Biometric relationships and sex ratio for red-spotted shrimp Farfantepenaeus brasiliensis (Latreille, 1817) (Decapoda, Penaeidae) from the coast of Sergipe, northeastern Brazil

Abstract

The goal of this study was to estimate the biometric relationships and sex ratio for the red-spotted shrimp Farfantepenaeus brasiliensis (Latreille, 1817) from the coast of Sergipe, northeastern Brazil. A total of 132 specimens of F. brasiliensis were collected and analyzed (65 females and 67 males) from May 2015 to May 2016. The overall sex ratio did not differ significantly from 1:1. Female size ranged from 20.73-46.43 mm of carapace length (CL), whereas male size ranged from 20.75-32.47 mm CL. Females were larger (34.78±6.05 mm CL, 152.61±19.68 mm of total length (TL)) and heavier (27.45±11.12 g of wet weight (WW)) than males (27.07±2.56 mm CL, 123.77±11.37 mm TL and 14.48±3.98 g WW). All biometric relationships were different between females and males. Both TL vs CL and WW vs TL relationships indicated positive allometric growth. Data shown here correspond to an adult population exploited by local shrimp fleets and are an important contribution to the biological knowledge of this species, especially in northeastern Brazil, where there is no information available.

Keywords
Adult; allometry; length; pink shrimp; weight

INTRODUCTION

Marine shrimps of the genus Farfantepenaeus Burukovsky, 1997 are one of the most exploited fishing resources along the Brazilian coast due to their high market value (Dias-Neto, 2011; Boos et al., 2016). Despite this commercial importance, all species belonging to this genus are commonly known as “camarões-rosa” (= pink shrimps) along the Brazilian coast (Paiva et al., 2001; Dias-Neto, 2011). However, the exploitation rate of each species is variable along the coast due to their different latitudinal distributions. For example, Farfantepenaeus subtilis (Pérez-Farfante, 1967) can be found from the Caribbean Sea (20°N) to Cabo Frio (Rio de Janeiro, southeastern Brazil, 23°S), whereas Farfantepenaeus paulensis (Pérez-Farfante, 1967) occurs from Ilhéus (Bahia, northeastern Brazil, 14°S) to Mar del Plata (Argentina, 38°S) (Pérez-Farfante, 1988; Costa et al., 2003). On the other hand, Farfantepenaeus brasiliensis (Latreille, 1817) shows a wider distribution than the two previously mentioned species, occurring from North Carolina (USA, 35°N) to Rio Grande do Sul (southern Brazil, 33°S) (Pérez-Farfante, 1988; Costa et al., 2003).

The total catch of Farfantepenaeus spp. in Brazil was about 10.3 thousand tonnes in 2011, the last year for which catch statistics were available at a national level (Brasil, 2011). This amount was surpassed only by the Atlantic seabob shrimp, Xiphopenaeus kroyeri (Heller, 1862), with total catches of 15.4 thousand tonnes (Brasil, 2011). The total catch of Farfantepenaeus spp. on the coast of Sergipe (11°S) was about 176.6 tonnes in 2014 (Araújo et al., 2016). However, it should be noted that in both national and local reports, there was no discrimination of catch by species, making it difficult to assess the exploitation status of each stock (Dias-Neto, 2011; Valentini et al., 2012; Freire et al., 2015).

Farfantepenaeus brasiliensis can be found from shallow water to 366 m of depth, inhabiting detritus-rich muddy to gravel-sandy bottoms, and reach a size at first maturity at about 150 mm of total length (Boos et al., 2016). This species has a type II life cycle, in which reproduction and egg release take place offshore and the post-larvae migrate to estuarine areas where they grow up to juvenile/subadult stages and then move back offshore to complete their life cycles (Dall et al., 1990). Therefore, F. brasiliensis fishery may target two population groups: juveniles/subadults in estuaries and inshore habitats and/or adults offshore (D’Incao et al., 2002).

Several population studies have estimated biometric relationships for penaeids and these relationships have been highlighted as an important tool for assessing the status of shrimp stocks, as very often different morphological traits may reflect environmental influences or overfishing impacts (Leite-Jr and Petrere-Jr, 2006; Dumond and D’Incao, 2010; Segura and Delgado, 2012; Silva et al., 2015, 2018; Reis-Jr et al., 2019). Biometric relationships for F. brasiliensis have already been estimated for southern and southeastern Brazil (Mello, 1973; Villela et al., 1997; Branco and Verani, 1998; Albertoni et al., 2003; Leite-Jr and Petrere-Jr, 2006; Carvalho et al., 2019) as well as for the Gulf of Mexico (Arreguín-Sánchez, 1981; Pérez-Castañeda and Defeo, 2002; May-Kú et al., 2006). Nevertheless, there is no information about this species for northeastern Brazil. Here, F. brasiliensis from the coast of the Sergipe was studied to understand the biometric relationships and sex ratio of this species. Additionally, data from the present study were compared with those obtained from other populations to assess the variation of the traits studied along its distribution range.

MATERIAL AND METHODS

Samples of 3 kg each were collected monthly from each category separated manually by fishermen onboard, based on shrimp size (‘espigão’ - small size, ‘escolha’ - medium size, and ‘pistola’ - large size), before landing in the Port of Aracaju, state of Sergipe, from May 2015 to May 2016. No sample was collected during the closed seasons, which correspond to April 1st to May 15th and December 1st to January 15th (Brasil, 2004a). Artisanal shrimp trawlers along the coast of Sergipe are 8-13 m long and operate with double nets with a mesh size of 21 mm in the cod end (Fig. 1). It was not possible to collect information on local depth for each sample collected, as they were obtained from fishing boats at the port, right before landing. Each sample was taken to the Laboratório de Ecologia Pesqueira (LEP) of the Departamento de Engenharia de Pesca e Aquicultura (Depaq) at the Universidade Federal de Sergipe (UFS) and kept frozen until processing. Within each category, each specimen was identified according to Costa et al. (2003) and Teodoro et al. (2016), and all F. brasiliensis present in all samples were separated.

Figure 1.
Map showing the approximate location of the Port of Aracaju and the shrimp fishing ground on the coast of Sergipe.

Each specimen of F. brasiliensis had its total length (distance from the tip of the rostrum to the tip of the telson, TL, mm), rostrum length (distance from the tip of the rostrum to the postorbital margin, RL, mm), and carapace length (distance from the postorbital margin to the mid-dorsal posterior edge of the carapace, CL, mm) measured with a digital caliper (precision: 0.01 mm) and was weighed (wet weight; WW, g) with a scale (precision of 0.0001 g). All individuals were sexed by checking the thelycum in females and the petasma in males. The maturation stage of the gonads was classified according to Peixoto et al. (2003). Females were defined as immature (I), developing (II), mature (III) and spent (IV), and males as immature (non-linked petasma) and mature (linked petasma).

A chi-square test (χ²) with the Yates correction (Zar, 2010) was applied to test if the sex ratio differed statistically from 1:1. The mean length and wet weight of females and males were compared using t-tests with different variances (Zar, 2010). The relationships TL vs CL, TL vs RL and RL vs CL were fitted using linear equations (Y=a+bX), while a power equation was fitted to both WW vs TL and WW vs CL relationships (Y=a∙X^b; Froese, 2006). All relationships were estimated for females and males, separately, and then compared using an analysis of covariance (Zar, 2010). The hypothesis of isometry was tested for the length-length relationships (b=1) and weight-length relationships (b=3) using t-tests (Froese, 2006; Zar, 2010). All statistical tests were performed with a significance level of 5%.

RESULTS

In all samples collected from May 2015 to May 2016, there were only 132 specimens of F. brasiliensis and all of them were analyzed here (65 females and 67 males). The overall sex ratio did not differ significantly from the expected 1:1 (χ²=0.007, p=0.93).

Regarding the maturation stage, all males were mature while most female individuals (48%) were in development. Mature and spent females were respectively 35% and 15% and only one individual (21.09 mm CL) was immature.

Female size ranged from 20.73 to 46.43 mm CL, whereas male size ranged from 20.75 to 32.47 mm CL (Fig. 2). Females were larger and heavier (34.78±6.05 mm CL, 152.61±19.68 mm TL, and 27.45±11.12 g WW) than males (27.07±2.56 mm CL, 123.77±11.37 mm TL, and 14.48±3.98 g WW) (Tab. 1). The TL/CL ratio was significantly lower for females than males (Tab. 1).

Figure 2.
Number of Farfantepenaeus brasiliensis (Latreille, 1817) per carapace length class (CL) caught on the coast of Sergipe, northeastern Brazil, from May 2015 to May 2016.

Table 1.
Length and weight data for females (F) and males (M) of Farfantepenaeus brasiliensis (Latreille, 1817) caught on the coast of Sergipe, northeastern Brazil. TL = total length, RL = rostrum length, CL = carapace length, WW = wet weight, n = sample size, SD = standard deviation, t = Student t-value comparing mean values between females and males, p = probability associated with t-values.

Variable	Sex	n	Mean ± SD	Range	t	p
TL (mm)	F	41	152.61±19.68	100.01−189.18	8.12	<0.01
	M	41	123.77±11.37	98.91−145.90
RL (mm)	F	40	19.95±2.43	13.42−24.50	6.08	<0.01
	M	40	17.27±1.38	14.00−19.70
CL (mm)	F	65	34.78±6.05	20.73−46.43	9.47	<0.01
	M	67	27.07±2.56	20.75−32.47
TL/CL ratio	F	41	4.30±0.16	3.90−4.72	10.52	<0.01
	M	41	4.63±0.12	4.45−5.04
WW (g)	F	65	27.45±11.12	5.73−57.27	8.86	<0.01
	M	67	14.48±3.98	6.31−25.66

All morphometric relationships were significantly different between females and males (Tab. 2). Both TL vs CL and TL vs RL indicated a pattern of positive allometry, while the RL vs CL relationship indicated negative allometry (Tab. 2). On the other hand, WW vs TL relationship indicated a positive allometry pattern and WW vs CL relationship showed negative allometry (Tab. 2).

Table 2.
Intercept (a) and slope (b) for length-length and weight-length relationships for females (F), and males (M) of Farfantepenaeus brasiliensis (Latreille, 1817) caught on the coast of Sergipe, northeastern Brazil. TL = total length, CL = carapace length, RL = rostrum length, WW = wet weight, n = sample size, CI = confidence interval, r² = coefficient of determination, p (b) = values of p for the analysis of covariance used to compare values of b from relationships estimated for females and males.

Relationship	Sex	n	a (CI)	b (CI)	r²	Allometry	p (b)
TL=a+b∙CL	F	41	29.167 (23.258−35.076)	3.459 (3.296−3.623)	0.979	+	<0.001
	M	41	16.161 (8.129−24.193)	4.029 (3.724−4.322)	0.950	+
TL=a+b∙RL	F	41	15.915 (5.228−26.601)	2.468 (2.277−2.659)	0.946	+	<0.001
	M	41	25.202 (12.289−38.115)	2.255 (1.961−2.548)	0.861	+
RL=a+b∙CL	F	41	4.489 (3.071−5.909)	0.436 (0.397−0.476)	0.928	˗	<0.001
	M	41	5.561 (3.245−7.877)	0.436 (0.351−0.522)	0.735	˗
WW=a∙TLb	F	41	3.96x10-6 (1.90x10-6−8.24x10-6)	3.134 (2.988−3.279)	0.979	=	0.040
	M	41	1.66x10-6 (0.47x10-6−6.46x10-6)	3.303 (3.021−3.585)	0.935	+
WW=a∙CLb	F	65	0.002 (0.001−0.003)	2.652 (2.546−2.757)	0.975	˗	<0.001
	M	67	0.001 (0.001−0.002)	2.892 (2.726−3.057)	0.949	=

DISCUSSION

All individuals of F. brasiliensis smaller than 25 mm CL are considered juveniles in southeastern Brazil (Costa and Fransozo, 1999; Costa et al., 2008). In this study, 28 specimens (22% of total) caught were lower than 25 mm CL, however only one female individual was immature. In this way, our data corresponds to an adult population, which is exploited by the shrimp fleet along the coast of Sergipe.

The largest specimens observed on the coast of Sergipe (11°S) (189.18 and 145.90 mm TL for females and males, respectively) were smaller than those recorded in southeastern Brazil (23°S). Mello (1973) recorded maximum sizes of 210 and 180 mm TL for females and males, respectively, while Leite-Jr and Petrere-Jr (2006) found values of 260 and 219 mm TL (both studies in coastal waters off the state of São Paulo). Larger values (266 and 219 mm TL for females and males, respectively) were also recorded off Mexico (21°N) (Arreguín-Sánchez, 1981). All three studies also analyzed data from shrimp fisheries landings. These variations in body size can be influenced by habitat conditions (e.g., water temperature and food/nutrient supply) correlated with latitude (Castilho et al., 2007), where an increase in body size is commonly observed with increasing latitude (Blackburn et al., 1999), in this case in both hemispheres.

The sex ratio of 1:1 observed here is very common in penaeids (Dall et al., 1990), especially for populations occupying relatively constant environments (Geisel, 1972). Likewise, sexual dimorphism is a very common trait in penaeids, with females being larger than males, which has also been observed for F. brasiliensis in our study, and also by Mello (1973), Arreguín-Sánchez (1981), Rabelo-Neto (1985), Leite-Jr and Petrere-Jr (2006), Freitas-Jr et al. (2011), and Souza et al. (2019). This dimorphism reflects a reproductive strategy, as larger size for females allows for larger gonads and higher fertility and, thus, maximizes egg production (Gab-Alla et al., 1990). As most organs of shrimps, including the reproductive ones, are located in the carapace, differences between sexes in the TL/CL ratio are expected, with females having a proportionally larger carapace than males as an adaptation to maximize egg production. Indeed, these differences in the TL/CL ratio are not observed when analyzing juveniles (Carvalho et al., 2019).

Morphometric relationships vary according to the length/age, with younger, smaller individuals having higher slopes than older, larger individuals requiring different equations for conversions (Arreguín-Sánchez, 1981; Primavera et al., 1998; Reis-Jr et al., 2019). Most of the morphometric data available for F. brasiliensis are from juveniles/subadults inhabiting coastal habitats such as estuaries and coastal lagoons (Villela et al., 1997; Branco and Verani, 1998; Pérez-Castañeda and Defeo, 2002; Albertoni et al., 2003; May-Kú et al., 2006; Freitas-Jr et al., 2011; Carvalho et al., 2019), which differ from the equations for adults such as the ones obtained here in the coast of Sergipe.

All length-length relationships estimated here, except for RL vs CL, showed positive allometry corroborating previous records for this shrimp species in southeastern-southern Brazil (Rabelo-Neto, 1985; Leite-Jr and Petrere-Jr, 2006). The same pattern has also been reported for other penaeids (Leite-Jr and Petrere-Jr, 2006; May-Kú and Ardisson, 2012; Segura and Delgado, 2012; Reis-Jr et al., 2019) suggesting that positive allometry for TL vs CL relationship is a common trait for these shrimps. Regarding the RL vs CL relationship, May-Kú et al. (2006) also reported a negative allometry for juveniles of F. brasiliensis off Mexico. However, studies carried out with X. kroyeri indicated differences in the RL vs CL relationship between juveniles and adults, with faster increase (positive allometry) being observed in juveniles (Moraes et al., 2018). Faster increase of the rostrum length during the juvenile stage could be a strategy of this species to avoid predation, as observed in X. kroyeri (Moraes et al., 2018)

The weight-length relationships showed contrasting patterns. When the WW vs TL relationship was estimated, the allometry was positive for both sexes indicating a higher increase in weight than expected by the increase in total length. Mello (1973) recorded a positive allometry for adult males and negative for adult females caught along the southeastern Brazilian coast. Later, an inverse pattern was recorded for the same region (Rabelo-Neto, 1985; Leite-Jr and Petrere-Jr, 2006). According to Leite-Jr and Petrere-Jr (2006), the positive allometric growth presented by females is associated with the gain in weight due to gonad development. For the Mexican coast, this relationship was negative for both sexes (Arreguín-Sánchez, 1981). When the WW vs CL relationship was estimated, the allometry was negative.

It is expected that species with wide distribution ranges and complex life cycles (estuary-open sea migrations), such as F. brasiliensis, experience different environmental conditions that will be reflected in different biological traits such as body size and allometric growth. Differences may also result from differential fishing effort (Couto et al., 2013). However, it is very difficult to assess the exploitation status of the stock off Sergipe due to data scarcity.

Farfantepenaeus brasiliensis is overexploited or threatened with overexploitation along the Brazilian coast and a sustainable management plan should be implemented (Brasil, 2004b). One of the tools suggested was the establishment of closed seasons, which could vary among regions. The closed season defined for Sergipe according to Executive Order N. 14, which was published on October 14th/2004 (Brasil, 2004a) by the Brazilian Ministry of Environment, is from April 1st to May 15th and from December 1st to January 15th, but this measurement is for all shrimps caught in the region. More recently, F. brasiliensis was classified under the category “insufficient data” in a recent evaluation process conducted by the Chico Mendes Institute for Biodiversity Conservation (ICMBio) (Boos et al., 2016). Thus, our results contribute to better knowledge on some biological traits of the red-spotted shrimp F. brasiliensis, especially for northeastern Brazil where there was no previous data.

ACKNOWLEDGEMENTS

The authors are grateful to Diógenes Lopes and artisanal fishers from Aracaju (Sergipe) for the collection of samples used in this study and to Ana Claudia Gaspar and Aline Alves dos Santos Dias for helping with sample processing. This work was supported by the Projeto de Monitoramento Participativo de Desembarque Pesqueiro (PMPDP) and Fundação de Apoio à Pesquisa e Extensão de Sergipe (FAPESE) (Grant numbers 2600.0094374.14.4 and 2600.0099827.15.4). J. Reis-Júnior and T.M.R.R. Barreto thank FAPESE/PMPDP for their scholarships.

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• Zoobank:

  http://zoobank.org/urn:lsid:zoobank.org:pub:A54F80CC-F507-4BF8-973E-9E3C4F03BCFF

Publication Dates

• Publication in this collection
  08 Feb 2021
• Date of issue
  2021

History

• Received
  01 Apr 2020
• Accepted
  23 Sept 2020