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. 2025 Jun 16;20(6):e0326304.
doi: 10.1371/journal.pone.0326304. eCollection 2025.

Landscape to microhabitat: Uncovering the multiscale complexity of native and exotic forests on Terceira Island (Azores, Portugal)

Sébastien Lhoumeau  1 , Rui B Elias  1 , Dominik Seidel  2 , Rosalina Gabriel  1 , Paulo A V Borges  1   3   4
Affiliations

Landscape to microhabitat: Uncovering the multiscale complexity of native and exotic forests on Terceira Island (Azores, Portugal)

Sébastien Lhoumeau et al. PLoS One. .

Abstract

This study aims to identify the structural and compositional differences between native and exotic woodlands on Terceira Island, Azores. Based on landscape, habitat, and microhabitat analyses, remnants of native forests appeared to be associated with less accessible terrains. A more homogeneous structural complexity is exhibited, derived from the numerous branching patterns of the endemic vascular plant species. In contrast, exotic forests exhibit structural heterogeneity driven by mixed non-indigenous vascular plant species as a result of human actions such as afforestation and latter invasion of exotic tree species, after abandonment of the agricultural use. The ground and canopy layers in exotic forests were more invaded by non-indigenous species, while the understory demonstrated greater resilience by being mostly composed of indigenous species. Our findings highlight the structural and ecological differences between native and exotic woodlands, reflecting the historical transformation of forest cover in the Azores. These insights emphasize the importance of long-term monitoring and structural assessments in informing conservation efforts aimed at preserving native forests and managing invasive species in exotic woodlands.

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Conflict of interest statement

The authors have declared that no competing interests exist.

Figures

Fig 1
Fig 1. Location of sampling sites.
Map of the study area on Terceira Island, Azores, showing the location of the 18 forest sites surveyed, including nine within native forest and nine within exotic woodlands. Protected areas are indicated based on data from [50], and land use classifications are provided by the Azorean Government.
Fig 2
Fig 2. Visual representation of the group of variables considered.
The organisation is based on the spatial extent from landscape scale (Spatial traits) to plot scale (stand traits) micro-habitat (canopy, understory and ground traits).
Fig 3
Fig 3. NMDS Plot of Native vs. Exotic Forests on Terceira Island.
Non-metric Multidimensional Scaling (NMDS) plot showing the separation between native and exotic forest plots on Terceira Island, Azores, based on 49 variables. Each point represents an individual plot, with ellipses indicating the forest types (native (green) and exotic (yellow)). The clear spatial separation between the two ellipses reflects distinct ecological characteristics of native versus exotic forest structures, as supported by a significant ANOSIM result (R = 0.9036, p < 0.001).
Fig 4
Fig 4. NMDS Plots of Traits by Forest Type.
Non-metric Multidimensional Scaling (NMDS) plots for each trait considered. Each point represents an individual plot, with ellipses indicating the forest types (native (green) and exotic (yellow)). ANOSIM statistics and significance are mentioned for each graph.
Fig 5
Fig 5. Visual comparison of native and exotic woodlands.
Comparison of forest structures between native forest (left, TER-NFSB-T164B) and exotic woodlands (right, TER-PRIBS-T06) on Terceira Island. Visually, the native forest displays a complex vertical structure with diverse strata and species, while the exotic forest exhibits a reduced species diversity and overall structural complexity.
See this image and copyright information in PMC

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