Using the Alternative Stable States Framework to understand the persistence of secondary shrubland within a Mexican oak forest with livestock and agricultural legacies

Many forests exist in human-modified mosaic landscapes subject to logging, agriculture, and livestock grazing. These disturbances can reduce ecosystem resilience and ultimately result in crossing thresholds, leading to the transition of the ecosystem into new stable states with different plant communities. The interaction between these disturbances and forest succession is still poorly understood, despite the growing need to manage these human-modified landscapes for biodiversity and productivity. We studied abrupt changes in vegetation composition in a Central Mexican oak forest and explored how these changes could indicate the coexistence of various alternative stable states caused by agricultural and livestock legacies. We established 36 strip transects covering 9000 m2 in patches with various land use legacies where we sampled vegetation, soil characteristics, and human disturbance. We assessed future and past tendencies in vegetation spatial patterns by evaluating seedling composition and analyzed time series of NDVI signals recorded by LANDSAT images (1972–2019). We utilized NMDS, PERMANOVA, and ANOVA analyses to assess if stable states can be distinguished based on historical and current variations in vegetation composition and structure in patches with different management legacies. We then assess potential factors that may contribute to differences in plant community composition by fitting edaphic, topographic, and anthropogenic disturbance variables to the ordination analyses. We identified three distinct plant communities: oak forest (dominated by late-successional Oak species), secondary shrubland (dominated by Vachellia-Dodonaea association), and a transitional forest-shrub community (a mixture of the first two communities). Time-series analyses and seedling composition provide evidence of long-term community segregation. Variation partition analysis demonstrated that anthropogenic disturbance (particularly livestock browsing), had a greater impact on species composition differentiation than environmental variables. A state-and-transition model is proposed, where cattle browsing and agricultural legacies trigger the transition from oak forests to a stable secondary shrubland community. Three potential self-reinforcing and interacting feedbacks are suggested: (i) micro-environmental filtering, (ii) plant-trait mediated assembly process, and (iii) cascading effects on tree recruitment caused by cattle browsing. The proposed state-and-transition