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Latin american journal of aquatic research

versión On-line ISSN 0718-560X

Lat. Am. J. Aquat. Res. vol.44 no.4 Valparaíso set. 2016 

Research Article


Impact of the clam Arca zebra artisanal fishery upon the population of the neogastropod Voluta musica in eastern Venezuela

Impactos de la pesca artesanal de la almeja Arca zebra sobre la población del neogastrópodo Voluta musica en el oriente de Venezuela


Ana Carolina Peralta1, Patricia Miloslavich1, Alvar Carranza2 & Gregorio Bigatti3

1Departamento de Estudios Ambientales, Centro de Biodiversidad Marina Laboratorio de Biología Marina, Universidad Simón Bolívar, Caracas, Venezuela
Centro Universitario Regional Este-CURE, Sede Maldonado Universidad de la República, Uruguay, Área Biodiversidad y Conservación Museo Nacional de Historia Natural, Montevideo, Uruguay
LARBIM. Centro Nacional Patagónico (CENPAT)-CONICET, Puerto Madryn, Argentina

Corresponding author: Ana Carolina Peralta (
Corresponding editor: José Álvarez Pérez

ABSTRACT. An important ark clam (Arca zebra) artisanal fishery takes place in the east region of Venezuela. Besides the target species, trawling extracts a significant bycatch of several mollusk species including the gastropod Voluta musica, a threatened species according to the Venezuelan Red List of Endangered Species. In this paper we evaluate: 1) the composition of mollusk species in the bycatch, 2) the number of individuals of V. musica caught as bycatch and, 3) the abundance and size structure of the exploited population. Each fishing night, about 27,830 m2 are trawled by each boat extracting on the average 607 kg of A. zebra meat and 19 kg of V. musica (whole shelled animal). This fishery activity potentially captures ~30,000 kg of ark clam meat and 922 kg of V. musica in one week (~95 snails/fishing night/boat). The size structure of the V. musica population at the ark clam bed is significantly smaller than in other nearby sites not impacted by trawling fishing activities. Non-targeted species extracted in the bycatch but consumed by the local inhabitants include the gastropods Chicoreus brevifrons, Phyllonotus margaritensis, P. pomum, Fasciolaria tulipa, Strombus pugilis, a few Trochidae species, and the bivalves Pinctada imbricata, Spondylus americanus, Anadarafloridana, A. notabilis, and Trachicardium muricatum. Individuals of V. musica along with several invertebrates are discarded. We recommend that V. musica should not be neglected as a conservation target, and despite that A. zebra fisheries are considered "artisanal", that more strict regulations should be established on it.

Keywords: Arca zebra, Voluta musica, bycatch, endangered species, artisanal fishery, size structure, Caribbean Sea.

RESUMEN. La pesca artesanal de la almeja arca (Arca zebra) es una de las pesquerías de mayor importancia en el oriente de Venezuela. En esta pesquería se capturan incidentalmente varias especies de moluscos, incluyendo el gasterópodo Voluta musica, considerada amenazada según el Libro Rojo de la Fauna Venezolana. En este trabajo se evalúa: 1) la composición de especies de moluscos en la captura incidental, 2) el número de individuos de V. musica capturados como pesca incidental y 3) la estructura de tallas y abundancia de la población explotada. En cada faena de pesca se arrastran 27.830 m2 por cada barco, extrayendo en promedio 607 kg de A. zebra (sin concha) y 19 kg de V. musica (con concha). La flota completa captura ~30.000 kg de A. zebra y 922 kg de V. musica en una semana (~95 caracoles/noche de pesca/barco). La estructura de tallas de la población de V. musica en el banco de A. zebra es significativamente menor comparada con sitios cercanos no afectados por la pesca de arrastre. Otras especies capturadas incidentalmente son consumidas por los habitantes locales e incluyen los gasterópodos Chicoreus brevifrons, Phyllonotus margaritensis, P pomum, Fasciolaria tulipa, Strombus pugilis, y unas pocas especies de Trochidae, bivalvos Pinctada imbricata, Spondylus americanus, Anadara floridana, A. notabilis, Trachicardium muricatum. Los individuos de V. musica y otros invertebrados son desechados completamente. V. musica debe ser considerada como objeto de conservación, y a pesar que la captura de A. zebra es considerada "artesanal", se deberían establecer regulaciones más estrictas sobre esta especie.

Palabras clave: Arca zebra, Voluta musica, captura incidental, especies amenazadas, pesca artesanal, estructura de tallas, Mar Caribe.



The ark clam Arca zebra (Bivalvia, Arcidae), is distributed along the western shores of the Atlantic Ocean from North Carolina and Bermuda to Brazil (Abbott, 1974). A large natural bed of ark clams occurs on rocky bottoms between 1 and 20 m depth near the coastal population of Chacopata in the Araya Peninsula, Venezuela. This bed covers an area of 70-80 km2 and has been intensively exploited by local artisanal fishermen since 1940 (Lodeiros et al., 2005). The annual production varies between 15,792 and 33,986 ton, although it has reached 40,000 ton year-1 (Mendoza, 1999; Lodeiros et al., 2005). It is the most important economical income for the artisanal fisheries in the country, after the collapse of the sardine fishery, being most of the production destined for canning and local consumption (Trujillo, 1997; Novoa et al., 1998; Gómez-Gaspar, 1999; Diaz et al., 2002). The fishing method is non selective and the catch is classified after landing, and thus many non-targeted species are incidentally caught (bycatch) and landed (Peralta, 2012). After fishing operations, all the organisms landed are cooked together with the clam A. zebra. The shelled mollusks (several bivalve species and gastropods) are extracted from their shells (Gómez-Gaspar, 1999), while the remaining organisms lacking commercial importance such as sea stars, sea urchins, sponges, corals and the gastropod V. musica are discarded with all the empty shells (Peralta, 2012).

Voluta musica (Linnaeus, 1758) is an endemic volutid gastropod restricted to the Venezuelan and Colombian Caribbean and some Caribbean Islands (Clench & Turner, 1964; Gibson-Smith, 1973). It is a gonochoristic species with internal fertilization and development occurring in egg capsules, which contain extra-embryonic food sources in the intracapsular fluid allowing the embryos to hatch as crawling juveniles (Penchaszadeh & Miloslavich, 2001). The species is listed in the Venezuelan Red List of Endangered Species categorized as data deficient (Rodríguez & Rojas-Suárez, 2008), however, in the updated 2015 edition of the list, the species will be assigned a category of "vulnerable" (Peralta & Miloslavich, in press) based on the impact that the clam Arca zebra fisheries along the Araya Peninsula (eastern Venezuelan coast) is having on it and some critical reproductive aspects of the species, such as low fecundity, and a significant proportion of the population affected by imposex (Peralta et al., 2012, 2014).

Although bycatch is widely recognized as a threat to marine biodiversity and fisheries sustainability (Pauly et al., 2002), few studies have quantified bycatch of invertebrates and the effect on their populations in South America (Riestra et al., 2006; Escolar et al., 2009; Morsan, 2009) and specifically, gastropod bycatch is poorly registered (but see Carranza, 2006; Riestra et al., 2006; Carranza & Horta, 2008). The bycatch of benthic invertebrates may have negative impacts on some species, but some other (scavenger gastropods, crabs, and echinoderms) may increase population densities (Collie et al., 1997; Juan et al., 2007; Escolar et al., 2009). In these cases, the opportunistic species would be favored by the increase of food availability, as a result of discarded organisms (Britton & Morton, 1994; Morton, 1995, 2006; Escolar et al., 2009). This may not be the case in the A. zebra fishery, since bycatch is discarded after landing (and on land), and thus we expect a negative impact on the population of V. musica.

In this scenario, this paper aims to: a) provide bycatch and abundance estimates for V. musica on the A. zebra fishery in the Araya Peninsula, b) compare the abundance and size structure of this population with other populations not affected by the A. zebra fishery in the region, and c) report the composition of mollusk species in the bycatch of the A. zebra fishery.


Study area

Sampling was performed in the ark clam bed of "El Mosquito-La Canal", northeastern of Araya Peninsula (Fig. 1). This bed is located in the Venezuelan eastern upwelling ecoregion, characterized by a significant seasonality in water temperature as a response to oceanographic upwelling processes induced by tidal currents, wind, coastal geomorphology and sea-floor bathymetry. The surface water temperature varies between 22°C during December to April and 28°C during May to November (Miloslavich & Klein, 2008).


Figure 1. Study area: northeastern of Araya Peninsula (10°49’4"N, 63°52’7"W).
Bycatch per unit of effort (BPUE) (ind m-2 expressed as the number of V. musica
captured incidentally by one fishing night.



Samples were taken during the ark clam fishing season (February-May, 2011). The fishing gear used to capture the ark clam was a small trawl net 1.5 m wide x 1 m high with 8 cm mesh size, trawled from 7 m long wood boats operated by a crew of 4-5 fishermen. Fishing operations took place during the night (23:00-7:00 h) and consisted of several trawls performed during a few minutes each (7-10 min). One of us accompanied the fishermen during 10 different nights and recorded with a GPS the geographic coordinates at the beginning and the end of each trawl to estimate Total Swept Area (TSA). Organisms captured on each fishing night were kept by the fisherman in polyethylene bags on board.

After landing, the bags were weighted with all the content in order to quantify total landings. From these bags, all V. musica were manually separated, their shell length measured and their whole bodies weighed with a digital scale of 0.001 mm precision. The rest of the benthic invertebrates were collected and identified either as "target species" (A. zebra) or as "others".

Data analysis

In order to characterize the fishing operations, we estimated the mean swept area (SA) for a total of 100 random trawls. The mean SA was calculated using the fishing gear dimensions and geographic positional system data (GPS) at the beginning and end of each trawl. The total swept area by fishing night (TSA) was estimated as the product of total number of trawls/night/boat (TL), registered on board, multiplied by the calculated mean swept area (SA):

TSA = TL x mean SA

Thus, fishing effort is expressed in TSA/night, since we are unable to associate V. musica bycatch to an individual trawl. BPUE (bycatch per unit effort) is expressed as the number of V. musica/TSA/night. Since the capturability coefficient (q, i.e., the fraction of the biomass that is caught by unit of fishing effort) was assumed to equal 1, BPUE is considered a surrogate for abundance of V. musica in the fishing area, and is here reported as ind m-2. We compared the size structure of the population of the fishing area "El Mosquito-La Canal" with three other V. musica populations located few kilometers from the ark clam bed (data from Peralta et al., 2014), where no artisanal ark clam fisheries operate: Isla Caribe (10°41’19"N, 63°51’05"W), Isla Lobos (10°41’26"N, 63°52’28"W) and Bajo Cuspe (10°43’53"N; 63°51’10"W). Mean size of individuals from each population were compared using a one-way ANOVA, after checking ANOVA assumptions.


The mean number of trawls released by one boat/fishing night (TL) was 33 (±10). The mean swept area (SA) was 792.23m2 (±183.55) and the total swept area by fishing night (TSA) was 27,829.35 m2 (±10,553.21) (Table 1). BPUE is expressed as the number of individuals of V. musica captured incidentally by one fishing night/boat and mean BPUE was 95 ± 60 ind m-2 (Table 1, Fig. 1).


Table 1. Swept area (SA) evaluation and Voluta musica bycatch at El Mosquito-La Canal ark
clam bed. SD: standard deviation.


According to BPUE data, the density of V. musica population found in the Arca zebra bed from "El Mosquito-La Canal" was 0.0041 ± 0.002 ind m-2. The total biomass of Arca zebra meat captured by fishing night/boat was 606.75 ± 99.83 kg, while the total biomass of V. musica captured incidentally by fishing night was 19.20 ± 13.10 kg. In each fishing night we registered 12 artisanal fishing boats, and according to the fisherman, the fishing activity takes place usually four days a week (depending on weather and boat conditions). Therefore, according to the data registered, the capture of a single boat represents ~2,400 kg of ark clam meat and ~76.80 kg of V. musica per week. Extrapolation of these data suggests that the whole fleet (12 boats) can capture ~30,000 kg of ark clam meat and 921.60 kg of V. musica in one fishing week. Fishing takes place during ten months in a year (A. zebra fishing is closed during August and December following the National Fishing Laws). In this scenario, and with the V. musica BPUE estimated (~95 snails/fishing night by one boat) we can estimate that the bycatch of V. musica is 4,560 ind week-1 and 182,400 ind year-1. This represents 36,864 kg of V. musica by catch in a year-round basis.

Size structure

The shell length of incidentally captured individuals of V. musica varied from 39.4 to 87 mm for males and 36.2 to 100.7 mm for females (Fig. 2). Table 2 shows the mean size of female and male snails incidentally captured and the mean size of individuals sampled in three other V. musica populations located ~4 km from the ark clam bed where no artisanal ark clam fisheries operate. The ANOVA analysis showed significant differences between female (F = 40.52; P < 2.10"16) and male (F = 57.43; P < 2.10"16) sizes at the four populations, being "El Mosquito-La Canal" the site with the smaller sizes in females and males (Tukey test, 95% family-wise confidence level). Sex ratio in "El Mosquito-La Canal" ark clam bed did not differ from the 1:1 ratio (X2 = 24.5; P < 0.05; n = 341) with 51% being female and 49% males.


Figure 2. Population size structure of Voluta musica
in the Arca zebra bycatch.


Table 2. Mean size of V. musica at El Mosquito-La Canal area (Arca zebra bank) in
comparison to other nearby sites. *ANOVA shows significant differences in mean size
for females (F = 40.52; P < 2.10-16) and male (F = 57.43; P < 2.10-16) (Tukey multiple
comparisons, 95% family-wise confidence level).


Bycatch composition

In addition to V. musica, we registered the capture of other mollusks in the ark clam fishery. These were consumed directly by the local population, and also represented a source of extra income as some of these by-products were sold locally. The fishery’s community reserves for themselves the following species: the gastropods Chicoreus brevifrons, Phyllonotus margaritensis, P. pomum, Fasciolaria tulipa, Strombus pugilis, a few Trochidae species and the bivalves Pinctada imbricata, Spondylus americanus, Anadara floridana, A. notabilis, and Trachicardium muricatum Neither V. musica nor any echinoderm, cnidarian, polychaeta or crustacean found in the bycatch was used as by-product, and was discarded with the empty A. zebra shells.


Although Arca zebra fishing activity occurs in the Araya Peninsula since 1940, few works dealing with benthic community composition have been performed, and most of these only provide species lists of bivalves and gastropods associated to the A. zebra beds in Chacopata (Prieto et al., 2001; Acosta et al., 2007; Licet et al., 2009). In this work, we present the bycatch mollusk species composition for the first time, which includes species listed in the Venezuelan Red Book (the gastropods Voluta musica, Phyllonotus margaritensis, Strombus pugilis, and the bivalve Pinctada imbricata) (Rodríguez & Rojas-Suárez, 2008).

Since the ark clam fishery does not use a selective fishing gear, a significant number of benthic invertebrates are retained in the trawl net. The bycatch of Voluta musica represents a significantly high number of captured individuals (~95 ind/night/boat), which given the low fecundity of the species (Peralta et al., 2012) could cause a reduction in population as has been observed in other species of exploited Caenogastropods (e.g., Torroglosa & Giménez, 2010).

It has also been shown that benthic invertebrate bycatch generates a high mortality of larger individuals, affecting size structure and productivity of benthic communities (Duplisea et al., 2002; Queirós et al., 2006). Thus, this fishery may cause an important impact on the population structure as suggested by the size structure analysis, which showed significant smaller individuals (males and females) in the A. zebra bed compared with neighboring non-fishing sites. In this study, only 1 of 174 females was found measuring over 100 mm in length. Rangel et al. (2011) report a length range of V. musica specimens collected in the same region between 5 and 104 mm, with an average of 56.4 ± 28.3 mm, however, no distinction was done between males and females. A reduction in individual size has been reported previously for other impacted gastropod species, as is the case for a population of the gastropod Cymbiolacca pulchra (Volutidae) in an Australian reef due to the impact of dredging (Catterall et al, 1992).

Our work also showed a higher frequency of capture within the range of the minimal size of sexual maturity, calculated for both sexes in a neighboring site (Isla Caribe): 52 mm for males and 61 mm for females (Peralta, 2012). The probable capture of a significant number of immature individuals is here inferred by the size structure of V. musica captured incidentally, with a mean of 54.61 mm for males and 63.78 mm for females. This could mean that the 40.23% of females and 22.16% of males from bycatch are below the minimal size of sexual maturity. According to Stearns (1992), this kind of impacts could lead to a new steady state in which the population could prevail despite the impact, producing certain responses as: 1) a reduction in the minimum size of sexual maturity, 2) an increase in growth rates, and 3) a higher reproductive effort. Torroglosa & Giménez (2010) demonstrated that the size at first maturity of Zidona dufresnei (Volutidae) has decreased 10 years after fishery exploitation. Blanchard et al. (2004) and Juan (2007) reported several examples based in life histories optimization depending on the energy distribution and the reproductive success, demonstrating that in some cases removing the larger individuals of a population can lead to a type of directional natural selection, resulting in a new and smaller size of sexual maturity. Thus, a reduction in size at sexual maturity size could be occurring in the analyzed population but this still need to be investigated.

In addition to the direct negative impacts on V. musica populations, densities of species that are part of the benthic community impacted by trawling activities may change depending on their ecological function. Opportunistic species may be favored by disturbed environments, while long-live top predators may decline in number during high intensity disturbance (Blanchard et al., 2004). On the other hand, fishing gear effects may favor scavengers and opportunistic carnivores attracted by crushed organisms, and eliminate fragile species (Britton & Morton, 1994; Taylor, 1994; Morton, 1995, 2006; Ramsay et al., 1997, 1998). V. musica is a carnivore species capable of shifting to scavenging habits when fishing discards are available (Von Cosel, 1976; Peralta, 2012), suggesting a plasticity to overcome environment disturbance. However, in the present work, V. musica densities were lower (0.003 ind m-2) than reported for other neighboring sites without trawling disturbance (0.040.25 ind m-2) (depending on time period and location) suggesting an impact on this gastropod population besides the late attainment of sexual maturity (Peralta et al., 2012). However, presumed effects of spatial clustering could be biasing our results, since aggregations of individuals of V. musica are known to occur due to a) mating behavior seasonality, b) oviposition seasonality, c) feeding behavior, and d) selection of an oviposition area (Peralta, 2012; Peralta et al., 2012). V. musica density in the fishing area was low as well as observed for other edible volutids from Patagonia (Bigatti et al., 2008; Bigatti & Ciocco, 2008; Zabala et al., 2013). The authors suggest that the fishing of these species must be regulated due to very low population density plus a slow individual grow rate, late sexual maturity and direct development.


This paper reports new threats for V. musica populations in the northeastern Venezuelan coast that are worth considering not only in future re-evaluations of the status of the species under the Venezuelan Red List Fauna, but also for future IUCN assessments. Clearly, the data we present here shows that there is a clear negative effect of the bycatch in the population structure, however, the species is still present after more than 30 years of trawling, which could be explained by the upwelling phenomena that occurs in this region and that somehow could sustain the population. On the other hand, the A. zebra fishing activity catches incidentally other animals like star fishes that are mollusk predator, therefore if the star fish is "getting off the picture" then their prays such as V. musica could somehow balance the consequences of the trawling impact. Nevertheless, under the precautionary principle, the species should receive some degree of protection. Furthermore we recommend that the establishment of future marine protected areas and/or current management schemes should include V. musica populations as an important conservation target, since this is a fragile species, characterized by low fecundity and density (Penchaszadeh & Miloslavich, 2001; Peralta et al., 2012) and may be threatened by multiple stressors including imposex (Peralta et al., 2014) and bycatch. In addition, it would be also necessary to control the A. zebra fishing activity, particularly regarding regulations on fishing effort (Total Swept Area and duration of fishing seasons). Onboard (or even after landing) discard of live V. musica will probably be the most effective strategy, but depends on the implementation of public awareness campaigns.


We sincerely acknowledge funding by PROVITA and Decanato de Investigación y Desarrollo (USB). Many thanks to the entire fisherman team from Chacopata and Guayacán who share their work with us and let us travel with them on their boats for fishing observation. To the woman and children who work for the arc clams processing in Chacopata (Las Tres Bellezas), for their help on separating our samples from the fishing capture. Centro de Investigaciones Ecológicas de Guayacán (UDO) for field work facilities and support to authors.



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Received: 2 March 2015;
Accepted: 29 May 2016


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