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Reproductive Cycle of the Rock Oyster, Striostrea Prismatica (Gray, 1825) from Two Locations on the Southern Coast of Ecuador

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Reproductive Cycle of the Rock Oyster, Striostrea Prismatica (Gray, 1825) from Two Locations on the Southern Coast of Ecuador Aquaculture Research, 2014, 1–11 doi:10.1111/are.12601 Reproductive cycle of the rock oyster, Striostrea prismatica (Gray, 1825) from two locations on the southern coast of Ecuador Alfredo Loor & Stanislaus Sonnenholzner Escuela Superior Politecnica del Litoral (ESPOL), Centro Nacional de Acuicultura e Investigaciones Marinas (CENAIM), Guayaquil, Ecuador Correspondence: A Loor, Escuela Superior Politecnica del Litoral, Centro Nacional de Acuicultura e Investigaciones Marinas, Km 30.5 Vıa Perimetral, P.O. Box: 09-01-5863, Guayaquil, Ecuador. E-mail: [email protected] Abstract Introduction The reproductive cycle of the rock oyster Striostrea The rock oyster, Striostrea prismatica (Gray, 1825) prismatica was determined at two fishing areas, Gen- (Mollusca, Bivalvia, Otreidae), is a commercially eral Villamil and Ayangue (located on the southern valuable species along the Pacific Ocean coast of coast of Ecuador), between May 2012 and April America, from La Paz, Baja California (Mexico), to 2013. Monthly sampling campaigns were performed Mancora, Tumbes (Peru) (Poutiers 1995). It is at both locations. The tissues were histologically found on rocky intertidal and shallow subtidal sub- examined to determine gonadal index (GI), oocyte strates down to 7 m of depth (Fournier 1992; Coan development, follicular area coverage and sex ratio. & Valentich-Scott 2012). The meat is highly appre- Surface seawater temperature, salinity and chloro- ciated as food plate and the valves are used by phyll a concentration were measured during sam- craftsmen, thus significantly contributing to the plings. Our results show a similar annual economy of coastal communities of Ecuador. Never- reproductive pattern at both locations. The GI theless, in coastal fishing villages such as Ayangue reached maximum values during the summer. Oys- (Santa Elena), a shortage of individuals and the ters reached highest ripeness in January and Febru- extraction of smaller oyster are reported (Artisanal ary, while spawning occurred in February–March. Fishermen 2012, personal communication), sug- Gametogenesis was linked to a consistently increas- gesting a possible over-exploitation of the resource. ing follicular area (from 1.3 Æ 0.7% to 85.8 Æ To mitigate human and environmental impacts on 7.8%) and associated to surface seawater tempera- S. prismatica populations, laboratory studies focus- ture. Spawning coincided with warm water tempera- ing on broodstock maturation, spawning and larval ture fluctuations and a seawater salinity decrease. development have been recently conducted (Loor No correlation was found with Chlorophyll a concen- 2012; Arguello-Guevara,€ Loor & Sonnenholzner tration. The sex ratio of sampled populations was 2013). However, studies to determine the reproduc- 1:1, suggesting that oysters sizing more than 10 cm tive cycle of this oyster in Ecuadorean waters have in shell length present a stable sex proportion in the not been undertaken to our knowledge, which are population. The diameter of mature oocytes was sig- essential for proper management policies of the nificantly reduced (32.7%) during histologically resource and efforts for its restoration. preparations in comparison to fresh oocytes. Our The reproductive cycle of marine bivalves vary study provides useful information of environmental among species and locations (Chavez-Villalba, factors that may control the observed gametogenesis Barret, Mingant, Cochard & Le Pennec 2002; and spawning activity of S. prismatica. Serdar & Lok€ 2009), and is regulated by exoge- nous and endogenous factors (Mackie 1984; Keywords: Striostrea prismatica, gametogenesis, con- Barber & Blake 2006). Water temperature and dition index, sex determination, Crassostrea iridescens food availability are the main factors that control © 2014 John Wiley & Sons Ltd 1 Reproductive cycle of Striostrea prismatica A Loor & S Sonnenholzner Aquaculture Research, 2014, 1–11 gonadal development (Giese & Pearse 1974; Frıas- Materials and methods Espericueta, Paez-Osuna & Osuna-Lopez 1997; Chavez-Villalba, Cochard, Le Pennec, Barret, Study areas and biometric data samples Enrıquez-Dıaz & Caceres-Mart ınez 2003; Fabioux, The samples of S. prismatica were obtained at Huvet, Souchu, Le Pennec & Pouvreau 2005), and General Villamil (2°38034″ S; 80°26009″ W) and influence fertility and gamete quality (Gabbott Ayangue (1°58005″ S; 80°45021″ W) between 1983; Utting & Millican 1997; Park, Lee, Choy, May 2012 and April 2013. Both sites are located Choi & Kang 2011). Microscopy and histology on the southern coast of Ecuador (Fig. 1). We provide accurate analysis of gametogenic stages, sampled fifteen oysters off the coast at water while oocyte size is determined by image analysis depths between 3 and 6 m every month. Only (Lango-Reynoso, Chavez-Villalba, Cochard & Le oysters having shell length of 10 cm or larger Pennec 2000; Meneghetti, Moschino & Da Ros were considered in the study. Oysters were trans- 2004; Enrıquez-Dıaz, Pouvreau, Chavez-Villalba & ported in plastic coolers to the Centro Nacional de Le Pennec 2009). Acuicultura e Investigaciones Marinas (CENAIM) The annual reproductive cycle for S. prismatica at San Pedro de Manglaralto (Santa Elena – Ecua- formerly referred to as Crassostrea iridescens and/or dor). Fouling organisms on oyster shells were Ostrea iridescens, has been investigated in other removed by brushing and scrubbing. Oyster were latitudes (Cuevas-Guevara & Martınez-Guerrero measured with a calipre to the closest 1 mm to 1979; Fournier 1992; Frıas-Espericueta et al. determine shell dimensions (length, height and 1997). Histological analyses carried out by Cue- width) and soft body weight was determine in an vas-Guevara and Martınez-Guerrero (1979) and electronic balance with precision 0.1 g. Dimen- Fournier (1992) on S. prismatica oysters collected sional terms were applied according to Coan and year round in the Northwest coast of Mexico and Valentich-Scott (2012). Pacific coast of Costa Rica, respectively, evidenced a clear seasonal reproductive cycle, where temper- ature and salinity appear to modulate the gameto- Environmental parameters genesis and trigger their spawning. Warm tropical Water samples were collected at both locations waters of the Panama Bay and cold subtropical during oyster sampling for temperature, salinity waters of the Humboldt Current from the south, and chlorophyll a determination. Temperature and seasonally influence the oceanography of the salinity were measured on site with a mercury Ecuadorean coast. These water masses display thermometer and refractometer respectively. marked thermohaline variability (Cucalon 1989), Chlorophyll a was analysed in the laboratory fol- showing an appealing opportunity for studying the lowing the spectrophotometric method (Greenberg, effect of environmental parameters on the repro- Clesceri & Eaton 1992). ductive cycle of marine invertebrates, which can be applied to aquaculture techniques. Determinations of physical and chemical param- Histology eters that regulate broodstock conditioning are Transverse 5-mm-thick cross-sections of the poster- essential for aquaculture projects of S. prismatica. ior visceral mass (between the pericardial cavity A one-year baseline study was performed in wild and digestive gland) from each oyster were sliced, populations of S. prismatica at two contrasting placed in histology cassettes and fixed in Davidson’s environmental locations of the southern coast of solution for 48 h (Howard & Smith 1983), dehy- Ecuador; Ayangue, characterized by coastal drated in a graded series of ethanol solutions, marine waters; and General Villamil, characterized cleared in xylene and embedded in paraffin. Paraffin by a mixture of coastal marine and estuarine blocks were then sliced into 4 lm on a rotary waters of the Gulf of Guayaquil. Our study aimed microtome, mounted on slides and stained with two objectives: (i) to determine the reproductive Haematoxylin-eosin for analysis. Pictures were cycle of wild S. prismatica by determining mean taken with a Olympus CH-2 microscope coupled to monthly Gonadal index (GI), follicular area, oocyte a Nikon E995 camera. development and population sex ratio; and (ii) to Oyster gonads were analysed microscopically for correlate reproductive patterns with changes in sex differentiation. Gonadal development was environmental factors. 2 © 2014 John Wiley & Sons Ltd, Aquaculture Research, 1–11 Aquaculture Research, 2014, 1–11 Reproductive cycle of Striostrea prismatica A Loor & S Sonnenholzner Figure 1 Map of the southern Ecuadorian coast showing sam- pling sites (grey circles). classified into six stages according to the classifica- Lango-Reynoso et al. (2000) and Rodrıguez- tion system described by Cuevas-Guevara and Jaramillo, Hurtado, Romero-Vivas, Ramırez, Manz- Martınez-Guerrero (1979) and Fournier (1992) for ano and Palacios (2008): Striostrea prismatica: Resting (S0), Early develop- pffiffiffiffiffiffiffiffiffiffi ¼ =p ment (S1), Late development (S2), Ripe (S3), TD 4a Spawning (S4) and Spent-Reabsorbing (S5). A total of 45 oocytes located at the centre of the follicle with visible nucleus were measured at 10 randomly selected regions per slide. A digital Mean gonadal index calibration was previously carried out to scale Monthly mean Gonadal Index (GI) was determined measurements in lm units. for each sampling site. The GI ranged between 0 Fresh ripe oocytes were also measured and (if all oysters in the monthly sample had a resting compared to histologically processed oocytes. Fresh stage S0) and 1 (if all oyster were in the ripe
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