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Latin american journal of aquatic research

On-line version ISSN 0718-560X

Lat. Am. J. Aquat. Res. vol.45 no.2 Valparaíso May 2017 

Short Communication


Reproduction of Cortez oyster, Crassostrea corteziensis (Hertlein, 1951) in a growing area in the central Mexican Pacific coast


Mariana Mena-Alcántar1, Oscar I. Zavala-Leal1,2, Carlos A. Romero-Bañólos1,2, Javier M.J. Ruiz-Velazco1,2, José T. Nieto-Navarro1,2, Deivis Palacios-Salgado1,2 & Juan M. Pacheco-Vega1,2

1 Posgrado en Ciencias Biológico Agropecuarias, Universidad Autónoma de Nayarit, México
Escuela Nacional de Ingeniería Pesquera, Universidad Autónoma de Nayarit, México

Corresponding author: Oscar I. Zavala-Leal (
Corresponding editor: Cesar Lodeiros

ABSTRACT. The state of Nayarit is the main oyster producer in the Mexican Pacific, this derived from artisanal crops of Crassostrea corteziensis and collecting of wild seeds. Despite its importance, there are no studies on the reproduction of this species in Boca de Camichin, Nayarit, Mexico, which is the area with the highest production (around 90%). The aim of this study was to evaluate sex ratio, condition index (CI), reproductive cycle, and recruitment size for reproduction and its relationship with environmental factors. Sixty oysters were sampled monthly at La Palicienta Lagoon for one year. The organisms were weighed and measured, then fixed with 10% formalin for histological analyses. Subsequently, the gonads were processed by histology. Sex ratio was different from 1:1, with a predominance of females. The condition index (CI) was higher from March to July. Oyster spawning was most prevalent in May, August, and October. The temperature had a positive relationship with reproduction. Recruitment size at reproduction was 57.1 mm. The results of this study could supply useful information about reproduction of the species at La Palicienta and seed collection in Boca de Camichin, Nayarit.

Keywords: Crassostrea corteziensis, reproduction, seed collection, condition index, temperature influence, Mexican Pacific.


In Mexico, the production of oysters is one of the most important fishing activities (Sevilla, 1993). However, a stagnation in oyster production has been observed at the national level from 2000 (51, 315 ton) to 2013 (42, 524 ton) with a tendency to decrease. Nayarit is the third national producer of oysters in Mexico and is the first producer in Mexico's Pacific Coast, reaching in 2013 a total production of >2, 400 ton, which means 43% of the total production of this littoral (SAGARPA, 2015). This production derives mainly from artisanal crops of Cortez oyster Crassostrea corteziensis. In Nayarit State, the aquaculture of C. corteziensis consists of the collection of seeds in sea shell substrates, which are attached to nylon monofilament cords or metal wire strings hanging on mangrove wood posts until they reach an adequate size for consumption (60-70 mm) (Chavez-Villalba et al, 2005); however, to our knowledge, no studies have been reported in the zone about oyster reproduction. To support wild seed collection for aquaculture purposes and commercial fishing exploitation of Cortez oyster, it is important to make research on their reproductive cycle and the environmental factors that govern its reproduction.

The reproductive cycle and the influence of abiotic variables have been characterized on the reproduction of C. corteziensis in many coastal lagoons of Mexico (Cuevas-Guevara & Martínez-Guerrero, 1979; Chávez-Villalba et al., 2008; Rodríguez-Jaramillo et al., 2008; Mazón-Suástegui et al., 2011). For example, in Las Guasimas Lagoon, Sonora State, Mexico, temperature has been reported as the main exogenous factor affecting the reproductive cycle (Chávez-Villalba et al., 2008). Mazón-Suástegui et al. (2011) reported that maximal reproductive activity of C. corteziensis occurs when there is a peak in the content of inorganic particulate matter in Bahía de Agiabampo, Sonora, Mexico. However, in the estuarine system of Boca de Camichín; which is the area that produces over 90% of Cortez oyster in Nayarit, there are not studies on the reproductive biology of this species. The aim of this study was to evaluate the reproduction (sexual proportion, size at first maturity, condition index, and gonadic maturity stages) of Cortez oyster and its relationship with environmental variables in La Palicienta Lagoon, which is the seed collection site in the estuarine system of Boca de Camichin, Nayarit, Mexico.

Oysters for this study were sampled at La Palicienta Lagoon, Boca de Camichin, Nayarit, Mexico (21°43' 26'' - 21°45'41''N and 105°29'2'' - 105°30'22''W). Sixty wild oysters with lengths sizes between 20 and 100 mm were sampled monthly from mangrove roots between September 2013 and August 2014. The organisms were washed and scrubbed to eliminate epibionts, then the size (wide, length, height) and total weight of each oyster were registered. The oyster meat was removed from the shell and weighed. Then, 30 specimens from each sample were dried in an oven (65°C/48 h) to determinate dry tissue weight, and the remaining 30 were fixed in 10% formalin and stored for histological analyses. Also, wet and dry shell weights were determined. During each sampling, temperature and salinity were recorded with a potentiometer and a refractometer, respectively. We obtained 2 L-1 of water at the time of the sampling for chlorophyll-a (Chl-a) determinations using standard spectrophotometry (Parsons et al., 1984). Seston (organic and inorganic particulate matter) was obtained by calcination and gravimetry (Chavez-Villalba et al., 2005). Sex ratio was estimated by dividing the number of females among the numbers of males. A Chi-square (χ2) test was applied to determine whether the sex ratio differed from 1:1. The tissues and shells of 30 specimens were used to calculate the condition index (CI) described by Walne & Mann (1975). The Condition Index (CI) was calculated as the proportion of dry tissue weight (DTW) with respect to dry shell weight (DSW) by the following formula: CI = w1×1000 w2-1, where w1 is the DTW and w2 is the DSW. Gonadal developmental stages were estimated by histology (Hematoxylin-Eosin). Were considered five stages (undifferentiated, development, ripe, spawning, and post-spawning) according to George-Zamora et al. (2003). The monthly relative-frequency of the stages of gonadal development of males and females were calculated. Recruitment size at reproduction was considered when 50% of the specimens presented gonadal maturity (Mazon-Suastegui et al., 2011) and spawning stages, using a logistic model. The relative frequency and frequency-interval accumulated length (5 mm) was calculated. The logistic model was adjusted as follows:

where: M%: percentage of mature specimens, a: intercept, b: slope, c: constant, and L: total length. To determine the relationship of each stage of the reproductive cycle (relative frequency) with each of the measured variables, a Spearman correlation analysis was applied. Statistical analyzes were performed with Statistica 7.1 software (StatSoft).

Environmental parameters recorded during the sampling period of this study are shown in Table 1. The surface water temperature ranged from 25.5°C (December) to 33.0°C (July). Chl-a concentration ranged between 0.000 (April and August) to 17.506 mg m-3 (February). Salinity was lower in summer (4 g L-1 in September) and higher in late winter and early spring (33-35 g L-1 from February to April). Finally, particu-late organic matter varied from 0.0016 to 0.0317 mg L-1 between September 2013 and August 2014 and inorganic matter from 0.0064 to 0.1637 mg L-1 for the same months (Table 1).


Table 1. Environmental factors average values during the sampling period
(September 2013 to August 2014), at La Palicienta Lagoon, Boca de Camichín,
Nayarit, México. OM: organic matter, IM: inorganic matter.


However, according to the x2, sex ratio was only different from 1:1 in October (x2 = 12.0; P = 0.00), March (x2 = 8.0; P = 0.00), April (x2 = 3.77; P = 0.05), and July (x2 = 4.27; P = 0.03). In May, hermaphrodite organisms were found in a percentage of 1.28% with respect to the number of samples obtained. Our results did not show a positive relation between CI and the reproductive process (Table 3). For the sexual proportion was observed that females predominated (Table 2).


Table 2. Females (F), males (M), and sexually
undifferentiated (U) Cortez oyster Crassostrea
at La Palicienta Lagoon, Boca de
Camichín, Nayarit, México (Sept/2013 to Aug/2014).
*Denoted significant difference (P < 0.05).


Table 3. Correlation between environmental
variables and reproduction of Cortez oyster
Crassostrea corteziensis
at La Palicienta
Lagoon, Boca de Camichin, Nayarit, Mexico
(Sept/2013 to Aug/2014). CI: Condition Index.
*Denoted significant difference (P < 0.05).


A resting period from December to February was observed (Fig. 1). Two peaks of oyster spawning occurred during September-November, and March August, but mass spawning peaks occurred in May, August and October (Fig. 1).


Figure 1. Gonadic developmental stages of Cortez oyster
C. corteziensis
in La Palicienta Lagoon, Boca de Camichín,
Nayarit, México (September 2013 to August 2014).


Temperature plays an important role in the reproductive cycle, followed by Chl-a. Spawning stage was predominant in months when the temperature is normally higher (R = 0.90; P = 0.00) (Table 3). The concentration of Chl-a showed a negative relationship with gametogenesis (R = -0.76; P = 0.00) during the reproductive period (Table 3). CI was related with salinity (R = 0.60; P = 0.03) (Table 3).

The remaining variables showed no correlation with CI in C. corteziensis in La Palicienta Lagoon (Table 3). The interval of length at which mature organisms occurred in Cortez oyster was 28-96 mm. However, recruitment size at reproduction (L50) was 57.1 mm (Fig. 2).


Figure 2. Recruitment size at reproduction of the Cortez oyster
C. corteziensis
at La Palicienta Lagoon, Boca de Camichín,
Nayarit, México (September 2013 to August 2014).


This work recorded that female generally predominate throughout the year. This finding is similar to that reported by several authors in oysters of the genus Crassostrea (Rodriguez-Jaramillo et al., 2008; Lenz & Boehs, 2011). The age and size of the organisms can be a determining factor for sex predominance, because the younger and smaller individuals are usually males and the older and larger organisms are females (Velez, 1991). In this sense Lango-Reynoso et al. (2006) report that hermaphrodites comprise the intermediate stage of male-to-female transition in C. gigas. This can be attributed to the protandrous condition that some species of this genus undergo (Chavez-Villalba et al., 2008). In this work, it was found a 1.3% hermaphroditic specimens in May, which is the month of greatest reproductive activity for C. corteziensis, at La Palicienta Lagoon. The proportion of hermaphrodites (1.5%) is similar to that reported for this species in other areas of distribution (Chavez-Villalba et al., 2008) but greater than other species of the same genus, such as C. virginica (0.83%) (George-Zamora et al., 2003), and C. rhizophorae (0.8 to 1.3%) (Lenz & Boehs, 2011). These authors attribute hermaphroditism to the age of specimens and to environmental stimuli; however, Paniagua-Chavez & Acosta-Ruiz (1995) attributed this to the fact that in hermaphroditism, complete removal and reabsorption of gametes of the most recent reproductive cycle and new gametogenesis do not occur. The condition index (CI) can evaluate parameters, such as nutritional status, temporally changes in nutritional reserves, and marketable quality (Chavez-Villalba et al., 2007) and further found about a relationship between the CI and sexually mature bivalves have been reported (Okumus & Stirling, 1998). However, in this study, no correlation between the CI and sexual maturity was found in C. corteziensis, as reported by Chávez-Villalba et al. (2007) and Lango-Reynoso et al. (1999) for C. gigas. In this study, an increased CI was found in the months of January and February, when the highest values of chlorophyll-a were also found, this coinciding with the point made by Thompson & MacDonald (1991), who showed the temporal separation between the reserves accumulation and gamete production in temperate bivalves, as noted by Chávez-Villalba et al. (2005) for C. corteziensis in Bahía de San Francisco, Sonora. We coincide with the last authors in the sense that the autumn-winter period is the time when some temperate species such as C. corteziensis build reserves, which are subsequently employed for gametes development.

Predominant spawning of species occurs in May, August, and October. Development stages is predominant in March, April, and July, and a resting period occurs from December to February. This is similar to that found for this species both in Nayarit and in other localities of the Mexican Pacific (Cuevas-Guevara & Martínez-Guerrero, 1979; Chávez-Villalba et al., 2008; Rodríguez-Jaramillo et al., 2008; Mazón-Suástegui et al., 2011). The occurrence of development stage and spawning during these months is attributable to the increase in water temperature, which is the environmental factor that exerts most control over the physiology of oysters (Chávez-Villalba et al., 2005). Temperature has been proved to be the main environmental factor regulating reproduction in marine bivalves (Thompson et al., 1996). For C. corteziensis, several authors have reported a temperature-related pattern in different parts of the Mexican Pacific; partial spawning occurs during temperature descents and steep increases, while mass spawning takes place at the maximal annual temperatures reported (Rodríguez-Jaramillo et al., 2008; Mazón-Suástegui et al, 2011). In this study, spawning was observed from March to November, but was more important in May and October, when abrupt temperature increases were recorded. The temperature range within C. corteziensis spawned at La Palicienta Lagoon, Boca de Camichín, Nayarit, México was 25.9-33°C. Cuevas-Guevara & Martínez-Guerrero (1979) found that temperature is a determining environmental factor for spawning and that C. corteziensis only spawn at temperatures above 25.5°C. In addition to temperature, it has been reported that this is related with other environmental factors. In this work, a negative correlation between gonadal development and Chl-a concentration was observed which coincides with the lowest temperature recorded. Similar results are reported by Rodríguez-Jaramillo et al. (2008) and Chávez-Villalba et al. (2008) for C. corteziensis and by Chávez-Villalba et al. (2002) for C. gigas. Higher levels of Chl-a and particulate organic matter corresponded to a period of gonadal rest in winter, during which the oyster increased its nutrient and energy reserves, which can be employed to start the gametogenic cycle. Recruitment size at reproduction has been defined in different ways. Mazón-Suástegui et al. (2011) defined it as the time when 50% of the population is in a stage of maturity or when the sum of the stages of maturity and spawning is ≥50%. We define recruitment size in the same manner as these authors. Different sizes at maturity have been established for C. corteziensis; Mazón-Suástegui et al. (2009) found developing gametes from 10 mm shell height in a hatchery cultivated juvenile population. Chávez-Villalba et al. (2008) found mature gonads between 45 and 50 mm in shell height, and Mazón-Suástegui et al. (2011) found mature males from 42 mm in shell height and females from a shell height of 54 mm, under culture conditions in the field, and from hatchery seed production. In this study, we found that recruitment size at reproduction for C. corteziensis is 57.1 mm in shell height. The first sexual maturity depending on a complex interaction of endogenous and exogenous cues including genetic mechanisms triggered at the onset of gametogenesis (Cruz et al., 2000), temperature variations and availability of food related to the geographic range of populations of the same species (Chung et al., 2005).

In conclusion, this study shows that food availably was not related with sexual maturity or spawning, while the temperature is the environmental factor that determines the spawning events in Cortez oyster Crassostrea corteziensis in the study area. Reproductive activity occurs between 25.9 and 33°C but with greater intensity when abrupt temperature changes occur reaching average temperatures between 30 and 31°C. Therefore, it is recommended to producers in La Palicienta Lagoon to place the seed collectors when the surface temperature reaches 29°C to wait for the spawns and to collect the oyster seed.


The authors wish to thank the SCPPGyA "OSTRJCAMICHIN", S.C. de R.L. de C.V. for donating the oysters, and the Sistema Producto Ostión of Nayarit State for their support and the facilities provided. M.M.-A. is the recipient of a CONACyT Master Fellowship. I.Z.-L., J.M.J.R.-V., J.T.N.-N., D.P.-S., and J.M.P.-V. are SNI Fellows.



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Received: 29 August 2016;
Accepted: 19 January 2017


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