Fish assemblages in tropical estuaries of northeast Brazil: A multi-component diversity approach
Introduction
Estuaries, interfaces between land and sea, are the site of a range of hydrological (e.g. river discharge), oceanographic and anthropogenic processes. These complex processes create habitats with intense thermal and salinity gradients and variable nutrient and pollutant concentrations (Wolanski, 2007). They notably drive fish patterns of distribution in space and time, and shape the diversity of assemblages (Elliott et al., 2007, Potter et al., 2015). Estuaries are ecologically essential as feeding and breeding grounds, providing appropriate habitats for different stages of the fish life cycle. In addition, they are a migratory pathway for both anadromous and catadromous species (Elliott et al., 2007, Potter et al., 2015). Furthermore, the high socio-economic value of their products, especially as a source of income and food, is widely recognized (Glaser and Diele, 2004, Isaac et al., 2009). However, global environmental change and direct human pressures impact the water quality and biodiversity of estuarine ecosystems (Garcia et al., 2003, Halpern et al., 2008, Borja et al., 2010, Li and de Jonge, 2015). Notably, major changes in fish diversity from the effects of mangrove removal, shrimp farming and agricultural development have been recently documented for estuaries worldwide in terms of assemblage composition, decrease in number of species, feeding groups and larger fishes (Blaber, 2008, Blaber, 2009, Blaber, 2013, Weerts and Cyrus, 2002, Singkran and Sudara, 2005, Oribhabor and Ogbeibu, 2010, Viana et al., 2010, Viana et al., 2012).
Aquatic resources are of fundamental importance for human food and for the development of Brazil, which has one of the longest coastlines in the world (9th longest worldwide) (Burke et al., 2001). Recently, Pinheiro et al. (2015) stated that the Brazilian aquatic biodiversity is in peril, notably with regard to fishes, in many cases due to habitat degradation. However, it is widely acknowledged that the success of many fisheries resources is dependent on the quantity and quality of the habitat (Lindall and Thayer, 1982; Auster et al., 1996). The northeast is one of the most densely populated coastal regions in Brazil. In Pernambuco state, the degradation of coastal ecosystems is most severe around the main urban centre, Recife, mainly due to domestic pollution, industrial activity and habitat degradation and loss. These coastal ecosystems are also threatened by fishing (Elfes et al., 2014) because of the impact on the habitat (Turner et al., 1999) and/or fishes at the top of the trophic networks (e.g. trophic downgrading, Estes et al., 2011). Along the Pernambuco coast are located 34 fishing communities, with ca. 12 000 fishermen (Lessa et al., 2006). To supply food and satisfy economic needs, aquatic resources are intensively exploited using multiple techniques and multispecies artisanal fisheries (Frédou et al., 2006, Frédou et al., 2009a, Frédou et al., 2009b). The majority of stocks are either fully or over-exploited, and there is little room for expansion into new fisheries (MMA, 2006). Despite all these threats, the degree of dependency on and use of estuarine ecosystems, even in urban areas, remains very high (MMA, 2002).
The diversity of the threats and their intensification in the Pernambuco estuaries (Lira et al., 2010a, Lira et al., 2010b) highlight the need to monitor fish biodiversity for management and sustainable use purposes. While several studies dedicated to the diversity of fish assemblages have been carried out in this area (Paiva et al., 2009, Paiva and Araújo, 2010), there is still a need to assess and quantify their state in the main estuaries subjected to contrasting human pressure and environmental conditions. For this purpose, the diversity of fish assemblages can be monitored more exhaustively by a multi-component diversity approach (e.g. Wilsey et al., 2005; Mérigot et al., 2007, Gaertner et al., 2010, Lefcheck et al., 2014). This approach consists in assessing different components, or facets, such as species number and evenness, and also components that explicitly take into account the degree of difference among species. This reflects the taxonomic relatedness between species, on the basis of which diversity indices can be more sensitive to environmental and/or human drivers (Clarke and Warwick, 1998). Monitoring a single diversity component using one or few diversity indices leads to an incomplete description of the diversity of assemblages, and in turn important responses in the face of environmental and/or human drivers could be missed (e.g. Stuart-Smith et al., 2013, D’agata et al., 2014).
In this context, the aim of this work is to investigate the fish diversity assemblages on the basis of a multi-component approach, comparing tropical estuaries subjected to contrasting environmental conditions and human pressure. Firstly, complementary diversity indices were quantified for each estuary. Secondly, the typology of fish assemblages among these estuaries was characterized using a recent multivariate analysis, the Double Principal Coordinates Analysis (DPCoA), that explicitly takes into account species differences (here, taxonomic relatedness). Overall, this study provides an assessment of the state of assemblages among the studied tropical estuaries of northeast Brazil, and demonstrates that a multi-component approach of this kind can be useful for ecosystem-based fishery management in estuarine ecosystems.
Section snippets
Study area
Four estuaries located along the coast of Pernambuco State were studied: Itapissuma, Suape, Sirinhaém and Rio Formoso (Fig. 1). The estuarine complex of Itapissuma is located to the north of and closest to the Recife and Olinda urban centres, and is the most strongly influenced by landscape alterations, industrial discharges, shrimp farming and high fishing pressure (CPRH, 2003, Lira et al., 2010a, Gondim, 2015). The other three estuaries are located south of Recife: the Suape estuary has
Patterns of fish assemblage indices
A total number of 122 fish species were identified in the 34 samples of the study area. The species accumulation curve did not stabilise towards asymptotic values (Fig. 2), as is usually the case in ecological studies, due to the large number of rare species present in natural communities. However, the slope at the end of the curve suggested that the number of additional samples necessary to collect the species more exhaustively with the fishing gear used should only be slightly higher. It is
Patterns of fish diversity in estuaries
Our results highlighted that fish assemblages differ among the four estuaries studied in Pernambuco state. Dominance-evenness profiles varied between Itapissuma and Suape compared to those of Rio Formoso and Sirinhaém (Table 2 and Fig. S1 in Supplementary Materials). In addition, Itapissuma had higher mean number of species and individuals per sample than the other estuaries (Fig. 3), but individuals were more closely related from a taxonomic point of view, especially compared to Sirinhaém and
Acknowledgments
We thank three anonymous reviewers for their constructive comments on an earlier version of the manuscript. We thank all students and researchers who participated in field collections. We are also grateful to the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for providing a research grant to F Lucena Frédou, T. Frédou and B. Ferreira. This work was carried out under the National Institute on Science and Technology in Tropical Marine Environments - INCT-AmbTropic (CNPq
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