Bryophyte Diversity and Community Composition in Cacao Agroforestry in Relation to Distance from Native Forest in Brazil

Understanding how landscape and local factors influence biodiversity in agroforestry systems is essential for conservation in fragmented tropical landscapes. We investigated the relative importance of forest proximity and local environmental variables (canopy openness, temperature, and air humidity) in shaping bryophyte assemblages in shaded cacao agroforests in the Brazilian Atlantic Forest. Bryophytes were sampled on 218 trees across six agroforestry sites, totaling 1304 occurrences and 87 taxa. Generalized linear mixed models and multivariate analyses were used to assess patterns of species richness, occurrence frequency, and composition. Moss richness and occurrence frequency increased with forest distance, whereas liverworts showed no clear relationship. Liverwort richness was positively associated with temperature, while occurrence frequency was positively associated with temperature and negatively with air humidity. When accounting for spatial nesting via site-level block permutations, neither forest distance nor microclimate significantly structured species composition (p ≥ 0.313), reflecting site-level spatial heterogeneity. Bryoflora was dominated by generalist taxa and species with intermediate desiccation tolerance, and assemblages showed taxon-specific relationships with local environmental conditions. Our findings suggest the hypothesis that long-established, shaded cacao agroforests may support diverse bryophyte assemblages in fragmented tropical landscapes, a pattern that requires formal evaluation with native forest reference controls.