## 2 Materials and Methods  
### Study Area  
Payne's Creek National Park (PCNP) in the Toledo District of southern Belize covers approximately 152 km\({}^{2}\) of low elevation land that borders the Caribbean Sea (Figure 1). The park is located at approximately 16.3\({}^{\circ}\) N 88.6\({}^{\circ}\) W in Belize's southern coastal plain. Belize has a subtropical tropical climate with mean monthly minimum/maximum temperatures from 16/28 \({}^{\circ}\)C in winter to 24/33 \({}^{\circ}\)C in summer [28]. The Koeppen climate classification of most of Belize, including PCNP, is \"tropical monsoon\" [29], which is characterized by a distinct wet and dry season. The majority of rainfall falls from June to November in Belize [29]. For this study, the dry season is considered as January-May and the wet season is June-December. The dry season average precipitation in PCNP is approximately 498 mm and 2208 mm in the wet season. The minimum/maximum precipitation in PCNP in the dry season was 269 mm/790 mm (1987/1990) and 1647 mm/2903 mm (2004/1979) in the wet season [30; 31; 32]. Mean annual rainfall ranges from 1524 mm in the north and 4064 mm in the south. This variability occurs due to orographic lifting over steep slopes and the seasonal migration of the Inter-Tropical Convergence Zone [29]. Belize contains upland savannas known as the Mountain Pine Ridge and lowland savannas of the northern and southern coastal plains. This study focuses on Belize's lowland savannas. The lowland savannas of Belize are located between dense broadleaf rainforests inland and mangrove swamps seaward in the relatively flat, poorly drained coastal plain [33]. Vegetation in these savannas are a mosaic of grassland and pine or oak trees, and pockets of broadleaf forest [23; 28; 34]. Both upland and lowland savannas are in areas with weathered, acidic and nutrient-poor soils, but the upland savannas are well-drained while those in the lowlands frequently experience flooding and drying with the wet and dry seasons, respectively [35]. The lowland area is underlain by the Yucatan limestone platform, which was shallow sea during the Pliocene (2-13 Myr BP). Subsurface clay sits atop the limestone platform, resulting in widespread poor drainage in the lowlands. Sands and gravels from the nearby Maya Mountains eroded and accumulated on the clay and formed the current lowland coastal plain [35]. Savannas in the low-lying area usually occur on nutrient-poor soils underlain by the sands and gravels [28]. Undulations in the lowland topography result in improved drainage and the dominance of woody vegetation, such as pine and oak. Dense broadleaf forests occur along rivers in the area due to increased nutrients in alluvial deposits originating in the Maya Mountains [28]. At an even finer scale, disturbance feedbacks in the form of fire and hurricanes control the boundaries between grasses and trees. This is partly because the dominant pine species, _Pinus caribaea_, most readily regenerates on sunlit mineral soils. These soils become available after fire from the removal of grass cover and organic matter, or because old trees are killed and stand density is reduced [36]. Similarly, strong winds from hurricanes can reduce stand density and allow sunlight to penetrate to the surface [36]. The Toledo Institute for Development and Environment (TIDE) manages PCNP and uses fire management practices in the fire-adapted tropical pine savanna and grassland ecosystems. While the pine savanna is fire adapted, too many fires would eventually lead to a grassland without pine by scorching saplings attempting to regenerate. Too few fires results in a thick herbaceous layer  
Figure 1: Map of Payne’s Creek National Park (PCNP), showing nearby countries in Central America. Image of PCNP is a true color composite from Landsat 5 TM taken in March 2011. Shapefile of Central America from Natural Earth. Shapefile of Belize and its rivers from Biodiversity and Environmental Resource Data System of Belize. Shapefile of Payne’s Creek National Park from Protected Planet.