Zoogeography of Elasmobranchs in the Colombian Pacific Ocean and Caribbean Sea

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Zoogeography of Elasmobranchs in the Colombian Pacific Ocean and Caribbean Sea Neotropical Ichthyology, 14(2): e140134, 2016 Journal homepage: www.scielo.br/ni DOI: 10.1590/1982-0224-20140134 Published online: 07 July 2016 (ISSN 1982-0224) Zoogeography of Elasmobranchs in the Colombian Pacific Ocean and Caribbean Sea Andrés Felipe Navia, Paola Andrea Mejía-Falla and José Sergio Hleap1 In order to investigate zoogeographical patterns of the marine elasmobranch species of Colombia, species richness of the Pacific and Caribbean and their subareas (Coastal Pacific, Oceanic Pacific, Coastal Caribbean, Oceanic Caribbean) was analyzed. The areas shared 10 families, 10 genera and 16 species of sharks, and eight families, three genera and four species of batoids. Carcharhinidae had the highest contribution to shark richness, whereas Rajidae and Urotrygonidae had the greatest contribution to batoid richness in the Caribbean and Pacific, respectively. Most elasmobranchs were associated with benthic and coastal habitats. The similarity analysis allowed the identification of five groups of families, which characterize the elasmobranch richness in both areas. Beta diversity indicated that most species turnover occurred between the Coastal Pacific and the two Caribbean subareas. The difference in species richness and composition between areas may be due to vicariant events such as the emergence of the Isthmus of Panama. It is unlikely that the Colombian elasmobranch diversity originated from a single colonization event. Local diversification/speciation, dispersal from the non-tropical regions of the Americas, a Pacific dispersion and an Atlantic dispersion are origin possibilities without any of them excluding the others. Para conocer los patrones zoogeográficos de los elasmobranquios marinos de Colombia, la riqueza de especies de Pacífico y Caribe y sus subáreas (Pacífico costero, Pacífico oceánico, Caribe costero y Caribe oceánico) fue analizada. Las áreas compartieron 10 familias, 10 géneros y 16 especies de tiburones y ocho familias, tres géneros y cuatro especies de batoideos. Carcharhinidae tuvo la mayor contribución a la riqueza de tiburones mientras que Rajidae y Urotrygonidae tuvieron la mayor contribución a la riqueza de batoideos en el Caribe y el Pacífico, respectivamente. La mayoría de los elasmobranquios estuvieron asociados con hábitats bénticos y costeros. El análisis de similitud permitió la identificación de cinco grupos de familias que caracterizan la riqueza de elasmobranquios en ambas áreas. La diversidad beta indicó que el mayor recambio de especies se produjo entre el Pacífico costero y las dos subáreas del Caribe. La diferencia en la riqueza y composición de especies entre las áreas puede ser debida a eventos vicariantes tales como el surgimiento del Istmo de Panamá. Es poco probable que la diversidad de elasmobranquios en Colombia se originara de un único evento de colonización. Eventos locales de diversificación/especiación, dispersión desde las regiones templadas de América, del Pacífico y del Atlántico son orígenes posibles que no se excluyen entre sí. Keywords: Batoids, Beta diversity, Historical biogeography, Richness, Sharks, Vagility. Introduction Dispersal in elasmobranchs depends on active swimming of individuals, contrary to most teleost fishes, in which it Despite the high diversity of elasmobranchs (more than occurs by means of eggs and larvae. Dispersal is then closely 465 species of sharks and 539 batoids), grouped in 178 linked to vagility, which is in turn inversely correlated with genera and 51 families (Nelson, 2006; White & Last, 2012; speciation (Bush, 1975), taxa diversity (Hrabik et al., 2005) Dulvy et al., 2014), there are few zoogeographical studies and endemism (Anderson, 1994). Specifically for sharks, it carried out in this group, and most are focused on the is proposed that the species vagility is directly proportional order Rajiformes (Ishihara, 1991; Last & Yearsley, 2002; to body size, with oceanic species more able to disperse Musick et al., 2004; Last & White, 2011). Elasmobranchs than coastal ones (Musick et al., 2004). Also, it has been are distributed in all the world oceans, from the poles to suggested that vicariant events, such as the Pangea breakup, tropical waters, and from the surface to over 3,000 m depth. played a decisive role in the evolution of distribution Many species have circumglobal distribution, but most are patterns currently observed in some benthic sharks (Musick restricted to a specific geographical area (McEachran & et al., 2004). For batoids, and based on the proposal that Aschliman, 2004; Musick et al., 2004). their fossil record date from the late Jurassic to Paleocene Fundación Colombiana para la Investigación y Conservación de Tiburones y Rayas, SQUALUS. Carrera 60A 11-39, Cali, Valle del Cauca, Colombia. (AFN) [email protected], (PAM-F) [email protected], (JSH) [email protected] 1 Neotropical Ichthyology, 14(2): e140134, 2016 2 Zoogeography of the elasmobranchs in Colombian (Cappetta, 1987), the observed distribution patterns should Material and Methods reflect the Gondwana partition during the final period of the Mesozoic Era (Pielou, 1979; Hallam, 1981; Pitman III et al., Based on the latest elasmobranch checklist of 1993; Last & Yearsley, 2002). Colombia (Mejía-Falla et al., 2007) and specific additions Colombian marine and freshwaters contain 128 species to elasmobranch richness (Mejía-Falla et al., 2011), a (64 sharks and 64 batoids; Mejía-Falla et al., 2007, 2011; database with the distribution of each species in the marine Lasso et al., 2013). Most studies on elasmobranchs in waters of Colombia (Pacific Ocean and Caribbean sea) Colombia have been directed towards biology (e.g. Mejía- was constructed. Since most records do not indicate the Falla et al., 2012, 2014a; Grijalba-Bendeck et al., 2012; exact location of collection or capture, it was assumed that Payán et al., 2011), ecology (e.g. Navia et al., 2007, 2010, the presence of a species is continuous between its most 2011; López-García et al., 2012; Mejía-Falla & Navia, 2011; extreme records. For those species with a single known Mejía-Falla et al., 2014b), fisheries (Acevedo et al., 2007; record, that location was considered the single point of Grijalba-Bendeck et al., 2007), genetics (Hleap et al., 2012), distribution of that species. Since the analyzed waters and only Díaz (1984) carried out a zoogeographical study of correspond to both oceanic and coastal environments, four sharks in the Colombian Pacific. The goal of this manuscript subareas were considered when analyzing the richness was to investigate zoogeographical patterns of the marine and distribution of the elasmobranchs: coastal Caribbean, elasmobranch fauna of Colombia, and hypothesize its origin oceanic Caribbean, coastal Pacific and oceanic Pacific and affinity with other geographic regions. (Fig. 1). Fig. 1. Study area indicating the territorial limits of the Colombian seas. The dotted lines indicate the partitioning into the subareas (Coastal Caribbean, Oceanic Caribbean, Coastal Pacific and Oceanic Pacific) used in this study. Neotropical Ichthyology, 14(2): e140134, 2016 A. F. Navia, P. A. Mejía-Falla & J. S. Hleap 3 Data Analysis. Information on species richness for each of co-variation between two geographical areas (or subareas) group (sharks and batoids) in each area (Pacific and indicate greater faunal similarity and therefore, higher Caribbean) was plotted comparatively to assess both the permeability to each other. It is also an indicator of a decrease overall (both areas) and relative (to each area) contribution in beta diversity or species turnover between the compared of each family to the richness and distribution patterns of areas (or subareas). A presence-absence matrix in the four elasmobranch in Colombian seas. The overall contribution sub-areas was used following Smith & Bermingham (2005). was measured as the number of species of a given family over Rajidae is one of the groups with the highest zoogeographic the total number of recorded species in both areas, while the complexity in Colombia, making it difficult to test any relative contribution was measured as the number of species distributional hypothesis. For this reason, a phylogeographic of a given family over the total number of species in the area method was used to test the hypothesis of the affinity of being evaluated. A similarity analysis at the family level Rajidae to any given region. A phylogenetic tree mapped was performed using Morisita’s similarity index (since it is into a world map was rendered. For this, the sequence and independent of the sample size), and the UPGMA clustering geographic information was downloaded from BOLD algorithm (Unweighted Pair Group Method with Arithmetic systems public records database (Ratnasingham & Hebert, mean; Krebs, 1999). Furthermore the bootstrap (1000 2007) (S1 - Available only as online supplementary file replicates) value of each group was reported. This analysis accessed with the online version of the article at http://www. was performed in PAST software (Hammer et al., 2001). The scielo.br/ni). The data was filtered to gather only sequences shared and exclusive species as well as the genera richness with geographical information. BOLD systems contain in each area were also evaluated. Likewise, the richness by only cytochrome oxidase subunit 1 DNA sequences. Once occupied habitat (coastal, coastal-oceanic, oceanic and deep the sequences and metadata were obtained, a codon-based water), and the habits of the species (bentopelagic, benthic alignment and curation were
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