*2.1. Study Area*

The study area was Togo (Figure 1A). It is a coastal country in West Africa that is bordered by Burkina Faso to the north, the Atlantic Ocean to the south, Benin to the east, and Ghana to the west. It belongs to the Sudano-Guinean zone, which is a climatic zone that is located south of the Sahara Desert in the continental and coastal areas, which extend from West Africa to Central Africa. With an area of 56,600 km2, Togo has a population of 7,264,637 inhabitants unequally distributed in the administrative regions with proportions of 42.16% in Maritime, 22.16% in Plateaux, 9.99% in Centrale, 12.44% in Kara and 13.26% in Savanes [29]. It experiences a tropical Sudano-Guinean climate with rainfall ranging from 900 to 1100 mm year<sup>−</sup><sup>1</sup> in the northern regions (distinct wet and dry seasons), and from 1000 to 1600 mm year<sup>−</sup><sup>1</sup> in the southern regions (with four seasons), and an average temperature of 27 ◦C [30].

**Figure 1.** (**A**) Geographical location of the study area; (**B**) ecological zones and elevations.

Due to its position in the Dahomey Gap (a remarkable interruption in the extent of continuous tropical rainforest covering Central to West Africa), Togo has a low forest cover with a deforestation rate of 0.73% per year for the period from 1990 to 2000 [31]. To ensure the protection of the country's forest resources, 14.2% of its territory was classified between 1939 and 1957 as 83 protected areas (classified forests, national parks, and reserves). Yet, human populations seeking arable land and wood for energy have encroached upon nearly one-third of these areas [32]. Vegetation formations are composed of the Sudano-Guinean forest that is located in the mountainous areas of the country, gallery forest along main rivers, dry forest or dense tree savannah in the northern half, and tree savannah in the south and center. The landscape variability of these ecosystems led [33] to subdivision of the country into five ecological zones (Figure 1B).

Ecological zones correspond to distinct ecosystems that are characterized by various plant formations and topographies. Following an update of their descriptions, these ecological zones have been summarized in [34] as follows:


• Zone V (or Southern Coastal Zone): This zone corresponds to the country's coastline with a sub-equatorial climate with two rainy seasons. The very degraded natural environment is strongly dominated by agrosystems, with relic mosaics of savannahs, halophytic or swampy grasslands, and mangroves. The main spontaneous ligneous species found there are *Lonchocarpus sericeus, Parkia biglobosa, Piliostigma thonningii, Dialium guineense, Holarrhena floribunda, Bridelia ferruginea Millettia thonningii,* and *Vitellaria paradoxa*. These natural ecosystems are highly degraded (85%) due to cultivation practices (59%) and the unsustainable exploitation of wood energy (18%) and urbanization (10%). Lands in the Coastal Zone have been heavily disturbed by vegetation fires (55%), which are often followed by extensive grazing (15%), and transhumance, woodcutting, and flooding (5%) [34].

The aforementioned descriptions indicate the continuation of high-intensity land degradation that has been observed across most of these zones since the 1990s [35]. Even in Zone IV, which is known as being the most extensively forested of the ecological zones, deforestation and forest degradation have been occurring in recent years due to the combined effect of the advancing agricultural front with slash-and-burn agriculture, wildfires, and logging [36].
