Recreational activities trigger changes in meiofauna and free-living nematodes on Amazonian macrotidal sandy beaches
Introduction
Sandy beaches occur in nearly all coastal zones in temperate and tropical areas (McLachlan and Turner, 1994), and historically these environments play an important role as locations for recreation upon which tourism development has been based (Orams, 2003). Beach recreation includes a wide array of activities, such as camping (Hockings and Twyford, 1997), use of vehicles (Schlacher and Thompson, 2008), and other recreational pursuits (Fanini et al., 2014). Overall, these activities can dramatically alter the natural physical characteristics of beaches through compaction, rutting, and displacement of the sand matrix (Anders and Leatherman, 1987; Priskin, 2003), which influence benthic communities either directly or indirectly (Brosnan and Crumrine, 1994; Brown and Taylor, 1999).
Meiofauna, and particularly free-living marine nematodes, comprise an important component of the benthic biota in both abundance and biomass. They are also closely related to other organisms, playing multiple roles, such as nutrient recycling and transferring energy and matter into benthic and pelagic trophic food webs (see Giere, 2009 for review). The meiobenthic fauna that inhabits sandy beaches is generally dominated by harpacticoid copepods and nematodes, with the dominance of one group over the other depending on sediment grain size (see Giere, 2009 for review). Although the presence of nematodes is independent of sediment characteristics (e.g. grain size, sorting, shape) (Vanaverbeke et al., 2000), they are generally more abundant in fine sands, while harpacticoid copepods are more important in coarse sediments (McLachlan and Brown, 2006; Giere, 2009).
Benthic organisms on sandy beaches are one of the most effective tools for assessing environmental variations in the habitat (Coull and Chandler, 1992; Giere, 2009; Félix et al., 2016). Meiofauna can particularly be used as a biological indicator of anthropogenic impacts due to their small size, simple body structure, high abundance, rapid generation time, and direct benthic development (Kennedy and Jacoby, 1999; Zeppilli et al., 2015). These characteristics make these organisms more suited to be used as bioindicators on sandy beaches when compared to traditionally studied organisms, such as the macro- and megafauna (Kennedy and Jacoby, 1999). In addition, these organisms provide different responses to different types of impacts, both qualitatively and quantitatively, (e.g. density and richness changes, replacement of organisms) (Giere, 2009; Moreno et al., 2011); therefore, it can be used as a sensitive indicator of several impacts (Coull and Chandler, 1992; Sun et al., 2014).
However, most studies on the effects of recreational activities in sandy beaches refer to macrofauna (both at the community and population levels) (Schlacher et al., 2007; Schlacher and Thompson, 2012; Costa et al., 2020), and little is known about the effects of these activities on meiofauna community in these environments (Johnson et al., 2007). The results of the few studies available showed a general negative effect of recreational activities on abundance, diversity, and composition of the meiobenthic community (e.g. Moellmann and Corbisier, 2003; Gheskiere et al., 2005; Pereira et al., 2017). Also, recreational activities are known to have a negative effect on beach communities as they may increase sediment compactness (Ugolini et al., 2008). The degree of sediment compaction can be affected by the intensity of recreational activities (Schlacher et al., 2007), and it correlates with several sediment physical properties (Hsu et al., 2009). Thus, sites with high compaction, reflecting harder substrates, might be unfavorable to a wide range of organisms and meiofauna is particularly susceptible, as these organisms inhabit the sediment interstitial space (Giere, 2009).
Macrotidal sandy beaches of the Brazilian Amazonian coast, with their natural environments distributed along a coastline of 3900 km (Klein and Short, 2016), have considerable potential for tourism (Pereira et al., 2016a, b). In general, these beaches are overcrowded during vacation (July) and some bank holidays, especially in the second half of the year (Sousa-Felix et al., 2017). Information available on the biodiversity of Brazilian sandy beaches, particularly in northern Brazil (Amazonian coast), is insufficient to ensure their preservation (Amaral et al., 2016). Furthermore, although meiofauna is a powerful tool to assess human-induced impacts (Schratzberger et al., 2009; Netto and Valgas, 2010), studies that aim to evaluate the effect of recreational activities on meiofauna community in macrotidal tropical sandy beaches are scarce (e.g. Martínez et al., 2020).
In this context, the present study assessed the impact of recreational activities on the structure and composition of meiofauna community and nematode assemblages in three Amazonian macrotidal sandy beaches with different levels of tourism (high, intermediary, and low) before, during, and after an episode of high tourist occupancy (vacation in July). The following hypothesis was tested: high intensity of recreational activities (human trampling and vehicle traffic) causes changes in the structure and composition of the meiobenthic community and nematode assemblages by decreasing species richness and density, particularly in the urban beach during vacation.
Section snippets
Study area
The study area is located in the Atlantic Coastal Sector of North Brazil, one of the most densely populated areas of the Amazonian region (Sousa et al., 2011) with about 40.000 residents, where economy is based on fishing and tourism, which may receive > 280.000 beachgoers during July (IBGE, 2018). Local climate is equatorial humid with annual mean temperature and rainfall of 26–27 °C and 2500–3000 mm, respectively, and approximately 75–85% of rainfall during rainy season (January–May) (
Environmental parameters and human beach use
All environmental parameters are shown in supplementary material 1. In general, the number of beach users (vehicles and beachgoers) was higher during Vacation in all beaches compared to the other periods (beach × month interaction). Both Atalaia and Farol-Velho showed significant differences compared to Corvinas during Vacation (F(1.84) = 6.38; p < 0.05) with higher values of beach users on both beaches compared to Corvinas; however, differences were not detected between Corvinas and
Discussion
Meiofauna density and richness had different patterns among periods. Overall, the Atalaia and Farol-Velho followed the same temporal pattern: sharp reduction in density and richness, with significant changes in community structure between Before Vacation and Vacation months, followed by an increase and restoration of community composition (density and richness) in the After-vacation months. Conversely, the Corvinas beach showed more stability throughout the months, with no significant changes
Conclusion
The results reported in this study showed a similar pattern to those found in previous studies evaluating the effect of recreational activities on sandy beaches as well as in other coastal environments. Thus, the initial hypothesis that recreational activities trigger changes in the structure and composition of meiofauna and nematodes, reducing the richness and abundance of the community, was confirmed. Furthermore, the vulnerability of some taxa studied here, particularly tardigrades and
CRediT authorship contribution statement
Thuareag Monteiro Trindade dos Santos: Conceptualization, Methodology, Validation, Formal analysis, Investigation, Data curation, Writing - original draft. Marcelo Petracco: Conceptualization, Methodology, Resources, Writing - review & editing. Virág Venekey: Conceptualization, Methodology, Resources, Writing - review & editing, Supervision, Funding acquisition.
Funding
This research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors.
Declaration of competing interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
The authors declare that they have no conflicts of interest.
Acknowledgements
This study was funded in part by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - Brasil (CAPES) - Finance Code 001. The authors are grateful to Adrielle Lopes, Afonso Quaresma, Ana Paula Danin, Diego Garcia, Felipe Souza, Gabriel Soares, Keuli Campelo, Leonardo Morais and Roseanne Figueira for their assistance in the field. The first author thanks CAPES for the postgraduate research studentship (Brazil). Thanks also to the anonymous reviewers for their comments, which helped
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