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Shell Shape Variation of Queen Conch Strombus Gigas (Mesograstropoda: Strombidae) from Southwest Caribbean

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Shell Shape Variation of Queen Conch Strombus Gigas (Mesograstropoda: Strombidae) from Southwest Caribbean Shell shape variation of queen conch Strombus gigas (Mesograstropoda: Strombidae) from Southwest Caribbean Edna J. Márquez, Natalia Restrepo-Escobar & Francisco L. Montoya-Herrera Facultad de Ciencias, Universidad Nacional de Colombia, Medellín, Colombia. Calle 59A No 63 – 20 Bloque 19 A Laboratorio 310, Medellín, Colombia; [email protected], [email protected], [email protected], [email protected] Received 15-X-2015. Corrected 10-VI-2016. Accepted 11-VII-2016. Abstract: The endangered species Strombus gigas is a marine gastropod of significant economic importance through the Greater Caribbean region. In contrast to phenotypic plasticity, the role of genetics on shell variations in S. gigas has not been addressed so far, despite its importance in evolution, management and conservation of this species. This work used geometric morphometrics to investigate the phenotypic variation of 219 shells of S. gigas from eight sites of the Colombian Southwest Caribbean. Differences in mean size between sexes and among sites were contrasted by analysis of variance. Allometry was tested by multivariate regression and the hypothesis of common slope was contrasted by covariance multivariate analysis. Differences in the shell shape among sites were analyzed by principal component analysis. Sexual size dimorphism was not significant, whereas sexual shape dimorphism was significant and variable across sites. Differences in the shell shape among sites were concordant with genetic differences based on microsatellite data, supporting its genetic background. Besides, differences in the shell shape between populations genetically similar suggest a role of phenotypic plasticity in the morphometric variation of the shell shape. These outcomes evidence the role of genetic back- ground and phenotypic plasticity in the shell shape of S. gigas. Thus, geometric morphometrics of shell shape may constitute a complementary tool to explore the genetic diversity of this species. Rev. Biol. Trop. 64 (4): 1585-1595. Epub 2016 December 01. Key words: geometric morphometrics, phenotypic plasticity, phenotypic stocks, fishery management, Colombian Caribbean. The queen conch Strombus gigas Lin- & Baqueiro-Cárdenas, 2006). Thus, the mea- naeus, 1758 is a large marine gastropod of sures based on shell length and lip thickness to significant economic importance through the estimate the size at sexual maturity are used as Greater Caribbean region (Theile, 2005). In a parameter for management regulations and this endangered species, the genetic patchi- sustainability of fisheries (Wenner, Fusaro, & ness seems to characterize the natural popula- Oaten, 1974; Conand, 1981; 1989; Appeldoorn, tions since studies using neutral markers have 1988). However, the morphometric studies shown both gene flow (Mitton, Berg, & Orr, have not been used to define phenotypic stocks 1989; Campton, Berg Jr, Robinson, & Glazer, so far, this information allows to determine the 1992) and genetic structure either at isolated way the stock responds to exploitation. This sites or at micro-scales across different Carib- issue is relevant since the ontogenic rates influ- bean areas (Mitton et al., 1989; Tello-Cetina, ence many population attributes that are inti- Rodriguez-Gil, & Rodríguez-Romero, 2005; mately related to population dynamics (Garrod Márquez et al., 2013). & Horwood, 1984). The morphometrics studies in S. gigas On the other hand, the morphometric stud- has been used to solve ecological questions of ies have documented sexual size dimorphism fisheries interest (Randall, 1964; Ávila-Poveda in S. gigas shell (Randall, 1964; Galindo-Pérez, Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 64 (4): 1585-1595, December 2016 1585 2009) and other members of the genus Strom- is unknown, although this information may bus such as S. gibberulus, S. flammeus (Abbott, complement the genetic studies in this region 1949), S. pugilis (Colton, 1905; Galindo-Pérez, (Márquez et al., 2013). The genetic control 2009), S. canarium (Cob, Arshad, & Idris, of the shell shape cannot be overlooked in 2008), and S. costatus (Galindo-Pérez, 2009). populations of S. gigas from Colombian San Similarly, the sexual shape dimorphism in the Andrés archipelago because they are structured shell has been reported in S. gigas, S. costa- in three genetically different groups. Further- tus (Galindo-Pérez, 2009), S. pugilis (Colton, more, both the bathymetry of San Andrés 1905; Galindo-Pérez, 2009) and other no phy- archipelago (Andrade, 2001) that limit the logenetically related snails like Buccinun unda- queen conch dispersion among sites, as well as tum (Hallers-Tjabbers, 1979), Nucella lapillus the environmental and fishing variable condi- (Son & Hughes, 2000), Pomacea canaliculata tions, may induce phenotypic differences in the (Estebenet, Martín, & Burela, 2006) and Buc- queen conch shells. cinanops globulosus (Avaca, Narvarte, Martín, & Van der Molen, 2013). MATERIALS AND METHODS In addition, evolutionary studies of shell in S. gigas have evidenced plastic responses Specimens and study area: A total of to environmental variations (Alcolado, 1976; 219 shells of adult individuals of S. gigas were Stoner & Davis, 1994; Martín-Mora, James, & assessed using geometric morphometric analy- Stoner, 1995) and to predators under controlled sis. These samples were collected at different conditions (Delgado, Glazer, & Stewart, 2002). sites in the San Andrés archipelago, which Likewise, the plastic responses to environ- are separated by depths ranging from 100 to mental variations have been found in other 1 500 fathoms that impede the dispersal of snail species (Pascoal et al., 2012; Kistner & juveniles and adults among these sites (Fig. 1). Dybdahl, 2013; Gustafson, Kensinger, Bolek, In this area, the population genetics of S. gigas & Luttbeg, 2014; Solas, Hughes, Márquez, & shows a moderate genetic structure among Brante, 2015). During ontogeny, these plastic three regions of the San Andrés archipelago: responses to environmental heterogeneity con- Southern (South-South-West and East-South- stitute a key factor in the potential of species to East atolls); Northern (Roncador, Queena and colonize, survive and reproduce; abilities that Serrana atolls) and most Northern, near to allow them to persist under diverse environ- Jamaica (Serranilla atoll, Alice shoal and Bajo mental conditions and expand its distribution Nuevo atoll) (Márquez et al., 2013). On the range (Stearns, 1989). However, it remains to other hand, the fishing pressure is differential address whether the variations in shell shape of among the sites because some of them are S. gigas reflect genetic differences as well as subjected to artisanal (South-South-West and it occurs in other gastropods (Johannesson & East-South-East atolls, Roncador, Serrana) and Johannesson, 1996; Conde-Padín, Caballero, industrial fisheries (Queena, Serranilla atoll, & Rólan-Alvarez, 2009; Martínez-Fernández, Alice shoal and Bajo Nuevo atoll). Paes de la Cadena, & Rolán-Alvarez, 2010; Zieritz, Hoffman, Amos, & Aldridge 2010; Geometric morphometrics: Ten land- Pascoal et al., 2012). marks of type II (Bookstein, 1991) were identi- Thus, in this work, geometric morpho- fied on digital photographs of shells (Fig. 2). To metric analysis was used to address the effect reduce peripheral optical distortion, each shell of genetics and geographic origin on S. gigas was photographed in the centre of the visual shell size and shape across a broad area in field, and landmarks were digitized twice on the Colombian Southwestern Caribbean (San the set of 219 shells. Digital precision was Andrés archipelago). In this area, the spa- estimated by using the “Repeatability” index tial phenotypic variation of S. gigas shell (individual variance / total variance) in a model 1586 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 64 (4): 1585-1595, December 2016 85º0’0” W 80º0’0” W 75º0’0” W 19º0’0” N 19º0’0” 80º0’0” W 79º0’0” W 78º0’0” W 77º0’0” W 76º0’0” W 15º0’0” N 15º0’0” 16º0’0” N 16º0’0” 11º0’0” N 11º0’0” 15º0’0” N 15º0’0” 14º0’0” N 14º0’0” 13º0’0” N 13º0’0” 12º0’0” N 12º0’0” 11º0’0” N 11º0’0” Fig. 1. Sampling sites of queen conch S. gigas from San Colombian Andrés archipelago, Southwest Caribbean. Fig. 1. Sitios de muestreo de S. gigas en el archipiélago colombiano de San Andrés, Caribe suroccidental. Fig. 2. Ten landmarks type II measured as coordinates of S. gigas shells. Numbering on the landmarks denotes the arrangement followed during digitization. Fig. 2. Diez puntos anatómicos de referencia tipo II empleados como coordenadas para las conchas de S. gigas. La numeración de los puntos de referencia denota el arreglo seguido durante la digitalización. Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 64 (4): 1585-1595, December 2016 1587 II one-way ANOVA on repeated measures were conducted in R 2.15.2 (www.r-project. (Arnqvist & Mårtensson, 1998) using VAR org) and RWizard (Guisande, 2015). module of the software CLIC V.70 (Dujardin, 2013). Raw data of coordinates were submitted Shape variation: The effect of sex (Ser- to Generalized Procrustes analyses to generate ranilla atoll, Alice shoal and Bajo Nuevo atoll) ‘‘partial warp’’ scores and uniform components and geographic origin on shell shape of S. gigas (Rohlf, 1990; Rohlf & Slice, 1990) as shape from San Andrés archipelago was explored by variables, using the modules COO and MOG using principal component analysis. Statisti- of the software CLIC V.70 (Dujardin, 2013). cal significance between pair-wise Euclidean
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