Modeling the effects of Hurricane Hugo on spatial and temporal variation in primary productivity and soil carbon and nitrogen in the Luquillo Experimental Forest, Puerto Rico

Hurricanes account for much of the spatial and temporal variation in forest productivity and soil organic matter pools in many forest ecosystems. In this study, we used an ecosystem level model, TOPOECO, to simulate the effects of Hurricane Hugo (18 September 1989) on spatial and temporal patterns of gross primary productivity (GPP), net primary productivity (NPP), soil organic carbon (SOC) and nitrogen over the entire Luquillo Experimental Forest (LEF), Puerto Rico, a tropical rainforest. Our simulation results indicated that simulated annual GPP increased by an average of 30% five years after Hugo in the Tabonuco forest at low elevations where there was a fast recovery of the canopy, whereas simulated GPP decreased by an average of 20% in the Palm and Dwarf forests at high elevations as a result of the slow recovery of the canopy. Simulated annual NPP in the Palm and Dwarf forests also did not recover to pre-Hugo levels within 5 years. Simulated storages of SOC, CO₂ emission from decomposition of SOC and total soil nitrogen increased slightly but N mineralization rate increased significantly in all four vegetation types due to the massive input of plant materials from Hugo at low elevations and the slow decomposition at high elevations.     

Lack of Ecotypic Differentiation: Plant Response to Elevation,Population Origin,and Wind in the Luquillo Mountains,Puerto Rico1

How important is ecotypic differentiation along elevational gradients in the tropics? Reciprocal transplants of two shrubs, Clibadium erosum (Asteraceae) and Psychotria berteriana (Rubiaceae), and a palm, Preitoea acuminata var. montana (Palmaceae), were used to test for the effect of environment and population origin on growth and physiology in the Luquillo Experimental Forest of Puerto Rico. Two sites were used, one at Pico del Este (1000 m in cloud forest) and one at El Verde (350 m in lower montane rain forest). At the cloud forest site, plastic barriers were erected around a subset of the plants to examine if protection from wind affected survival or biomass accumulation. Survival of C. erosum and P. berteriana was not affected by site, population origin, or the presence of barriers. For P. acuminata var. montana, survival was higher for plants with barriers, but not affected by site and population origin. Plants of C. erosum and P. berteriana at El Verde grew larger than at Pico del Este, but there was no effect of population origin or barrier treatment on biomass accumulation for these species. For P. acuminata var. montana, there was no effect of environment, population origin, or barrier treatment on biomass accumulation. Light‐saturated photosynthetic rate (A_max) of C. erosum, P. berteriana, and P. acuminata var. montana, as well as leaf anatomical characteristics of C. erosum, were unaffected by environment, population origin, and barrier treatment. On balance, there seems to be little evidence of ecotypic differentiation in these species along the gradient.     

Growth Rings,Phenology, Hurricane Disturbance and Climate in Cyrilla racemiflora L., a Rain Forest Tree of the Luquillo Mountains,Puerto Rico1

The growth phenology of Cyrilla racemiflora L., the dominant tree species of the montane rain forest, (subtropical lower montane rain forest, sensu Holdridge) of the Luquillo Mountains of Puerto Rico was studied intensively during 1989, and then semiannually through mid-1993 to determine the periodicity of changes in xylem structure. Four trees at 770 m were monitored for flowering, branch elongation, leaf litterfall, and xylem cell growth and differentiation in the lower stem, and these events were related to local seasonal patterns of rainfall and temperature. Hurricane Hugo defoliated study trees in September, 1989. Bud-break and branch elongation in March, 1989 were followed by earlywood xylem cell production in the lower stem in April and the onset of flowering in May. Leaf litterfall was greatest between April and June, coinciding with peak branch growth and new leaf formation. Latewood xylem was produced in December. The general phenological pattern was synchronized between trees and over study years. Vessel diameter and density were monitored along with thickness of earlywood and latewood and the former converted to vessel lumen area, a measure of xylem conductance capacity. Annual growth rings were formed with periods of earlywood and latewood production coinciding with traditional summer (rainy) and winter (dry) seasons, respectively, in the Luquillo Mountains. Hurricane defoliation was followed by heavy flowering in 1990, a year of reduced branch elongation and annual xylem ring width, and was associated with maximum vessel lumen area, as was flowering in 1989, prior to the hurricane. Hurricane Hugo provided a perturbation that, through its elicited stress response, allowed for the demonstration of the interplay between flowering, branching, structural growth of xylem, and xylem function.     

Soil Phosphorus Fractionation during Forest Development on Landslide Scars in the Luquillo Mountains,Puerto Rico1

Mineral soils from a chronosequence of landslide scars ranging in age from 1 to more than 55 years in a subtropical montane rain forest of eastern Puerto Rico were used to determine the rate at which labile P capital recovers during primary succession. Nine organic and inorganic soil P fractions were measured using the Hedley sequential extraction procedure. Deep soil cores (9 m) from a nearby site were also analyzed to determine the distribution of P fractions below the solum. Litterfall P was measured for two years in the landslide scars to estimate allochthonous litter P inputs, and published precipitation data were used to estimate annual atmospheric inputs of P to the recovering forests. In the upper solum (0–10 cm), organic matter increased with landslide age, as did resin‐Pi, labile P (defined here as resin‐Pi + HCO₃‐Pi + HCO₃‐Po) and total organic P. Occluded P decreased with increasing landslide age. No significant changes in P concentrations or pools were observed in 10 to 35 or in 35 to 60 cm depth intervals across the chronosequence. Labile soil P increased to approximately two‐thirds of the pre‐disturbance levels in the oldest landslide scar (>55 yr). Thus, plants, their associated microflora/fauna, and P inputs from off‐site substantially altered the distribution of soil P fractions during forest recovery. Across the chronosequence, the increase in labile P accumulated in soil and biomass appeared to be greater than the estimated allochthonous inputs from litter and precipitation, indicating that as the forest developed, some occluded P may have been released for use by soil biota. Resin‐Pi and labile P were correlated with soil organic matter content, suggesting, as in other highly weathered soils, organic matter accumulation and turnover are important in maintaining labile P pools. Primary mineral P (apatite) was scarce, even in deep soil cores.     

Land-Use History and Forest Regeneration in the Cayey Mountains, Puerto Rico

Although deforestation continues to be a major threat to tropical biodiversity, abandonment of agricultural land in Puerto Rico provides an opportunity to study long-term patterns of secondary forest regeneration. Using aerial photographs from 1937, 1967, and 1995, we determined land-use history for 2443 ha in the Cayey Mountains. Pastures were the dominant land cover in 1937 and <20% of the area was classified as forest. Between 1937 and 1995, forest cover increased to 62% due to widespread abandonment of agriculture. To examine the effect of historic land use on current forest structure and species composition, we sampled secondary forests in 24 abandoned pastures, 9 abandoned coffee plantations and 4 old-growth forest sites. Sites were located on two soil types along an elevational gradient (125–710 m) and included a chronosequence from 4 to over 80 years old. After 25–30 years, basal area and species richness in secondary forest sites derived from abandoned pastures and coffee plantations were similar to old-growth forest sites. The species composition of secondary forests derived from abandoned pastures and coffee plantations remained distinct from old-growth forest. In addition to historic land use, age and elevation were important environmental variables explaining variation in secondary forest species composition. Non-indigenous species were common in recently abandoned pastures and coffee plantations, but their importance declined in the older sites. This study demonstrates that secondary forests on private land can be an important component of the conservation of tropical tree biodiversity. Received 16 June 1999; Accepted 8 October 1999.     

Forest Succession in Tropical Hardwood Hammocks of the Florida Keys: Effects of Direct Mortality from Hurricane Andrew1

A tree species replacement sequence for dry broadleaved forests (tropical hardwood hammocks) in the upper Florida Keys was inferred from species abundances in stands abandoned from agriculture or other anthropogenic acitivities at different times in the past. Stands were sampled soon after Hurricane Andrew, with live and hurricane‐killed trees recorded separately; thus it was also possible to assess the immediate effect of Hurricane Andrew on stand successional status. We used weighted averaging regression to calculate successional age optima and tolerances for all species, based on the species composition of the pre‐hurricane stands. Then we used weighted averaging calibration to calculate and compare inferred successional ages for stands based on (1) the species composition of the pre‐hurricane stands and (2) the hurricane‐killed species assemblages. Species characteristic of the earliest stages of post‐agricultural stand development remains a significant component of the forest for many years, but are gradually replaced by taxa not present, even as seedlings, during the first few decades. This compositional sequence of a century or more is characterized by the replacement of deciduous by evergreen species, which is hypothesized to be driven by increasing moisture storage capacity in the young organic soils. Mortality from Hurricane Andrew was concentrated among early‐successional species, thus tending to amplify the long‐term trend in species composition.     

Variation in nutrient characteristics of surface soils from the Luquillo Experimental Forest of Puerto Rico: A multivariate perspective

We assessed the effects of landscape features (vegetation type and topography), season, and spatial hierarchy on the nutrient content of surface soils in the Luquillo Experimental Forest (LEF) of Puerto Rico. Considerable spatial variation characterized the soils of the LEF, and differences between replicate sites within each combination of vegetation type (tabonuco vs. palo colorado vs. dwarf vs. pasture) and topographic position (ridge vs valley) accounted for 11–60% of the total variation in soil properties. Nevertheless, mean soil properties differed significantly among vegetation types, between topographic positions, and between seasons (wet vs dry). Differences among vegetation types reflected soil properties (e.g., bulk density, soil moisture, Na, P, C, N, S) that typically are related to biological processes and inputs of water. In forests, differences between topographic positions reflected elements (e.g., Ca, Mg, K, and Al) that typically are associated with geochemical processes; however, the nutrients and elements responsible for topographic differences in dwarf forest were different from those in other forest types. In pastures, differences between topographic positions were associated with the same soil properties responsible for differences among the other vegetation types. Pastures also had reduced N levels and different soil characteristics compared to undisturbed tabonuco forest. The only soil parameter that differed significantly between seasons was soil moisture. Soils of the LEF do not support the contention that N becomes limiting with an increase in elevation, and suggest that absolute pool sizes of N and P are not responsible for the reduction in productivity with elevation.     

Land‐use History Affects the Distribution of the Saprophytic Orchid Wullschlaegelia calcarata in Puerto Rico's Tabonuco Forest1

Human activities such as logging and agriculture can severely damage forest ecosystems by changing forest structure, ecosystem function, and biodiversity. These changes may have long‐lasting consequences, which influence forest recovery. We investigated the effect of past human disturbance on the current distribution of an understory, achlorophyllous orchid, Wullschlaegelia calcarata in Puerto Rico's tropical rain forest after 70 yr of recovery. Our study site was the 16‐ha Luquillo Forest Dynamics Plot located in the Luquillo Experimental Forest, which has four areas with differing intensity of land use that have been distinguished from variation in canopy cover seen in aerial photographs taken in 1936. We recorded orchids in six 10‐m‐wide, 500‐m‐long transects across four different areas of land‐use history. We found that the orchid was not present in an area of the plot which had <20 percent canopy cover in 1936, and was most abundant in the area with >80 percent canopy cover, which had been minimally impacted by human activity. Tree species composition varied among land‐use history areas, and our observations suggested that this variation might be influencing the local distribution of W. calcarata. We also measured leaf litter biomass and identified the leaves of litter in areas with and without the orchid. Litter with a high proportion of Buchenavia tetraphylla leaves had more orchids. Even though human disturbance ceased in 1932, land‐use history in the Luquillo Forest still casts a shadow over the distribution of W. calcarata.