Seed rain along a gradient of degradation in Caribbean dry forest: Effects of dispersal limitation on the trajectory of forest recovery

Questions Tropical dry forests that experience severe disturbances (e.g., fires) often remain degraded for long time periods, during which non-native grasses and trees dominate. One barrier to native tree regeneration in degraded areas may be seed dispersal limitation. To better understand how dispersal limitation influences recovery from degradation, we tested whether the mode and rates of seed dispersal differed in degraded sites dominated either by the exotic tree Leucaena leucocephala or open areas dominated by introduced pasture grasses. We also tested whether L. leucocephala stands facilitate the recruitment of native trees by increasing their seed input compared to open grass areas. Location Guánica Commonwealth Forest, Puerto Rico. Methods Seed rain was measured for one year in traps located within five vegetation types that ranged in degree of forest degradation from open grass to intact native forest. Results In open grass areas, seed rain density was similarly low for L. leucocephala and abiotically dispersed native trees (mean [95% CI] = 50.9 [15.1–171.0] vs. 34.2 [10.3–113.5] seeds m−2 year−1), whereas it was even lower for animal-dispersed native trees (0.14 [0.03–0.67] seeds m−2 year−1). Compared to open grass areas, L. leucocephala-dominated stands, even those with grass understories, had higher seed rain density of animal-dispersed trees (43.0 [12.9–143.6] seeds m−2 year−1), but not abiotically dispersed trees (20.8 [6.3–68.5] seeds m−2 year−1). Conclusions The dominance of L. leucocephala in disturbed Caribbean dry forests does not appear to be mediated by disproportionate seed arrival in open areas compared to native tree seeds. Rather, subsequent factors such as seed and seedling survival likely favor L. leucocephala in highly degraded areas. Since L. leucocephala stands increase the seed rain of animal-dispersed native trees, retaining them in highly disturbed Caribbean dry forests may facilitate the regeneration of native forests

Regional Forest Types - Tropical Dry Forests

Tropical dry forests occur in nearly every tropical country. This forest type provides critical habitat for large mammals and migratory birds, and patches of dry forest can support a high proportion of endemic plant and animal species, as well as being highly valued for agricultural and production forestry uses. Consequently, conservation and understanding of these forests need emphasis, yet conservationists and scientists still frequently overlook this ecosystem

Structural response of Caribbean dry forests to hurricane winds: a case study from Guanica Forest, Puerto Rico

Aim Tropical dry forests in the Caribbean have an uniquely short, shrubby structure with a high proportion of multiple-stemmed trees compared to dry forests elsewhere in the Neotropics. Previous studies have shown that this structure can arise without the loss of main stems from cutting, grazing, or other human intervention. The Caribbean has a high frequency of hurricanes, so wind may also influence forest stature. Furthermore, these forests also tend to grow on soils with low amounts of available phosphorus, which may also influence structure. The objective of this study was to assess the role of high winds in structuring dry forest, and to determine whether soil nutrient pools influence forest response following hurricane disturbance. Location Guánica Forest, Puerto Rico. Methods Over 2000 stems in five plots were sampled for hurricane effects within 1 week after Hurricane Georges impacted field sites in 1998. Sprout initiation, growth, and mortality were analysed for 1407 stems for 2 years after the hurricane. Soil nutrient pools were measured at the base of 456 stems to assess association between nutrients and sprout dynamics. Results Direct effects of the hurricane were minimal, with stem mortality at \u3c 2% and structural damage to stems at 13%, although damage was biased toward stems of larger diameter. Sprouting response was high – over 10 times as many trees had sprouts after the hurricane as before. The number of sprouts on a stem also increased significantly. Sprouting was common on stems that only suffered defoliation or had no visible effects from the hurricane. Sprout survival after 2 years was also high (\u3e 86%). Soil nutrient pools had little effect on forest response as a whole, but phosphorus supply did influence sprout dynamics on four of the more common tree species. Main conclusions Hurricanes are able to influence Caribbean tropical dry forest structure by reducing average stem diameter and basal area and generating significant sprouting responses. New sprouts, with ongoing survival, will maintain the high frequency of multi-stemmed trees found in this region. Sprouting is not limited to damaged stems, indicating that trees are responding to other aspects of high winds, such as short-term gravitational displacement or sway. Soil nutrients play a secondary role in sprouting dynamics of a subset of species. The short, shrubby forest structure common to the Caribbean can arise naturally as a response to hurricane winds

A link between hurricane-induced tree sprouting, high stem density and short canopy in tropical dry forest

The physiognomy of Caribbean dry forest is shorter, denser and contains a greater proportion of multi-stemmed trees than other neotropical dry forests. Our previous research, conducted after Hurricane Georges in 1998, has shown that dry forest trees sprout near the base following hurricane disturbance, even if the trees have not incurred structural damage. However, for these hurricane-induced sprouts to contribute to the physiognomy of the forest, they must grow and survive. We followed sprout dynamics and stem mortality on 1,407 stems from 1998, after Hurricane Georges, until 2005. The number of surviving sprouts and the proportion of sprouting stems decreased during the 7-year period, but the sprouting rate was still 3-fold higher and the proportion of sprouting stems 5-fold higher than before the hurricane. Mortality of non-sprouting stems (15.4%) was about the same as for sprouting stems (13.9%) after 7 years. The mean length of the dominant sprout surpassed 1.6 m by 2005, with over 13% of the dominant sprouts reaching subcanopy height. Sprout growth and survival varied among species. These results demonstrate that, despite some thinning, hurricane-induced sprouts survive and grow and that the unique physiognomic characteristic of Caribbean dry forests is related to hurricane disturbance

Water requirements for growth and survival of Swietenia macrophylla and Tabebuia heterophylla juvenile trees in relation to water production capacity of dew condensers1

Drought mortality of juvenile trees is a major cause for failure of reforestation projects. Portable devices such as passive radiative dew condensers can often provide 0.15 L/day of water in situ, possibly sufficient for tree survival until roots can access groundwater, allowing self-sustainability. To evaluate growth and survivability of juvenile trees of Tabebuia heterophylla Britton and Swietenia macrophylla King under such low water amounts, juvenile trees received approximately 0.033, 0.067, 0.134, 0.201 and 0.268 L/tree/day, representing fractions (relative evapotranspiration or RET) of 0.125, 0.25, 0.50, 0.75 and 1.00 of the evapotranspiration demand (ETo). The experiment lasted 60 days for S. macrophylla and 90 days for T. heterophylla. All T. heterophylla juvenile trees survived even at the lowest irrigation rate. However, S. macrophylla juvenile trees began dying at RET \u3c 0.5, with only 60 percent surviving at RET = 0.25 (0.067 L/day) and 100 percent mortality occurring at RET = 0.125 (0.033 L/day). Water requirements of 0.134 L/day, necessary for full survival of both species, were within the typical production capacity of 1-m2 dew condensers. However, a greater safetyfactor is obtained using drought tolerant species such as T. heterophylla, which can survive under water application rates as low as 0.03 L/day

Fire resistance in a Caribbean dry forest: inferences from the allometry of bark thickness

Trees’ resistance to fire-induced mortality increases with bark thickness, which varies widely among species and generally increases with stem diameter. Because dry forests are more fire-prone than wetter forests, bark may be thicker in these forests. However, where disturbances such as hurricanes suppress stem diameter, trees may not obtain fire-resistant bark thickness. In two hurricane-prone Caribbean dry-forest types in Puerto Rico—deciduous forest and scrub forest—we measured bark thickness on 472 stems of 25 species to test whether tree species obtain bark thicknesses that confer fire resistance, whether bark is thicker in the fire-prone scrub forest than in the deciduous forest, and how bark thickness in Caribbean dry forest compares with other tropical ecosystems. Only 5% of stems within a deciduous-forest stand had bark thickness that would provide \u3c 50% probability of top-kill during low-intensity fire. In contrast, thicker-barked trees dominated the scrub forest, suggesting that fires influenced it. Compared with trees of similar diameter in other regions of the tropics, bark in Caribbean dry forest was thinner than in savanna, similar to other seasonally dry forests, and thicker than moist-to-wet forests. Dry-forest species appear to invest more in fire-resistance than species from wetter forests. However, Caribbean dry forests remain highly vulnerable to fire because the trees rarely reach large enough diameters to be fire resistant

Spread of common native and invasive grasses and ruderal trees following anthropogenic disturbances in a tropical dry forest

Introduction A fundamental challenge to the integrity of tropical dry forest ecosystems is the invasion of non-native grass species. These grasses compete for resources and fuel anthropogenic wildfires. In 2012, a bulldozer from the Puerto Rico Electric Power Authority cleared a 570-m trail from a state road into a mature dry forest section of Guánica Forest to control a wildfire. We monitored colonization by a non-native invasive grass (Megathyrsus maximus), a highly invasive tree (Leucaena leucocephala), and a native grass (Uniola virgata), as well as natural regeneration, along the bulldozer trail. We determined whether bulldozing facilitated colonization by these species into the forest and the extent of spread. Results Distance from propagule source and temporal variations strongly influenced colonization by our three focal species. Megathyrsus maximus invaded along the trail from source populations by the state road. The establishment of new colonies of M. maximus seedlings went as far as 570 m inside the forest (i.e., at the end of the bulldozer trail), but we found most new colonies within 270 m of the road. Leucaena leucocephala exhibited a similar spreading pattern. Before disturbance, Uniola virgata was distributed widely across the forest, but the highest densities were found in areas near the latter portion (\u3e 401 m) of the bulldozer trail. Subsequently, the species formed new clumps along more than half of the trail (250 to 570 m), apparently colonizing from undisturbed patches nearby. Conclusions Bulldozing facilitated the invasion of non-native vegetation. The projected community assemblage will be more fire-prone than before since M. maximus carries fire across the landscape better than U. virgata, emphasizing the capacity of invasive plant colonization to alter local ecological processes after only a single wildfire and bulldoze event. Our results provide a valuable baseline for short-term vegetation response to anthropogenic disturbances in tropical semi-deciduous dry forests

Shifts in biomass and productivity for a subtropical dry forest in response to simulated elevated hurricane disturbances

Caribbean tropical forests are subject to hurricane disturbances of great variability. In addition to natural storm incongruity, climate change can alter storm formation, duration, frequency, and intensity. This model-based investigation assessed the impacts of multiple storms of different intensities and occurrence frequencies on the long-term dynamics of subtropical dry forests in Puerto Rico. Using the previously validated individual-based gap model ZELIG-TROP, we developed a new hurricane damage routine and parameterized it with site- and species-specific hurricane effects. A baseline case with the reconstructed historical hurricane regime represented the control condition. Ten treatment cases, reflecting plausible shifts in hurricane regimes, manipulated both hurricane return time (i.e. frequency) and hurricane intensity. The treatment-related change in carbon storage and fluxes were reported as changes in aboveground forest biomass (AGB), net primary productivity (NPP), and in the aboveground carbon partitioning components, or annual carbon accumulation (ACA). Increasing the frequency of hurricanes decreased aboveground biomass by between 5% and 39%, and increased NPP between 32% and 50%. Decadal-scale biomass fluctuations were damped relative to the control. In contrast, increasing hurricane intensity did not create a large shift in the long-term average forest structure, NPP, or ACA from that of historical hurricane regimes, but produced large fluctuations in biomass. Decreasing both the hurricane intensity and frequency by 50% produced the highest values of biomass and NPP. For the control scenario and with increased hurricane intensity, ACA was negative, which indicated that the aboveground forest components acted as a carbon source. However, with an increase in the frequency of storms or decreased storms, the total ACA was positive due to shifts in leaf production, annual litterfall, and coarse woody debris inputs, indicating a carbon sink into the forest over the long-term. The carbon loss from each hurricane event, in all scenarios, always recovered over sufficient time. Our results suggest that subtropical dry forests will remain resilient to hurricane disturbance. However carbon stocks will decrease if future climates increase hurricane frequency by 50% or more

Multi-temporal unmixing analysis of Hyperion images over the Guanica Dry Forest

This paper presents a methodology to analyze time-series data from Hyperion to study seasonal vegetation dynamics on the Guánica Dry Forest in Puerto Rico. Unmixing analysis is performed over ten near-cloud-free Hyperion images collected in different months in 2008. Abundance maps and endmembers estimated from the unmixing procedure are used to analyze the seasonal changes in the forest. Results from the analysis are compared with published knowledge of the Guanica Forest phenology

Stem-inhabiting fungal communities differ between intact and snapped trees after hurricane Maria in a Puerto Rican tropical dry forest

Hurricanes impact forests by damaging trees and altering multiple ecosystem functions. As such, predicting which individuals are likely to be most affected has crucial economic importance as well as conservation value. Tree stem-inhabiting fungal communities, notably rot-causing agents, have been mentioned as a potential factor of tree predisposition to hurricane damage, but this assumption remains poorly explored. To examine this relationship, we sampled the stem wood of intact and damaged trees shortly after Hurricane Maria in a Puerto Rican dry tropical forest in 2017. We categorized samples depending on two types: trees with intact stems and trees in which stems were snapped. We extracted fungal environmental DNA of wood from 40 samples consisting of four different tree species. Fungal community taxonomic and functional richness and composition was assessed using high-throughput DNA metabarcoding. We found that snapped trees harbored significantly higher fungal operational taxonomic unit (OTU) richness than the intact trees and that the composition of the stem-inhabiting fungal communities diverged consistently between intact and snapped trees. On average, snapped trees’ fungal communities were relatively enriched in “other saprotrophs” guild category and depleted in endophytes. Conversely, intact trees had high relative abundances of Clonostachys, a mycoparasitic endophyte, suggesting that endophytic fungi might act as biocontrols in tree stems. Overall, our results support the hypothesis that stem-inhabiting fungal communities could represent a predisposition factor of tree damage caused by hurricanes in tropical dry forests