Forest cover and heterogeneous pastures shape the diversity of predatory rove beetles in tropical riparian habitats
Introduction
In human–dominated landscapes, riparian vegetation acts as a reservoir for many terrestrial taxa and often differs in species composition from neighboring forests or other land use types, increasing regional diversity (Naiman, Decamps & Pollock, 1993; Sabo et al., 2005). At the landscape scale, vegetation associated with watercourses provides habitat and resources for feeding and reproduction in a large number of species (Fahrig et al., 2011; Ries, Fletcher, Battin & Sisk, 2004). Species diversity associated with riparian zones is therefore expected to be influenced both by habitat structural characteristics (e.g., canopy and leaf litter cover) and the surrounding landscape (e.g., land cover types, forest edge density) (Braun, Pires, Stenert, Maltchik & Kotzian, 2018; Marczak et al., 2010).
Riparian zones are complex ecosystems that link water and land environments and provide multiple ecological services such as habitat connectivity and maintenance of biodiversity, riverbank reinforcement, pollution mitigation, nutrient redistribution and delivery and amenity and recreation, all of which are of great value to basin functioning and human well-being (Naiman et al., 1993, Richardson et al., 2007). For these reason, riparian vegetation remnants are an important ecosystem for conservation planning in order to promote structural and functional landscape connectivity to guarantee both the persistence of species and integrity of ecological functions (Rudnick et al., 2012).
The presence and spatial configuration of native vegetation and other land uses such as crops and pastures are among the most important factors that influence species diversity in agricultural landscapes (Fahrig et al., 2011). Intense, but localized, human activity increases landscape heterogeneity, promoting higher species diversity at the local and regional level (Tscharntke et al., 2008). In contrast, landscape homogenization due to the dominance of certain land uses (e.g., treeless pastures) often brings negative impacts on species diversity (Fahrig et al., 2011; Watling, Nowakowski, Donnelly & Orrock, 2011). However, to understand the contribution made by habitat remnants such as riparian vegetation to biodiversity conservation, it is necessary to evaluate their interaction with the surrounding landscape characteristics, which requires studies that consider different spatio-temporal scales (Tscharntke et al., 2012).
Rove beetles (Staphylinidae) make up an important but little-known element of biodiversity in most terrestrial ecosystems, with approximately 65 thousand species described (Irmler & Lipkow, 2018). Necrophilous rove beetles are generalist predators associated with carrion, and some species are associated with specific habitat conditions (Dekeirsschieter, Frederick, Verheggen, Drugmand & Haubruge, 2013; Mądra, Konwerski & Matuszewski, 2014). Rove beetles represent a significant proportion of the predator insect fauna in riparian vegetation (Gutiérrez-Chacon, Zuñiga, Van Bodegom, Chara & Giraldo, 2009). In these environments, Staphylinidae predators have a diet comprised mainly of aquatic invertebrates (some species up 80%) and are therefore considered an important functional component of riverine systems by contributing to the transfer of biomass from aquatic systems to terrestrial food webs (Eyre, Lott & Luff, 2001; Paetzold, Schubert & Tockner, 2005, 2008). The diversity patterns of rove beetles have mainly been explained by microenvironmental conditions and habitat structure (Caballero, León-Cortés & Morón-Ríos, 2009; Irmler & Gürlich, 2007; Irmler & Lipkow, 2018), and a few studies have focused on evaluating the role played by the surrounding landscape supporting the diversity of this insect group (Dauber et al., 2005; Rusch, Birkhofer, Bommarco, Smith & Ekbom, 2014).
In this study, we examined how the diversity and species composition of predatory rove beetles vary in a tropical montane cloud forest (TMCF) landscape in central Veracruz, Mexico, with respect to: i) forest cover at the micro-basin scale (micro-basins dominated by forest and micro-basins dominated by pastures), ii) habitat condition (sites with native riparian vegetation and sites with heterogeneous pastures) and iii) habitat condition as a function of micro basin type. In addition, we determined which site and microsite environmental variables explained species composition. We tested the hypothesis that modification of the environmental conditions (e.g., microclimate and availability of microhabitats), due to the replacement of native riparian vegetation with cattle pastures, will reduce rove beetle diversity, and we expect this reduction to become more evident as the area of TMCF decreases at the micro-basin scale.
Section snippets
Study area
The upper basin of the La Antigua River is located on the eastern slope of the mountain Cofre de Perote in the central region of the state of Veracruz, in Mexico. The rivers and its tributaries drain in a southeasterly direction toward the Gulf of Mexico over an elevation range from 600 to 4200 m a.s.l. The climate is humid (temperatures vary from 10 to 22 °C and annual precipitation is between 600 and 2000 mm; Muñoz‐Villers & López‐Blanco, 2008). The basin has been recognized as a priority
Diversity patterns
A total of 1917 Staphylininae rove beetles were captured, representing 51 species from 19 genera (Appendix A: Table 2 in Supplementary material). Sampling coverage was greater than 97% for micro-basin type and close to 98% for habitat condition. For habitat, depending on the micro-basin type, the sample coverage reaches 95% (Appendix A: Table 2 in Supplementary material). The highest numbers of beetles were caught in the micro-basin with greater forest cover (15% more individuals compared to
Species diversity
Contrary to our hypothesis, all diversity measures were similar between micro-basin types and, although total species richness (0D) was similar between sites with riparian vegetation and heterogeneous pastures, when common (1D) and dominant (2D) species were considered, the rove beetle diversity increased two–fold in both habitat conditions regardless of the micro-basin type. It was also notable, and contrary to expectation, that pastures tended to be more diverse and contain a different set of
Conclusion
Our study supports the notion that heterogeneity within and among human land use contributes to buffering the impact of forest loss at landscape scale. Pastures with isolated and scattered trees and shrubs promote microhabitat availability along rivers and streams, which could function as a semi-natural cover that, combined with different riparian vegetation cover at the micro-basin scale, supports a complementary and diverse Staphylininae predatory rove beetle assemblage. The replacement of
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.
Acknowledgments
We thank José Luis Navarrete-Heredia for his generous help in rove beetle identification. Rosario Landgrave provided the maps of the study area. D. Méndez-Rojas acknowledges the scholarship from the Consejo Nacional de Ciencia y Tecnología (CONACyT No. 300335). Funding was provided by CONACyT (projects CB 2008-101542-F and CB-285962).
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