleets (Perez et al., 2002bPEREZ, J. A. A., WAHRLICH, R., PEZZUTO, P. R. & LOPES, F. R. A. 2002. Estrutura e dinâmica da pescaria do peixe-sapo Lophius gastrophysus no sudeste e sul do Brasil. Boletim do Instituto de Pesca, 28(2), 205-230.).

The monkfish fishery is relatively recent, compared to other relevant species, for Brazilian fisheries harvests (Perez et al., 2002aPEREZ, J. A. A., PEZZUTO, P. R., ANDRADE, H. A., SCHWINGEL, P. R., RODRIGUES-RIBEIRO, M. & WAHRLICH, R. 2002. O ordenamento de uma nova pescaria direcionada ao peixe-sapo (Lophius gastrophysus) no Sudeste e Sul do Brasil. Notas Técnicas da FACIMAR, 6, 65-83.). Until the mid-1990s, there was no demand for monkfish in the domestic market, and the fleet acted mostly upon traditional fishery resources distributed along the inner continental shelf (Botelho et al., 2009BOTELHO, M. L. L. A., Á VILA-DA-SILVA, A. O. & BASTOS, G. C. C. 2009. A evolução da pesca paulista de Lophius gastrophysus (Miranda-Ribeiro, 1915) na margem continental da costa brasileira entre 22° e 28°S. Boletim do Instituto de Pesca, 35(3), 413-427.). This scenario changed from the 2000s onwards due to the combination of two factors: the foreign vessel leasing program and the displacement of the national fleet towards the outer continental shelf and slope fishing grounds (Perez et al., 2002aPEREZ, J. A. A., PEZZUTO, P. R., ANDRADE, H. A., SCHWINGEL, P. R., RODRIGUES-RIBEIRO, M. & WAHRLICH, R. 2002. O ordenamento de uma nova pescaria direcionada ao peixe-sapo (Lophius gastrophysus) no Sudeste e Sul do Brasil. Notas Técnicas da FACIMAR, 6, 65-83.; Botelho et al., 2009BOTELHO, M. L. L. A., Á VILA-DA-SILVA, A. O. & BASTOS, G. C. C. 2009. A evolução da pesca paulista de Lophius gastrophysus (Miranda-Ribeiro, 1915) na margem continental da costa brasileira entre 22° e 28°S. Boletim do Instituto de Pesca, 35(3), 413-427.). Thus, the deep-sea fisheries of the monkfish emerged with expectations of high yields and productivity (Perez et al., 2002bPEREZ, J. A. A., WAHRLICH, R., PEZZUTO, P. R. & LOPES, F. R. A. 2002. Estrutura e dinâmica da pescaria do peixe-sapo Lophius gastrophysus no sudeste e sul do Brasil. Boletim do Instituto de Pesca, 28(2), 205-230.). As a result, the species experienced high exploitation rates without scientific projections of sustainable yields (Perez et al., 2002bPEREZ, J. A. A., WAHRLICH, R., PEZZUTO, P. R. & LOPES, F. R. A. 2002. Estrutura e dinâmica da pescaria do peixe-sapo Lophius gastrophysus no sudeste e sul do Brasil. Boletim do Instituto de Pesca, 28(2), 205-230.). Subsequently, studies conducted by the Department of Fisheries and Aquaculture (DPA) of the Ministry of Agriculture, Livestock and Supply (MAPA) and the REVIZEE Project (Recursos Vivos na Zona Econômica Exclusiva) indicated the vulnerability of monkfish stocks as a consequence of the strong fishing pressures (Brasil, 2006BRASIL. Ministério do Meio Ambiente. 2006. Relatório Executivo. Programa ReviZEE: avaliação do potencial sustentável de recursos vivos na Zona Econômica Exclusiva. Brasília: Ministério do Meio Ambiente.). In the mid-2000s, the species was included in the National List of Overexploited Fish Species (Botelho et al.,2009). Despite the species’ unfavourable scenario, few studies have been conducted to understand the population biology of L. gastrophysus.

Commercial landings by the fishing fleet are important sources of data for the development of population studies (Fonteles-Filho, 2011FONTELES-FILHO, A. A. 2011. Oceanografia, biologia e dinâmica populacional de recursos pesqueiros. Fortaleza: Expressão Gráfica e Editora.). Monkfish are often processed on board to obtain commercial cuts called “colas”, which consist of the tail portion of the fish and are the main processing products of the species (Perez et al, 2002aPEREZ, J. A. A., PEZZUTO, P. R., ANDRADE, H. A., SCHWINGEL, P. R., RODRIGUES-RIBEIRO, M. & WAHRLICH, R. 2002. O ordenamento de uma nova pescaria direcionada ao peixe-sapo (Lophius gastrophysus) no Sudeste e Sul do Brasil. Notas Técnicas da FACIMAR, 6, 65-83.). However, this procedure reduces the possibility of monitoring catches using length-based criteria (e.g., minimum and/or maximum catch sizes). Furthermore, it limits the estimation of relevant biological attributes for assessing stock status, which are usually based on length composition (Froese et al., 2018FROESE, R., WINKER, H., CORO, G., DEMIREL, N., TSIKLIRAS, A. C., DIMARCHOPOULOU, D., SCARCELLA, G., PROBST, W. N., DUREUIL, M. & PAULY, D. 2018. A new approach for estimating stock status from length frequency data. ICES Journal of Marine Science, 76(1), 350-351.).

The regulation of the monkfish fishery was instituted by Interministerial Normative Instruction MPA/MMA 3rd, of September 4, 2009 (Brazil, 2009), which defines fisheries management criteria and procedures. Among them, there is the need for a fishing permit issued by the government fisheries authority, definition of fishing exclusion areas, limitation of the fleet to nine Brazilian vessels, with a mandatory on-board observer, an annual catch quota of 1,500 tons, minimum depth of action of 250 meters and use of a fixed bottom stationary net only. Finally, the regulation states that renewal of the fishing permit is conditional on determination of technical feasibility given the state of exploitation of the stock (Brasil, 2009BRASIL. Ministério da Pesca e Aquicultura. 2009. Instrução Normativa Interministerial MPA/MMA nº 03, de 4 de setembro de 2009. Estabelece critérios e procedimentos para o ordenamento da pesca do peixe-sapo (Lophius gastrophysus), nas águas jurisdicionais brasileiras das regiões Sudeste e Sul. Diário Oficial da União, Brasília, DF, 09 set 2009, Seção 1, pp. 27.).

In this sense, studies on the population structure in terms of size, age and growth and mortality rates of species subject to exploitation are the technical framework that supports the management of fishing activities (Dias-Neto, 2010DIAS NETO, J. 2010. Gestão do uso dos recursos pesqueiros marinhos no Brasil. 2ª ed. Brasília: IBAMA (Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renovávaveis).; Fonteles-Filho, 2011FONTELES-FILHO, A. A. 2011. Oceanografia, biologia e dinâmica populacional de recursos pesqueiros. Fortaleza: Expressão Gráfica e Editora.). This set of information is important because it allows understanding the behavior of stocks against fishing pressure and the monitoring of attributes that influence the size of populations and, consequently, the abundance of resources (Dias-Neto, 2010DIAS NETO, J. 2010. Gestão do uso dos recursos pesqueiros marinhos no Brasil. 2ª ed. Brasília: IBAMA (Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renovávaveis).; Prince et al., 2020PRINCE, J., CREECH, S., MADDUPPA, H. H. & HORDYK, A. R. 2020. Length based assessment of spawning potential ratio in data-poor fisheries for blue swimming crab (Portunus spp.) in Sri Lanka and Indonesia: implications for sustainable management. Regional Studies in Marine Science, 36, 101309.). The present work aims to reconstruct the catch size structure of L. gastrophisus from colas landing data to determine its growth parameters, mortality rates and level of exploitation. Such information may help further understanding of species life history in Southwestern Atlantic-Brazil and contribute to decision-making in resource management.

METHODS

Sample collection and procedure

The study area comprises the continental shelf region between latitudes 21°56’S (Rio de Janeiro) and 24°45’S (São Paulo) (Figure 1), within the Southeastern-Eastern Brazil Marine Ecoregions (Spalding et al., 2007SPALDING, M. D., FOX, H. E., ALLEN, G. R., DAVIDSON, N., FERDANA, Z. A., FINLAYSON, M., HALPERN, B. S., JORGE, M. A., LOMBANA, A., LOURIE, S. A., MARTIN, K. D., MCMANUS, E., MOLNAR, J., RECCHIA, C. A. & ROBERTSON, J. 2007. Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. Bioscience, 57(7), 573-582.). The Brazil Current (BC) is formed by two surface waters and two water masses: (i) Coastal Water - CW; (ii) Tropical Water - TW. The South Atlantic Central Water (SACW) joins the BC at 20°S, between 400 and 500 m deep to the southwest-south. On the continental slope, between 1000 and 2000 meters deep, close to the parallel 25°S, flows the Intermediate Contour Current - ICC, with the Intermediary Antarctic Water mass - IAW; and further down, at approximately 3000 meters deep, the Deep Contour Current - DCC, carrying the North Atlantic Deep Water- NADW, also moving South. The most striking oceanographic feature in this region is the summer presence of the SACW on the inner continental shelf (around the municipality of Cabo Frio) separated by a strong thermocline from the surface layer of AT mixture. The inflection of the coastline associated with bathymetric changes (narrowing of the Cabo Frio platform towards the northeast) and the greater intensity of the northeast winds (NE) favors the coastal outcrop of cold waters (SACW) rich in nutrients, leading to increased local biological productivity (Albuquerque et al., 2012ALBUQUERQUE, A. L. S., BELÉM, A. L., PORTILHO-RAMOS, R. C., MENDOZA, U. & BARBOSA, C. F. 2012. Projeto ressurgência: processos geoquímicos e oceanográficos no limite entre as bacias de Campos e Santos. Boletim de Geociências Petrobras, 20(1-2), 193-210.). During the winter, SACW retracts and the water column on the inner and middle shelf acquires almost homogeneous characteristics, with the disappearance of the thermocline. The region is characterized by the predominance of sand, mud and clay bottoms (Albuquerque et al., 2012ALBUQUERQUE, A. L. S., BELÉM, A. L., PORTILHO-RAMOS, R. C., MENDOZA, U. & BARBOSA, C. F. 2012. Projeto ressurgência: processos geoquímicos e oceanográficos no limite entre as bacias de Campos e Santos. Boletim de Geociências Petrobras, 20(1-2), 193-210.).

Figure 1
Area of operation of the double-bottom trawling fleet based in Rio de Janeiro. Capture points (circles) and landing ports (squares).

The data used in the present study came from a sampling effort conducted at the landing docks of the municipalities of Niterói and São Gonçalo (Rio de Janerio State), monitoring the landings of the twin trawl bottom trawling fleet. In all, 34 vessels were monitored, with operating areas between 30 and 150m deep on the continental shelf of Southeast Brazil. The data matrix was composed of two sets of information: the first consisted of a biometric database with information on cola length and weight, designated as Cola length (CL) and Cola weight (CW), of 5,879 captured specimens between November/2013 and June/2015 at depths of 30 to 150 m. Cola length was measured from the insertion of the first and third post-cephalic spines of the dorsal fin to the end of the caudal fin (Figure 2). This on-board processing practice aims to maximize the yields, with cola being the marketable body portion of the monkfish, thus the main component landed in this fishery. The second dataset came from whole individuals of L. gastrophysus obtained monthly from landings, as part of a monitoring program by the state fisheries authority, FIPERJ (Fundação Instituto de Pesca do Rio de Janeiro) with the participation of boat skippers and fishermen. The catches took place from May/2013 to April/2015 at depths of 30 to 120 m. The specimens were collected randomly at the time of unloading in order to allow all strata of size to be represented. A total of 372 specimens were collected, from which measures of total length (TL) in cm and total weight (TW) in grams were taken. Subsequently, cola cuts were obtained from 292 whole individuals, which were measured (cm) and weighed (g) to obtain the cola length (CL) and cola weight (CW).

Figure 2
Cutting cola of Lophius gastrophysus resulting from processing ready to take measurements of length (cm) and weight (g).

Data analysis

Power regressions were constructed from CL and CW data obtained in laboratory from whole individuals, having TL and TW from the same individuals as dependent variables to generate reliable predictive equations. For this purpose, the following regressions were elaborated: TL = a x CLb, TL = a x CWb, TW = a x CLb, TW = a x CWb, where a = linear coefficient and b = slope. The fit of the data to the models was measured using the determination coefficient(R2). The equations that provided the best adjustments for the CL and CW data from the biometric database for conversion to TL and TW were chosen. From the converted values and data from whole individuals, the weight-length relationship was estimated according to the Huxley equation (1924HUXLEY, J. S. 1924. Constant differential growth-ratios and their significance. Nature, 114(2877), 895-896.): TW = a x TLb. where a = linear coeficient and b = slope coefficient (allometry cofficient). To verify whether the increments in weight are isometric (b = 3) or allometric (b <3 <b), the value of the slope was tested using the Student t-test (t=(b-3)SD), using a of 5% (Zar, 1999ZAR, J. H. 1999. Biostatistical analysis. 4th ed. New Jersey: Prentice Hall.).

From the TL values measured and estimated by the regression, the individuals were grouped into 1.0 cm length classes for analysis of the length frequency distributions. Possible differences in fish size depending on depth (class: 0-50 m, 51-100 m, 101-150 m) and austral seasons (summer: January-March; autumn: April-June; winter: July-September and spring: October-December) were investigated using the Kruskal-Wallis test. The biomass (kg) of the catch landed was estimated (regression equations) by depth class, season and the estimated age of the individuals (interval of 1 year). The growth parameters of von Bertalanffy (1938)VON BERTALANFFY, L. 1938. A quantitative theory of organic growth. Human Biology, 10(2), 181-213. were estimated using an iterative routine to minimize variances according to Gonçalves and Fontoura (1999)GONÇALVES, P. L. & FONTOURA, N. F. 1999. Dinâmica populacional de Palaemonetes argentinus na lagoa Fortaleza, Rio Grande do Sul, Brasil (Decapoda, Palaeminidae), Iheringia. Série Zoologia, 86, 171-186.. The starting point is to obtain the average lengths of the age groups/cohorts, obtained from the analysis of the frequency distribution of the length data, grouped monthly. To this end, the Battacharya’s method provided by the FISAT II FAO-ICLARM package was applied to identify the number of cohorts and estimate the respective average lengths. These average length values were used as seed input values to estimate growth parameters using the method of minimum residual variance (MINIVAR). This method allows quantification of the asymptotic length (L∞) and the instantaneous growth rate (k) with less variance from an estimated value of L∞ (Pauly empirical equation, 1983PAULY, D. 1983. Some simple methods for the assessment of tropical fish stocks. FAO Fisheries and Technical Papers, 234, 1-52.) and the assumed age of the first age group. To define the age of the first group, it was necessary to determine the interval of months between the reproductive/recruitment period and the appearance of the first age group in the samples and then divide this value by 12 (number of months in the annual cycle). From this data, the algorithm returns values of L∞ and k within biologically acceptable limits which are refined by the operator according to the biological characteristics of the species. Ages were assumed at 1 year intervals, taking into account the formation of an annual age ring observed in the species (Lopes, 2005LOPES, F. R. A. 2005. Reprodução, idade de crescimento do peixe-sapo (Lophius gastrophysus) (Ribeiro, 1915) na região sudeste e sul do Brasil [online]. MSc. Itajaí: UNIVALI (Universidade do Vale do Itajaí). Available at: http://www.dominiopublico.gov.br/pesquisa/DetalheObraForm.do?select_action=&co_obra=11484 [Accessed: 15 Jul 2020].
http://www.dominiopublico.gov.br/pesquis...
). The selection of the best set of values is made based on the lowest residual variance (RV) obtained. As a reference, RV values below 0.5% were considered acceptable. The k parameter was estimated by the equation k=[-ln{1-(LtL)}÷t]÷n, where Lt: average length (cm) of the age group with assumed age t (years); L∞: asymptotic length (cm); and n: number of age-length pairs. The residual variance was given by the equation: S2=[{L×(1-ek×t)-Lt}2÷(n-1)]. To verify the fit between parameters k and L∞ as well as consistency with the other estimates made for similar species, the growth performance index (φ) was estimated according to Pauly and Munro (1984)PAULY, D. & MUNRO, J. L. 1984. Once more on the comparison of growth in fish and invertebrates. ICLARM: International Center for Living Aquatic Resources Management, Fishbyte, 2(1), 1-21.: =logk+2×logL. Longevity (A0.95), defined as the time required to reach 95% of the asymptotic length, was estimated by the Taylor equation (1960TAYLOR, C. C. 1960. Temperature, growth, and mortality - the pacific cockle. Journal du Conseil, 26(1), 117-124.): A0.95=t0+(2.996÷k). To calculate the theoretical age at zero length (t0), a parameter of the von Bertalanffy growth equation, the Pauly formula (1979PAULY, D. 1979. Theory and management of tropical multispecies stocks-A review, with emphasis on the Southeast Asia demersal fisheries. ICLARM: International Center for Living Aquatic Resources Management, Studies and Reviews, 1, 1-35.) was used: log-t0=-0.392-0.275×logL-1.038×logk.

The total mortality coeficient (Z) was estimated through the linearized length capture curve according to the model presented by Sparre and Venema (1992)SPARRE, P. & VENEMA, S. C. 1992. Introduction to tropical fish stock assessment. Rome: FAO.. The natural mortality coeficient (M) was obtained by the empirical equation of Rikhter and Efanov (1976)RIKHTER, V. A. & EFANOV, V. N. 1976. On one of the approaches to estimation of natural mortality of fish populations. ICNAF: International Commission for the Northwest Atlantic Fisheries, 377, 1-12.: M=1.521÷(Tm50%×0.720)-0.155, where: Tm50% = age of first sexual maturation (corresponding to the length in which 50% of individuals reached sexual maturity). The Tm50% value was obtained using the von Bertalanffy (1938)VON BERTALANFFY, L. 1938. A quantitative theory of organic growth. Human Biology, 10(2), 181-213. (I) inverse equation using the 48.6 cm length of first sexual maturation (L50%) estimated by the empirical equation of Froese and Binohlan (2000)FROESE, 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(4), 758-73. (II): (I) tm50%=t0-(1k)×ln[1-(LL)]; (II) logL50%=0.8979×logL-0.0782.

The fishing mortality coeficient (F) was determined by the difference between Z and M (I) whereas the exploitation rate (E) was estimated by the quotient between F and Z (II): (I) F = Z - M; (II) E=FZ.

RESULTS

The power regressions to estimate TL and TW from the CL and CW data showed good adjustments for the conversion of biometric data (Table 1). Among all regressions, the following equations were selected: TL = 2.6282 x CL0.8488, n = 254, R2 = 0.964 and TW = 5.6032 x CW0.9254, n = 260, R2 = 0.956 (Fig. 3A and B).

Figure 3
Power regressions between the cola length and the total length (A) and between the cola weight and the total weight (B).

Table 1
Power regressions to estimate the total weight (TW) and total length (TL) from the cola weight (CW) and the cola length (CL). n = sample size, R2 = coefficient of determination.

The 372 whole individuals presented TL between 22.0 and 75.8 cm, mean ± standard error of 46.8 ± 0.49 cm and mode of 45.0 cm. The samples from the cola biometrics totaled 5,879 specimens, converted based on the regression TL = 2.6282 x CL0.8488, which generated a distribution of total lengths that ranged from 18.0 to 84.6 cm, mean 49.8 ± 0, 21 cm and modes between 47.0 and 49.0 cm. The frequency distribution by TL class (Fig. 4) for all individuals (i. e. whole and cola) indicated a predominance of specimens with total length between 45.0 and 51.0 cm and modes between 47.0 and 49.0 cm. Individuals smaller than 25.0 cm and larger than 70.0 cm were not abundant. The size structure showed significant variation between the seasons (Kruskal-Wallis, H = 885, p <0.05) with higher TL values in the spring months, both in 2013 and 2014. Significant differences were also found between the depth classes (Kruskal-Wallis, H = 50.27, p <0.05) with a clear tendency to increase the TL according to the depth increase, notably in the 101-150m depth range.

Figure 4
A = Total length frequency distributions of L. gastrophysus obtained directly from the sampling. B = Total length frequency distributions of L. gastrophysus resulting from the application of the conversion factors to the “colas" size structure.

The weight-length relationship for grouped sexes was determined from weight and length data of 5,905 individuals, from whole specimens and converted “colas”, being estimated at: TW = 0.0136 x TL3.0156, n = 5,905, R2 = 0.953. The Student t-test did not indicate a significant difference for the coefficient b (t = - 0.21825, p> 0.05) characterizing growth of the isometric type (b = 3) (Fig 5).

Figure 5
Weight/Length relationship of the monkfish Lophius gastrophysus in southeastern Brazil. Dark circles (estimated data) and light circles (biometric data).

The total monitored biomass landed was 12,379 kg. In 2013, 2,207 kg were landed during six months; in 2014, a total of 7,331 kg were landed in 11 months; and in 2015, 2,841 kg were landed in six months of sampling in the monitored ports. The estimated biomass values varied by depth classes, with more expressive values in the intermediate classes of 51-100 m and 101-150 m, with yields of 7,541 and 3,909 kg respectively, followed by the other classes of 0-50 and 151-200m with respectively 706 and 223 kg. Seasonally, there was an interannual pattern of higher yields in the spring/summer period followed by a decrease in autumn/winter (Table 2).

Table 2
Estimates of biomass (kg) by season and depth class (m).

From the length frequency distribution data, it was possible to estimate the growth parameters and identify the number of age groups (cohorts) present in the global sample. Modal decomposition using Battacharya’s method identified five cohorts and their respective average lengths. Biological data of the species available in the literature were used to refine the results. The routine of minimum residual variance (VR) adjusted to von Bertalanffy’s growth function returned 10 pairs of values (L∞ and k), with asymptotic lengths ranging from 92.0 to 92.9 cm and the instantaneous growth rate from 0.1587 to 0.1561 year-1. The pair that provided the lowest RV value (0.0623) was L∞ = 92.4 and k = 0.1576 year-1 (Fig. 6).

Figure 6
Minivar plots to estimate growth parameters of the von Bertalanffy growth function for the monkfish Lophius gastrophysus in southeastern Brazil.

Longevity was estimated at 17.20 years, the theoretical age t0 at 1.765 and the growth performance index (φ) at 3.13. Table 3 summarizes the values of the growth parameters and performance indexes obtained in this work and those recorded in the literature for similar species.

Table 3
Compilation of estimated growth parameter values for Lophius genus. Asymptotic length (L∞), growth constant (k), longevity, growth performance index (φ), sex and source of information. Data compiled at literature using only species with growth data available.

The estimated von Bertalanffy growth equation was: L=92.4×[1-e-0.158(t-1.765)] (Fig. 7A). From the growth parameters, the capture curve converted into linearized length was constructed using the entire structure in sample size considering a theoretical year (May to April) (Fig. 7B). The age of recruitment to fishing was approximately 6 years, which corresponded to 31% of the catches in the study area (Fig. 7B); catches under the age of 6 years totaled 29% whereas catches over 6 years totaled 40%. In terms of biomass, production at the age of 6 was 3,185.6 kg, equivalent to 25.7% of the total. The estimated biomass for ages under 6 years was 1,523 kg (12.3% of the total), while over 6 years was 7,670.7 kg (62.0%). Thus, specimens under the age of sexual maturity (i50 = 6 years) contributed considerably to the catches in numbers, however in biomass they contributed less proportionally.

Figure 7
A - Estimated von Bertalanffy growth curves with minimal residual variance for the monkfish L. gastrophysus in southeastern Brazil. B - Length converted catch curves applied to length frequencies data. The slope of the right descending arm (black dots) of the curve allows the estimation of total mortality (Z; R2 = 0,974). White dots (empirical data) the numbers of fish actually sampled (observed).

The instantaneous total mortality rate (Z) for the period 2013-2015 was estimated at 0.676 year-1. The natural mortality rate (M) estimated by the method of Rikhter and Efanov (1976)RIKHTER, V. A. & EFANOV, V. N. 1976. On one of the approaches to estimation of natural mortality of fish populations. ICNAF: International Commission for the Northwest Atlantic Fisheries, 377, 1-12. was 0.241 year-1. Fishing mortality (F) was 0.436 and the exploitation rate (E) equal to 0.644. Fishing mortality (F) was relatively higher than the natural mortality rate being the main vector of population mortality. Furthermore, the value of the exploitation rate suggests a state of overexploration of the stock.

DISCUSSION

In recent decades, fisheries scientists have used population size structure as a qualitative indicator of a species’ development, which can be understood as the relationships between body measurements (total weight - TW vs. total length - TL) or between age and length (Schwamborn et al., 2019SCHWAMBORN, P. R., MILDENBERGER, T. K. & TAYLOR, M. 2019. Assessing sources of uncertainty in length-based estimates of body growth in populations of fishes and macroinvertebrates with bootstrapped ELEFAN. Ecological Model, 393(C), 37-51.). Fish growth tends to be continuous throughout life, even after reaching sexual maturity. Thus, it is expected that random sampling of fish reflects the variations in size distribution of fish in the total stock, or part of the stock available in the catch area (King, 2007KING, M. 2007. Fisheries biology, assessment and management. 2nd ed. Oxford: Blackwell Science.). Based on this information, it is possible to quantify the impacts of fishing on fish stocks, ensuring the safe exploitation of resources. Obtaining biological information about stocks is commonly challenging and ends up making stock monitoring dificult. However, size structure data are relatively easy to collect and serve as a starting point for population investigations. Thus, our results are in line with the principles of fisheries management by using simple and easily visualized data (size and weight of fish) to build a predictive model for monitoring the fishery.

In the Southwestern Atlantic, the main forms of commercialization of landed monkfish are either gutted individuals or only the cola. In cases like this, it is common to use conversion equations for biometric measurements to estimate the actual size and weight of the fish caught. This approach is common for the genus Lophius, as can be seen in Almeida et al. (1995)ALMEIDA, F. P., HARTLEY, D. L. & BURNETT, J. 1995. Length-weight relationships and sexual maturity of goosefish off the northeast coast of the United States. North American Journal of Fisheries Management, 15, 14-25.. For L. gastrophysus, these conversions were applied by Perez et al. (2002a)PEREZ, J. A. A., PEZZUTO, P. R., ANDRADE, H. A., SCHWINGEL, P. R., RODRIGUES-RIBEIRO, M. & WAHRLICH, R. 2002. O ordenamento de uma nova pescaria direcionada ao peixe-sapo (Lophius gastrophysus) no Sudeste e Sul do Brasil. Notas Técnicas da FACIMAR, 6, 65-83. to convert cola weight into total weight and subsequently derive estimates of yield and production. In this work, we used the size structure to estimate the parameters of growth and mortality. Our estimates showed good adjustments demonstrating the effectiveness of these predictive equations, in agreement with the findings of Perez et al. (2002a)PEREZ, J. A. A., PEZZUTO, P. R., ANDRADE, H. A., SCHWINGEL, P. R., RODRIGUES-RIBEIRO, M. & WAHRLICH, R. 2002. O ordenamento de uma nova pescaria direcionada ao peixe-sapo (Lophius gastrophysus) no Sudeste e Sul do Brasil. Notas Técnicas da FACIMAR, 6, 65-83.. Therefore, the estimates of the structure in size (cola) from the landings of the fishing fleet could be used in determining parameters of the life history of this resource.

The estimated weight-length ratio showed values of the coefficients a and b similar to those reported in the literature for the species and its cogenerics (Froese and Pauly, 2020FROESE, R. & PAULY, D. 2020. FishBase [version 10/2016] [online]. Paris: Lancaster University. Available at: www.fishbase.org [Accessed: 17 Jul 2020].
www.fishbase.org...
), showing isometric growth. The differences observed in this and other studies can be attributed to a combination of factors such as: number of specimens evaluated, size structure, sex, gonadal maturity in addition to the preservation techniques used (Hossain et al, 2014HOSSAIN, M. Y., RAHMAN, M. M., AHMED, F., AHMED, Z. F. & OHTOMI, J. 2014. Length-weight and length-length relationships and form factor of three threatened fishes from the Ganges River (NW Bangladesh). Journal of Applied Ichthyology, 30(1), 221-224.). Morphometric relationships, mainly of fishery resources, are necessary to estimate unknown measures from known measures. In addition to these, it is also possible to estimate the condition of individuals from the linear coefficient related to the body shape (Froese, 2006FROESE, R. 2006. Cube law, condition factor and weight-length relationships: history, meta-analysis and recommendations. Journal of Applied Ichthyology, 22(4), 241-253.). In fish population dynamics, the most commonly used measures are length and weight. In this way, the parameters estimated here should be applied only within the specified length ranges, allowing good estimates of both total length and total individual weight/biomass landed from the monkfish colas.

Population parameters of the monkfish, including growth parameters, estimated from the reconstructed size structure, were similar to those presented by Lopes (2005)LOPES, F. R. A. 2005. Reprodução, idade de crescimento do peixe-sapo (Lophius gastrophysus) (Ribeiro, 1915) na região sudeste e sul do Brasil [online]. MSc. Itajaí: UNIVALI (Universidade do Vale do Itajaí). Available at: http://www.dominiopublico.gov.br/pesquisa/DetalheObraForm.do?select_action=&co_obra=11484 [Accessed: 15 Jul 2020].
http://www.dominiopublico.gov.br/pesquis...
and Costa et al. (2019)COSTA, M. R., TUBINO, R. A., ANDRADE-TUBINO, M. F., MARTINS, R. R. M., VIEIRA, F. & MONTEIRO-NETO, C. 2019. Lophius gastrophysus Miranda Ribeiro 1915 at the coast of Rio de Janeiro state. In: VAZ-DOS-SANTOS, A. M. & ROSSI-WONGTSCHOWSKI, C. L. D. B. (eds.). Growth in fisheries resources from the Southwestern Atlantic. São Paulo: Instituto Oceanográfico - USP (Universidade de São Paulo), pp. 210-213. for the Southwestern Atlantic . Our results were also consistent with growth parameters from cogeneric species with similar theoretical maximum lengths, available on the Fishbase platform (Froese and Pauly, 2020FROESE, R. & PAULY, D. 2020. FishBase [version 10/2016] [online]. Paris: Lancaster University. Available at: www.fishbase.org [Accessed: 17 Jul 2020].
www.fishbase.org...
), as well as their performance index values (Tab. 3). Some differences observed may be associated with the inverse relationship between k and L∞, i.e. if a fish grows rapidly, it will take less time to reach its maximum length, which will have a lower value than if it had grown slowly and reached a larger size (Fonteles-Filho, 2011FONTELES-FILHO, A. A. 2011. Oceanografia, biologia e dinâmica populacional de recursos pesqueiros. Fortaleza: Expressão Gráfica e Editora.). However, we cannot rule out other factors acting on growth, which were not measured here. The selectivity of the fishing gear and the different catch/fishing areas can determine variations in the population structure in terms of size and other vital processes of the species, regardless of the growth rate, which is genetically/physiologically determined for each species (Beverton and Holt, 1957BEVERTON, R. J. H. & HOLT, S. J. 1957. On the dynamics of exploited fish populations. Great Britain Fisheris Investment Service, 19(2), 1-533.; King, 2007KING, M. 2007. Fisheries biology, assessment and management. 2nd ed. Oxford: Blackwell Science.). Other estimates of the population dynamics of the species have also been determined and are closely linked to growth. They are the rates of mortality, longevity and exploitation. The estimated growth coefficient has a physiological interpretation associated with the availability of food and the space occupied by the species, suggesting that it has to adapt to different realities, with the consequence of specific population parameters for each population or strata (Lowe-McConnell, 1999LOWE-MCCONNELL, R. C. 1999. Estudos ecológicos de comunidades de peixes tropicais. São Paulo: EDUSP.). Our indirect estimates pointed to a relative age of 8.8 years, close to the findings of Camilo and Schwingel (2019)CAMILO, H. S. C. & SCHWINGEL, P. R. 2019. Lophius gastrophysus Miranda Ribeiro 1915 at the Brazilian Southwestern Atlantic. In: VAZ-DOS-SANTOS, A. M. & ROSSI-WONGTSCHOWSKI, C. L. D. B. (eds.). Growth in fisheries resources from the Southwestern Atlantic. São Paulo: Instituto Oceanográfico - USP (Universidade de São Paulo), pp. 206-209. who used the illicium to determine age, with the highest frequency between 6 and 11 years (30-75 cm TL). The study by Sherwood et al. (2013)SHERWOOD, G. D., GRABOWSKI, J. H. & BROWN, C. 2013. Tagging to assess monkfish movement: additional tagging to assess monkfish (Lophius americanus) movements and stock structure in the northeastern United States. Portland: Gulf of Maine Research Institute. with the cogeneric species L. americanus, using the illicium, also observed ages between 8 and 10 years, indicating that this structure is an age predictor for this species. Several authors agree that lofiiform fishes are sit-and-wait predators, attracting their prey by moving the illicium (Valentim et. al, 2007VALENTIM, M. F. M., VIANNA, M. & CARAMASCHI, E. P. 2007. Length structure of monkfish, Lophius gastrophysus (Lophiiformes, Lophiidae), landed in Rio de Janeiro. Brazilian Journal of Aquatic Science and Technology, 11(1), 31-36.). Even though a similar age pattern is observed at different latitudes, such feeding behavior (variable between sexes, size and type of habitat) (Sherwood and Grabowski, 2006SHERWOOD, G. D. & GRABOWSKI, J. 2006. Critical diet and habitat shifts in New England monkfish (Lophius americanus): insights from stable isotopes and relation to biological condition. Portland: Gulf of Maine Research Institute.) could explain the variable nature of the presence of age rings in different locations.

The obtained longevity of 17.2 years is directly associated with the high value of t0 and k, indicating that it is within the expected for the species, since Camilo and Schwingel (2019)CAMILO, H. S. C. & SCHWINGEL, P. R. 2019. Lophius gastrophysus Miranda Ribeiro 1915 at the Brazilian Southwestern Atlantic. In: VAZ-DOS-SANTOS, A. M. & ROSSI-WONGTSCHOWSKI, C. L. D. B. (eds.). Growth in fisheries resources from the Southwestern Atlantic. São Paulo: Instituto Oceanográfico - USP (Universidade de São Paulo), pp. 206-209. registered monkfish individuals up to 18 years of age. Other authors assessing the growth of cogeneric species through vertebrae (Armstrong et al., 1992ARMSTRONG, M. P., MUSICK, J. A. & COLVOCORESSES, J. A. 1992. Age, growth, and reproduction of the goosefish Lophius americanus (Pisces: Lophiiformes). Fishery Bulletin, 90(2), 217-230.), illicium (Sherwood et al., 2013SHERWOOD, G. D., GRABOWSKI, J. H. & BROWN, C. 2013. Tagging to assess monkfish movement: additional tagging to assess monkfish (Lophius americanus) movements and stock structure in the northeastern United States. Portland: Gulf of Maine Research Institute.) and otoliths Sun et al., (2020)SUN, Y., ZHANG, C., TIAN, Y. & WATANABE, Y. 2020. Age, growth, and mortality rate of the yellow goosefish Lophius litulon (Jordan, 1902) in the Yellow Sea. Journal of Oceanology and Limnology, 39(1-3), 1-9. also observed longevities equivalent to our findings. Comparing the estimated values of mortality M and F, we can see that fishing was the most important factor (F>M) for the southeast stock. Although a fishery targeting monkfish is relatively recent, it is known that one species is the target of accidental capture in shrimp fisheries (Vianna and Almeida, 2005VIANNA, M. & ALMEIDA, T. 2005. Bony fish bycatch in the southern Brazil pink shrimp (Farfantepenaeus brasiliensis and F. paulensis) fishery. Brazilian Archives of Biology and Technology, 48(4), 611-623.). Thus, it is possible that the high value of F is due not only to the targeted fishing but also to accidental captures. The estimated exploitation rate compared to the reference value equal to 0.5 (Gulland, 1983GULLAND, J. A. 1983. Fish stock assessment. A manual of basic method. Rome: FAO/Wiley Series on Food and Agriculture.) indicates that this stock is under pressure for fishing above the sustainable limit for the region, corroborating the state of overexploitation observed in past studies (Perez, 2006PEREZ, J. A. A. 2006. Potenciais de rendimento dos alvos da pesca de arrasto de talude do sudeste e sul do Brasil estimados a partir de parâmetros do ciclo de vida. Brazilian Journal of Aquatic Science and Technology, 10(2), 1-11.). Thangstad et al. (2006)THANGSTAD, T., BJELLAND, O., NEDERAAS, K. H., JÓNSSON, E., LAURENSON, C. H. & OFSTAD, L. H. 2006. Anglerfish (Lophius spp.) in Nordic waters. TemaNord, 2006, 570. conducted studies with L. budegassa and L. piscatorius stocks with a similar size structure to that reported in the present study and obtained the following mortality estimates: M = 0.15, F varying from 0.42 - 0.76 and Z varying from 0.57 - 0.91 for age groups between 4 and 11 years old. Comparing the estimated values for M and F in both regions, we can see that the fishing gear is decisive for estimating the exploitation rate by placing stocks close to or above the sustainable limits of exploitation. In this sense, we agree with Thangstad et al. (2006)THANGSTAD, T., BJELLAND, O., NEDERAAS, K. H., JÓNSSON, E., LAURENSON, C. H. & OFSTAD, L. H. 2006. Anglerfish (Lophius spp.) in Nordic waters. TemaNord, 2006, 570. who reported the need for more research to improve and standardize such estimates to obtain more accurate data on the total permitted catch size.

Tradicional trawler fishing grounds on the Rio de Janeiro coast are concentrated in the shallower area of total distribuition of monkfish (30-150m) during all seasons, especially due to the limited tecnology of the fishing vessels (Martins, 2017MARTINS, R. R. M. 2017. Dinâmica da pesca de arrasto de fundo no estado do Rio de Janeiro. DSc. Niterói: UFF (Universidade Federal Fluminense). Available at: https://app.uff.br/riuff/bitstream/handle/1/24864/Tese-Dina%cc%82mica%20da%20pesca%20de%20arrasto%20no%20RJ-2017.pdf?sequence=1&isAllowed=y [Accessed: 10 Jul 2020]
https://app.uff.br/riuff/bitstream/handl...
). The monkfish catches are higher in spring and summer, due to the reproductive cicle of the specie and SACW sincronization which promotes increased local biological productivity and, consequently, greater availability of food. In addition, it is recognized that the monkfish responds quickly to changes in ambient temperature, following the SACW intrusion on the continental shelf (Soares-Gomes and Fernandes, 2005SOARES-GOMES, A. & FERNANDES, F. C. 2005. Spatial distribution of bivalve mollusc assemblages in the upwelling ecosystem of the continental shelf of Cabo Frio, Rio de Janeiro, Brazil. Revista Brasileira de Zoologia, 22(1), 73-80.). This process increases the occurrence of juveniles in the region where the fishing fleet operates, as described by Valentim et al. (2007)VALENTIM, M. F. M., VIANNA, M. & CARAMASCHI, E. P. 2007. Length structure of monkfish, Lophius gastrophysus (Lophiiformes, Lophiidae), landed in Rio de Janeiro. Brazilian Journal of Aquatic Science and Technology, 11(1), 31-36., and observed in the present study 10 years later. Catches of fishes under the age of 6 years, the estimated age of monkfish recruitment, totaled 29.0% of total removed biomass, while the size structure showed that 45.9% of landings in RJ correspond to juvenile fish, in number of individuals, (TL < L50% = 48.6 cm). The high capture of juvenile individuals by the twin trawl bottom trawling fleet has already been documented and appears to be the result of the concentration of the fleet’s capture power in specimens smaller than 50.0 cm (Perez et al., 2002bPEREZ, J. A. A., WAHRLICH, R., PEZZUTO, P. R. & LOPES, F. R. A. 2002. Estrutura e dinâmica da pescaria do peixe-sapo Lophius gastrophysus no sudeste e sul do Brasil. Boletim do Instituto de Pesca, 28(2), 205-230.). This pattern was observed initially by Schwingel and Andrade (2002)SCHWINGEL, P. R. & ANDRADE, H. A. 2002. Capítulo 3: Aspectos biológicos e populacionais. In: PEREZ, J. A. A., ANDRADE, H. A., PEZZUTO, P. R., RODRIGUES-RIBEIRO, M., SCHWINGEL, P. R. & WAHRLICH, R. (eds.). Análise da pescaria do peixe-sapo no Sudeste e Sul do Brasil - Ano 2001. Itajaí: Convênio Ministério da Agricultura, Pecuária e Abastecimento - UNIVALI (Universidade do Vale do Itajaí). and later by Perez et al. (2002b)PEREZ, J. A. A., WAHRLICH, R., PEZZUTO, P. R. & LOPES, F. R. A. 2002. Estrutura e dinâmica da pescaria do peixe-sapo Lophius gastrophysus no sudeste e sul do Brasil. Boletim do Instituto de Pesca, 28(2), 205-230. on the Southeast-South coast of Brazil. On the coast of Rio de Janeiro, similar observations were made by Valentim et al., (2007)VALENTIM, M. F. M., VIANNA, M. & CARAMASCHI, E. P. 2007. Length structure of monkfish, Lophius gastrophysus (Lophiiformes, Lophiidae), landed in Rio de Janeiro. Brazilian Journal of Aquatic Science and Technology, 11(1), 31-36. and confirmed in this work. This pattern suggests a future loss of revenue and ecossytem health, due to the capture of a large number of low-value individuals which did not contribute to stock recomposition yet, decreasing the sustainability of these fishing practice and the monkfish as a fishery resource.

CONCLUSION

From the results raised here, we can conclude that: i) the conversion equations for body measurements are good predictors for the reconstruction of the species’ life history parameters and ii) the trawling fleet acting on the monkfish stock, even if considering it an accessory species, needs accurate management measures, which involve status and integrity monitoring of this stock since some parameters of its population dynamics already indicate a state of overexploitation of the catches made off the southeast coast. We recommend the development of an adequate management strategy for this fishery in view of minimizing the high catches of juveniles landed, maximizing the benefits of this fishery for food supply security.

ACKNOWLEDGMENTS

This work was supported by the FAPERJ (Grant # E-26/112.613/2012) and CNPq (Grant # 406249/2012-1). A. T. da Silva holds a Master’s degree scholarship from CAPES, C. Monteiro-Neto a Research Productivity Fellowship from CNPq (Grant # 305292/2016-1), and R. A. Tubino a post-doctoral fellowship PNPD-CAPES. We thank the trainees and staff from ECOPESCA-UFF and FIPERJ.

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Edited by

Associate Editor: Francesc Maynou

Publication Dates

  • Publication in this collection
    16 May 2022
  • Date of issue
    2022

History

  • Received
    27 July 2021
  • Accepted
    07 Jan 2022
Instituto Oceanográfico da Universidade de São Paulo Praça do Oceanográfico 191, CEP: 05508-120, São Paulo, SP - Brasil, Tel.: (11) 3091-6501 - São Paulo - SP - Brazil
E-mail: diretoria.io@usp.br