Landscape changes, especially habitat loss and degradation, are particularly significant in Brazil's major vegetation domains, which are characterized by unique topography, diverse aquatic systems, and high levels of endemism (Drummond et al., 2005; Sousa et al., 2010). Among the six major Brazilian vegetation domains, the Tropical Savanna (Cerrado), Tropical Dry Forests (Caatinga), and Atlantic Forests stand out for their ecological richness. The Tropical Savanna is a heterogeneous domain, that supports a wide variety of species (Sano et al., 2019), while the Atlantic Forest is globally recognized as one of the most biodiverse ecosystems on the Earth (Faria and Kaizer, 2020; Solórzano et al., 2021). Both the Tropical Savanna and Atlantic Forest are classified as global biodiversity hotspots (Mittermeier et al., 2011). Similarly, the Tropical Dry Forests in Brazil also exhibit high levels of endemism (Silva et al., 2014; Caetano et al., 2022). Despite their ecological importance, these domains continue to undergo significant degradation driven by human activities (Dutra et al., 2012; Girardi, 2014; Lopes et al., 2020).

Human activities, particularly those related to the uses and occupations of lands, as well as the removal of riparian forests, are major drivers of landscape transformations, with cascading effects on aquatic ecosystems and biodiversity losses (Valiente-Banuet et al., 2015; Mello et al., 2020). They directly and indirectly impact aquatic systems by modifying river channels, increasing sedimentation, and introducing pollutants (Walks, 2008; Ding et al., 2016; Singh et al., 2021). These changes disrupt species dispersal, reduce habitat connectivity, and convert native ecosystems into agricultural lands, ultimately decreasing regional dissimilarity and promoting landscape homogenization (Morante-Filho et al., 2015; Coelho et al., 2018; Cardoso et al., 2020; Ramos et al., 2022).

Among aquatic biodiversity, zooplankton, a community of invertebrates, including rotifers, cladocerans, and copepods (Elmoor-Loureiro et al., 2023), are highly sensitive to environmental changes and are excellent indicators of ecosystem health due to their short life cycles and high dispersal capacities (Castilho-Noll et al., 2023). Consequently, zooplankton are valuable for assessing the impacts of disturbances on natural ecosystems. Understanding the spatial distribution of zooplankton species contributes to understanding how natural ecological processes (Heino et al., 2015a) and anthropogenic stressors (Simões et al., 2022) filter the regional species pool. Changes in species composition and biodiversity serve as critical tools for identifying environmental modifications and assessing ecological quality (Chase et al., 2020).

Metrics such as beta diversity can reveal environmental heterogeneity and spatial variation, helping to measure how habitat transformation affects communities (Bomfim et al., 2024a). However, assessing human impacts on landscapes remains challenging due to varying degrees of disturbance. The Anthropic Transformation Index (ATI), developed by Lemeshev (1982) and applied by Costa et al. (2014); Ribeiro et al. (2017); Vieira et al. (2021) and others, helps quantify human pressure and the extent of landscape changes, which makes it possible to diagnose the environmental quality of occupied environments (Gouveia et al., 2013; Silva et al., 2021).

Given that aquatic environments are closely linked with terrestrial ecosystems, analyzing land use changes in riparian zones is crucial for understanding how terrestrial transformations influence aquatic biodiversity (Meier et al., 2015; Usio et al., 2017; Mwaijengo et al., 2020). But the relationship between spatial variation of land uses and zooplankton diversity remains underexplored, especially in neotropics (dos Santos et al., 2025; Balseiro et al., 2023; Bomfim et al., 2024b). Therefore, the present study investigated the relationships between landscape variations in regions close to aquatic environments and the diversity (alpha and beta) of zooplankton organisms in zones of native vegetation and zones influenced by human activities in the Atlantic Forest, Tropical Savanna, and Tropical Dry Forests.

To advance the understanding of biodiversity variation patterns (Gaston, 2000), it is essential to consider that freshwater systems provide valuable opportunities to identify general drivers associated with the properties of ecological communities (Heino et al., 2015b). Among these drivers, land-use change plays a critical role, as it directly modifies habitat and landscape characteristics, thereby influencing environmental filtering processes that shape community composition and diversity (Chase et al., 2020). Based on this framework, we hypothesize that zooplankton biodiversity responds to landscape variation along environmental gradients. Specifically, we predict that (i) alpha diversity is greater in landscapes with lower environmental stress near aquatic‒terrestrial interfaces; (ii) the landscape structures of the Tropical Savanna and Tropical Dry Forests are more similar to each other than to those of the Atlantic Forest; and iii) beta diversity increases with landscape heterogeneity as an indirect response to anthropogenic influences on zooplankton community composition.

The observed species richness (Magurran, 2003), individual density, Shannon diversity index, and evenness (Krebs, 1998) were used as measures of alpha diversity. To demonstrate the richness of unique and shared species in each vegetation domain, a Venn diagram was constructed using the online software Bioinformatic and Evolutionary Genomic, which is available at: https://bioinformatics.psb.ugent.be/webtools/Venn/.

To verify the spatial differences in the composition of the zooplankton community along the sampling sites between the vegetation domains, a permutational multivariate analysis of variance (PERMANOVA) (Anderson, 2001) was performed, followed by a principal coordinate analysis (PCoA) to visualize the variation in the composition of the zooplankton community between the sampling sites.

Beta diversity was calculated by permutational analysis of multivariate dispersion (PERMDISP) (Anderson et al., 2006), and the relationships with the vegetation domains were subsequently tested via analysis of variance (ANOVA). The beta diversity index was represented by the average distance to the centroid of each group calculated by PERMDISP. Pearson's correlation was used to assess the relationships between community attributes and ATI and land use categories (native areas × anthropic areas) and zooplankton beta diversity. ANOVAs were calculated to evaluate the differences in landscape indices (ATI and environmental heterogeneity) between the domains.

Finally, a BIOENV procedure was performed to identify the best combination of land use categories associated with the zooplankton community (Clarke and Ainsworth, 1993). The best combination was tested via the Mantel test (999 permutations).

All analyses were performed in R.4.1 (R Core Team, 2021), and the "vegan" and "BiodiversityR" packages and the "ggplot2" package were used to create the graphs. The significance for all analyses was set at p < 0.05.

The Atlantic Forest presented the highest alpha diversity and percentage of exclusive species and lowest ATI index, corroborating our first prediction. The high percentage of native areas within this vegetation domain, consisting of remaining forest patches, contributed to these findings, where it is expected to be found a wider range of microhabitats and environmental conditions capable of sustaining a more diverse array of species (Molina et al., 2017; Capellesso et al., 2022; Ribeiro et al., 2025). In contrast, the Tropical Savanna and Tropical Dry Forest, characterized by higher levels of habitat alteration, supported fewer taxa overall and showed a higher degree of community overlap. These findings are consistent with previous studies (e.g., Cortez-Silva et al., 2020; Bomfim et al., 2023; Santos et al., 2025) that highlight the influence of both local and regional factors on freshwater zooplankton biodiversity. While local attributes such as habitat quality (Gutierrez et al., 2022; das Candeias et al., 2022) and availability of refuges (Deosti et al., 2021) are key drivers of alpha diversity, regional elements, such as similarity in vegetation physiognomy (Dodds et al., 2019) and landscape connectivity (Pedruski and Arnott, 2011), appear to shape community composition across broader scales. Although zooplankton species partially overlap among the areas, they respond differently to environmental filters specific to each domain (Cabral et al., 2020). These differences arise because environmental filters vary across regions, influencing species selection and shaping their responses to environmental conditions (Agra et al., 2021; Diniz et al., 2021; Astorga et al., 2014).

The landscape composition surrounding the sampled aquatic systems varied markedly across the three vegetation domains. The Atlantic Forest exhibited the highest proportion of natural areas (73.9%) and the lowest ATI values, indicating relatively preserved conditions in its buffer zones. In contrast, the Tropical Dry Forest showed the greatest anthropogenic pressure, with only 47.8% of natural cover and the highest ATI values. These statistically significant differences indicate varying degrees of landscape transformation, likely driven by region-specific agricultural expansion and historical land-use policies, in a process known as frontier theory (Schielein and Börner, 2018), which is often associated with negative impacts on the both terrestrial (Foley et al., 2005; Phillips et al., 2017) and aquatic biodiversity (Dudgeon et al., 2006; Miserendino et al., 2011; Augusto et al., 2025).

The beta diversity increased in response to anthropogenic impacts, particularly in the Tropical Savanna and Tropical Dry Forest, which experienced the highest degrees of human transformation in the aquatic-terrestrial interface zones. This positive relationship indicates that an increase in the range of environmental gradients drives changes in the zooplankton community, revealing the deterministic influence of habitat heterogeneity on beta diversity (Stegen et al., 2013; Socolar et al., 2016). This also helps explain our third prediction, as anthropogenic transformations in the Tropical Dry Forest and Tropical Savanna induce eutrophication and other stressors, directly influencing zooplankton community composition (Diniz et al., 2021). For instance, the highest ATI values at Tropical Dry Forest sites reflect significant anthropogenic pressure, where livestock expansion is a main economic activity (Alves et al., 2009). Livestock and wood extraction are key drivers of spatial heterogeneity across the Tropical Dry Forest (Antongiovanni et al., 2018). These activities, combined with natural vulnerabilities like river intermittency, pose significant threats to biological diversity and primary productivity in these domains.

Thus, the land-use mosaics (including forest formation, agriculture, pasture, and cotton) surrounding aquatic environments promoted species turnover. This aligns with previous studies (Maloufi et al., 2016; Bomfim et al., 2024a), indicating that environmental changes in the landscape act as filters, selecting a local species from the regional pool. In other words, variations in local environmental conditions drive a high replacement of species (Braghin et al., 2015), likely because the environmental characteristics imposed by land use stress aquatic environments, limiting species establishment (Bomfim et al., 2024a; Duré et al., 2025). Moreover, forest cover affects zooplankton functional guilds by altering local environmental variables related to feeding strategies (Bomfim et al., 2023). Biodiversity is rarely governed by a single mechanism, especially when disturbance sources and intensities vary across regions (Hawkins et al., 2015; Socolar et al., 2016; Simões et al., 2022).

The combined analysis of landscape metrics (ATI, anthropogenic areas and environment heterogeneity) highlights the value of multiple approaches to better understand biodiversity variations. The different ways of measuring environmental variation on the basis of landscape metrics used in this study helped to understand the spatial distribution of zooplankton diversity, with an emphasis on ATI, which is driven by greater weights in areas with human activities (Rodriguez et al., 2017). The analysis that grouped land uses into native and anthropic areas also helped to reveal the gradients of change in species composition. Landscape heterogeneity through the standardization of categories captured the relationship of species composition variation only at the regional level. These discussions are pivotal for identifying the relationship between landscape environmental heterogeneity and aquatic biodiversity because the choice of spatial land-use metrics can bias conclusions of land-use impacts in river systems (Mwaijengo et al., 2020), and the sign and strength of correlations between environmental heterogeneity and biodiversity depend on the level of anthropogenic disturbance (Agra et al., 2021).

This study contributes to understanding the role of landscape structure in aquatic microbiodiversity distribution. Expanding the use of landscape metrics as indicators of ecosystem disturbance is crucial for increasing the explained variation in the plankton-environment relationship (Meier et al., 2015) and supporting decision-making about water resource management (Ding et al., 2016).

Land-use composition and anthropogenic pressure significantly influenced zooplankton community structure in lotic aquatic systems across different vegetation domains. While species richness and diversity were highest in more preserved landscapes, such as the Atlantic Forest, beta diversity was more responsive to land-use heterogeneity and degradation, particularly in the Tropical Savanna. Our findings confirm that landscape heterogeneity alters alpha and beta diversity patterns and that the use of multiple metrics is key to understanding the effects of landscape changes on aquatic biodiversity. The association between specific land-use categories and beta diversity highlights the importance of landscape-scale processes in shaping aquatic biodiversity.

We thank the FAPESB (Fundação de Amparo a Pesquisa do Estado da Bahia)for supporting the project. NRS thanks Nacional de Desenvolvimento Científico e Tecnológicofor their grants (Process: 308074/2022-0).