Population and growth of queen conch (Lobatus gigas Linnaeus, 1758) in the Sapodilla Cayes Marine Reserve of Belize
© Chan et al.; licensee Springer. 2013
Received: 27 November 2012
Accepted: 13 August 2013
Published: 22 November 2013
The conservation effectiveness of marine protected areas is of substantial dispute. In the Belize Barrier Reef, there are several zones with increasing human activity and harvesting of overexploited species. Marine protected areas are designed to protect endangered species and increase the population size of exploited species. The present study investigated the population and morphological characteristics of Lobatus gigas (Linnaeus, 1758) in the Sapodilla Cayes Marine Reserve (SCMR) at the southernmost tip of the Belize Barrier Reef to estimate the effectiveness of the marine protected area.
A total of 693 L. gigas conches were counted over the 7,200 m2 from 36 transect lines. The densities of juveniles and adult conches in the conservation zone were substantially higher (20.13 and 2.88 individuals/100 m2, respectively) than in the general use zone (5.29 juveniles and 0.58 adults individuals/100 m2). The shell length of adult individuals ranged from 14.4 to 18.5 cm in the conservation zone and from 14.6 to 18.1 cm in the general use zone. A comparison of shell length of juveniles and adults showed no significant differences between the conservation zone (11.39 ± 1.46 cm) and general use zone (11.62 ± 1.24 cm). There was a significant positive correlation for shell length with lip thickness in the conservation zone (Pearson's correlation, r = 0.729, p < 0.001) and general use zone (Pearson's correlation, r = 0.613, p = 0.02).
The function and effectiveness of marine protected areas are discussed using the results of the present study that identifies the primary objective which is to ensure a continuous recruitment of valuable targeted species by the protection of spawning stock biomass that is important for the management of marine protected areas in an environmentally sensitive ecosystem.
Overfishing has resulted in precipitous declines in many exploited populations in coastal ecosystems providing a threat to marine biodiversity (Dayton et al. 1995). As current fishery regulations appear to be inadequate to protect many exploited populations and maintain sustainable fisheries, researchers suggested that marine protected areas in which fishing is controlled and prohibited may be necessary to prevent overfishing of local species (Ballantine 1991;Bohnsack 1993;Lauck et al. 1998;Navarrete 2001). The effects of marine protected areas may increase larval dispersal and influence the movements of adults from the protected area to adjacent fishing habitats as species densities build up within the zones of the marine protected areas (Polacheck 1990;Carr and Reed 1993;DeMartini 1993). Marine protected areas have been promoted as a viable complement to classical forms of fishery management (Roberts and Polunin 1991;Dugan and Davis 1993). Hence, one of the primary objectives of marine protected areas is to ensure a continuous recruitment of commercially valuable targeted species to overfished areas via protection of spawning stock biomass (Rowley 1994). The mechanisms by which spawning stock biomass complements fishing operations in the adjacent fished areas are by the export of individuals from the marine protected areas by the spillover effects and the production of eggs from within the reserve that contributes to the dispersal of larvae outside the reserve by enhancing the recruitment effects (Sarkis and Ward 2009).
In Belize, L. gigas management is regulated by size restrictions. Individuals with a shell length less than 7 in. (178 mm) or a weight of less than 3 oz (85.049 g) are prohibited from harvest. The morphology of L. gigas is influenced by different habitats (Appeldoorn 1994;Martín-Mora et al. 1995) and predator (Delgado et al. 2002). Several fish species make use of discarded conch shells as shelter sites (Wilson et al. 2005). Clerveaux et al. (2005) reported that the morphology of the conch shell is highly plastic.
The shell of L. gigas provides an important function for estimating sexual maturity. L. gigas shells arrest growth when individuals become mature as adults and start to expand their shell lip thickness (Valle-Esquivel 1998). The thickness of shell lips generally increase with age. Therefore, the thickness of lips can be used in assessing age and growth rate of L. gigas (Valle-Esquivel 1998). Understanding the growth rate characteristics is an essential factor that can be used for an estimate of their population dynamics (Valle-Esquivel 1998).
SCMR of Belize has been established in 1996. As yet, comprehensive biological investigations in this area are still lacking. Thus, the status of population, morphology and characteristics of L. gigas in the SCMR is still relatively unknown. In addition, reproductive biological data should be collected to establish morphometric relationships (Carcamo 2006Valle-Esquivel 1998). Our objectives in the present study are aimed (1) to collect morphometric data on siphonal length and lip thickness of L. gigas, (2) to determine the correlation of morphometric data among different zones of protection, and (3) to obtain detailed information regarding population distribution of L. gigas in the SCMR.
The present study was carried out at the southernmost tip of the Belize Barrier Reef in the SCMR, which covers an area of approximately 15,619 ha (125 km2) and includes 14 sand and mangrove cayes. The location of the sampling areas is shown in Figure 1. The SCMR is rich in marine biodiversity since it includes various marine habitats such as mangroves and seagrass beds that provide habitat for a variety of marine organisms. The Belize Barrier Reef is the largest barrier reef in the western hemisphere. It extends for about 260 km, from the northern border of Belize with Mexico to the southern border with Guatemala.
There are three different zones in place at the SCMR, which include the following: conservation zone, preservation zone, and general use zone. Each of these zones has different rules and regulations. The underwater environments are diverse; the substrates/habitats of seabed and their classifications are presented in three categories: dense seagrass habitats, sparse seagrass habitats, and sand flat habitats (Stoner and Waite 1990;Friedlander et al. 1994;Tewfik et al. 1998).
Sampling methods and morphometric data obtained
Individuals of L. gigas were retrieved by rover diving from the three different zones: conservation zone, preservation zone, and general use zone in the SCMR during August 2009. L. gigas population size was estimated using a modified belt transect method. The modified belt transect method was used to collect abundance, distribution, and morphometric data. Each transect was 50 m in length and 4 m in width and started behind the reef crest and run to an approximate depth of 15 m, perpendicular to the shore. A total of 12 transects were laid for each investigated zone. The sites for the L. gigas survey were selected using the method of strategic site selection and stratified random sampling in the three different zones at the SCMR.
The L. gigas identified within the belt transects were counted, and their total shell siphonal length and shell lip thickness were measured by a plastic ruler and caliper; data was then recorded on a waterproof notepad. L. gigas were retransferred to the sampling area after measurements were taken. The shell length was classified as the distance from the spire tip to the anterior edge of the shell. The shell lip thickness was classified as the distance from the mid-lateral region on the lip side of the shell approximately 40 mm from the edge of the shell (Carcamo 2006). Therefore, based on siphonal length and shell lip thickness, L. gigas are characterized in two categories: juveniles and adults (Stoner and Waite 1990;Friedlander et al. 1994;Tewfik et al. 1998). Juveniles are identified with no shell lip or shell thickness less than 4 mm, and adults are identified with lip thickness greater than or equal to 4 mm (Valle-Esquivel 1998).
The monthly averaged sea surface temperature and seawater chlorophyll a values from SCMR during the sampling period of August 2009 were collected from the National Oceanic and Atmospheric Administration (NOAA) and the Sea-viewing Sensor (SeaWiFS).
For each zone in the SCMR, the data on shell siphonal length and shell lip thickness were analyzed to calculate the total densities of juvenile and adult identified in each zones of the marine protected area. The SPSS computer package version 13 was applied for statistical analysis. Since there was no L. gigas recorded in the preservation zone, Student's t test was applied to compare differences in shell length and lip thickness between the conservation zone and general use zone. Furthermore, linear regression was used to estimate the correlation between shell lengths with lip thickness in each investigated zone.
Population and densities of L. gigas
Substrate/habitat categories used in characterizing the sites surveyed and number and proportion of identified conch
Number of conch
Coarse sand bottom dominated by seagrass Thalassia spp. and Syringodium spp.
A: 69 (9.96%)
J: 483 (69.7%)
Coarse sand bottom without seagrass cover
General use zone
J: 127 (18.33%)
Coarse sand bottom with sparse seagrass cover by Thalassia spp. and Syringodium spp.
General use zone
A: 14 (2.02%)
Characteristics of shell morphology for L. gigas
The present study estimated densities of juveniles at 2,012.5 conchs/ha and adults at 287.5 conchs/ha in the conservation zone and densities of juveniles at 529.2 conchs/ha and adults at 58.3 conchs/ha in the general use zone in the SCMR during summer (season 3). Previous studies indicated that L. gigas populations vary seasonally (Stoner and Waite 1990). These authors recorded the highest density (6.73 conch/ha) occurring during the winter of 1987 (season 4) and the lowest record (0.10 conch/ha) occurring during the fall of 1989 in Florida. These comparatively higher densities were due to aggregations of juveniles. The Caribbean Fishery Management Council (1999) recorded the highest densities of L. gigas which was 830 conch/ha. Previous studies conducted in the Bahamas showed that juvenile L. gigas dominated the populations (Stoner and Sandt 1992;Caribbean Fishery Management Council 1999). Stoner and Waite (1990) reported that aggregations caused the high variances that influence the estimates of L. gigas population densities. Our results observed more than 90% juvenile individuals among L. gigas populations, confirming that the high proportion of juveniles is a common phenomenon among L. gigas in nature as in previous studies.
The present study recorded higher abundance of L. gigas in the conservation zone than in the general use zone. Previous studies reported juvenile L. gigas using exclusively shallow dense seagrass habitats as nursery grounds, as adults occupy sand flat habitats and sparse seagrass beds (Randall 1964;Alcolado 1976;Hesse 1979;Weil and Laughlin 1984;Iversen et al. 1987). L. gigas are known to use multiple essential habitats. In relation to the low abundance of adults identified in the marine reserve, this could be related to habitat selection or to the overexploitation of this species within the marine reserve (Carcamo 2006). However, illegal poaching of L. gigas is a general problem in Belize marine protected areas, as identified by fisheries officers. The association of juvenile L. gigas within dense seagrass habitats, sand flat habitats, and sparse seagrass habitats are well established (Alcolado 1976;Hesse 1979;Iversen et al. 1987;Randall 1964;Stoner and Ray 1996;Weil and Laughlin 1984). Results confirmed that juvenile L. gigas had higher abundances in dense seagrass habitat as in previous studies (Stoner and Ray 1993). Field experiments demonstrate that this resulted from habitat choice (Stoner and Waite 1990). The density of juvenile L. gigas was relatively high in dense seagrass habitats of the conservation zone, whereas the density of adult L. gigas was relatively low in this zone. In the general use zone, juvenile L. gigas density was also relatively high in the sand flat habitats and sparse seagrass habitats, and adult density was relatively low in this zone. In the preservation zone, there were no L. gigas being identified, which could be related to the habitat type (sand flat habitat, coral head, coral rubble, and gorgonian corals).
Possible factors of habitat effects have shown that the growth of juvenile L. gigas can be affected by the habitat type in which settlement of larvae is developed (Alcolado 1976;Stoner and Waite 1990;Stoner and Ray 1996). Previous studies conducted by Ray and Stoner (1995) suggested that juveniles' (2 to 5 months old) daily growth rate was higher in dense seagrass habitats than in sand flat habitats. In the present study, analyses were therefore performed that stipulate that dense seagrass habitats do provide a nursery ground for L. gigas. Thus, the habitat type in comparison of mean length and mean lip thickness does have a positive effect on L. gigas size and is in agreement with the aforementioned studies. Hence, the aspect of this study in comparison to the different zones clearly shows that individuals are more densely populated in the conservation zone and general use zone rather than the preservation zone. In addition, one of the most important findings of this research is that the conservation zone in the SCMR provides a better habitat for juvenile L. gigas inside dense seagrass beds.
This study found L. gigas individuals in the conservation zone 3.9 times more abundant than in the general use zone. As conservation work continues to be an urgent environmental need to control overfishing of marine biodiversity, marine protected areas have the potential to protect and ncrease the reproductive output of overexploited populations (Roberts and Polunin 1991;Rowley 1994). Previous studies indicated that the density and biomass of L. gigas within marine reserves result in a significantly increase population size (McField 2000;Huitric 2005). However, identifying the effects between habitat size and heterogeneity and reserve protection are important and often neglected during the process of marine protected area design (Paddack and Estes 2000). Consequently, the distribution and design of different habitat types within the marine reserve has an important role in the sustainable protection of endangered or otherwise valuable species.
The finding of Carcamo (2006) reported no significant correlation between total length and lip thickness. He suggested the fact that mature L. gigas cease their growth in relation to shell length but continue thickening their shell lips. However, previous studies clearly showed the differences of L. gigas shell length and lip thickness in Puerto Rico (Appeldoorn 1994), central Bahamas (Stoner and Ray 1996), and Xel-ha southeastern Mexico (Valle-Esquivel 1998;Navarrete 2001;Rios-Jara et al. 2001). We also identified a significant correlation between total length and lip thickness in adult individuals. Our findings are similar to the results identified in relation to growth rate studies in Mexico (Valle-Esquivel 1998). Environmental condition can also affect shell morphology (Caldeira and Wickett 2003 2005;Berge et al. 2006;Carcamo 2006;McCarthy 2007;Watson et al. 2009;Turley et al. 2010). Another study showed that the shell of L. gigas is characterized by high morphological plasticity (McCarthy 2007). Shell growth is also influenced by the environmental pressure of predators (Delgado et al. 2002) and habitats (Alcolado 1976).
L. gigas are preyed upon by a large variety of predators, including several species of other mollusks, crustaceans, fishes, and marine turtles. Predation pressure is particularly intense during the juvenile stages and diminishes as shell growth increases (Iversen et al. 1987). Possibly, juvenile L. gigas are consumed by upper trophic levels in the marine food web and likewise in the SCMR. This aspect needs further investigation. This study reveals high mean average lip thickness in the sparse seagrass habitat in the general use zone. The difference in growth rate may be explained by the different ecological factors and habitats being identified in various zones of the marine protected area (Stoner and Sandt 1992;Friedlander et al. 1994;Tewfik 1996). Five important factors have been identified by previous literature to influence L. gigas growth rate: habitat types, depth, temperature, salinity, and intra-specific competition (Friedlander et al. 1994;Tewfik 1996). Depth and temperature plays a very important role indirectly on the abundance and quality of food brought through light availability and have therefore shown to affect both density and population size within marine protected areas (Stoner and Sandt 1992;Friedlander et al. 1994;Tewfik 1996).
In conclusion, the present study identified several important aspects for the management of marine protected areas. If the primary objective of marine protected areas is to ensure that fishery activities continue, the protection of environmentally sensitive areas such as spawning sites needs to be protected also considering spawning seasons. The protection of spawning sites and spawning stocks is only the first step in marine conservation, but the main goal is to maintain and likewise improve fishery yields. Therefore, marine reserves are thought to be the best tools to protect species with limited geographic dispersal. In addition, the present study provides useful information of morphological characteristics and population status of the L. gigas population in the SCMR of Belize and will be valuable for future assessments of stock and fisheries harvesting management.
The authors are grateful to the manager of the Toledo Institute for Development and Environment (TIDE), Seleem Chan, for his technical support. The authors would like to acknowledge Dr. Leandra Cho-Ricketts and Dr. Edward Bold for their constructive suggestions. We thank the National Science Council of Taiwan, ROC, grant No. NSC102-2611-M-019-003 for financial support to J.-S. Hwang for partially supporting the salary to IC to complete this manuscript.
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