Scatena F.N.

Estimating fog deposition at a Puerto Rican elfin cloud forest site: comparison of the water budget and eddy covariance methods

Holwerda, F., R. Burkard, W. Eugster, F. N. Scatena, A. G. C. A. Meesters,
and L. A. Bruijnzeel (2006), Estimating fog deposition at a Puerto
Rican elfin cloud forest site: Comparison of the water budget and eddy
covariance methods, Hydrol. Processes, 20, 2669– 2692.

The deposition of fog to a wind-exposed 3 m tall Puerto Rican cloud forest at 1010 m elevation was studied using the water budget and eddy covariance methods. Fog deposition was calculated from the water budget as throughfall plus stemflow plus interception loss minus rainfall corrected for wind-induced loss and effect of slope. The eddy covariance method was used to calculate the turbulent liquid cloud water flux from instantaneous turbulent deviations of the surface-normal wind component and cloud liquid water content as measured at 4 m above the forest canopy. Fog deposition rates according to the water budget under rain-free conditions (0Ð11 š 0Ð05 mm h1) and rainy conditions (0Ð24 š 0Ð13 mm h1) were about three to six times the eddy-covariance-based estimate (0Ð04 š 0Ð002 mm h1). Under rain-free conditions, water-budget-based fog deposition rates were positively correlated with horizontal fluxes of liquid cloud water (as calculated from wind speed and liquid water content data). Under rainy conditions, the correlation became very poor, presumably because of errors in the corrected rainfall amounts and very high spatial variability in throughfall. It was demonstrated that the turbulent liquid cloud water fluxes as measured at 4 m above the forest could be only ¾40% of the fluxes at the canopy level itself due to condensation of moisture in air moving upslope. Other factors, which may have contributed to the discrepancy in results obtained with the two methods, were related to effects of footprint mismatch and methodological problems with rainfall measurements under the prevailing windy conditions. Best estimates of annual fog deposition amounted to ¾770 mm year1 for the summit cloud forest just below the ridge top (according to the water budget method) and ¾785 mm year1 for the cloud forest on the lower windward slope (using the eddy-covariance-based deposition rate corrected for estimated vertical flux divergence). Copyright  2006 John Wiley & Sons, Ltd.

Throughfall in a Puerto Rican lower montane rain forest: A comparison of sampling strategies

Holwerda, F.; Scatena, F.N.; Bruijnzeel, L.A. 2006. Throughfall in a Puerto Rican lower montane rain forest: A comparison of sampling strategies. Journal of Hydrology 327, :592- 602.

During a one-year period, the variability of throughfall and the standard errors of the means associated with different gauge arrangements were studied in a lower montane rain forest in Puerto Rico. The following gauge arrangements were used: (1) 60 fixed gauges, (2) 30 fixed gauges, and (3) 30 roving gauges. Stemflow was measured on 22 trees of four different species. An ANOVA indicated that mean relative throughfall measured by arrangements 1 (77%), 2 (74%), and 3 (73%) were not significantly different at the 0.05 level. However, the variability of the total throughfall estimate was about half as high for roving gauges (23%) as for fixed gauges (48–49%). The variability of stemflow ranged from 36% to 67% within tree species and was 144% for all sampled trees. Total stemflow was estimated at 4.1% of rainfall, of which palms contributed about 66%. Comparative analysis indicated that while fixed and roving gauge arrangements can give similar mean values, least 100 fixed gauges are required to have an error at the 95% confidence level comparable to that obtained by 30 roving gauges.

Disturbance and long-term patterns of rainfall and throughfall nutrient fluxes in a subtropical wet forest in Puerto Rico

Heartsill-Scalley, T.; Scatena,F.N.; Estrada,C.; McDowell,W.H.;Lugo,A.E. 2007. Disturbance and long-term patterns of rainfall and throughfall nutrient fluxes in a subtropical wet forest in Puerto Rico. Journal of Hydrology 333, :472- 485.

Nutrient fluxes in rainfall and throughfall were measured weekly in a mature subtropical wet forest in NE Puerto Rico over a 15-year period that included the effects of 10 named tropical storms, several prolonged dry periods, and volcanic activity in the region. Mean annual rainfall and throughfall were 3482 and 2131 mm yr1, respectively. Average annual rainfall and throughfall fluxes of K, Ca, Mg, Cl, Na, and SO4–S were similar but somewhat larger than those reported for most tropical forests. Rainfall inputs of nitrogen were comparatively low and reflect the relative isolation of the airshed. More constituents had seasonal differences in rainfall fluxes (6 out of 12) than throughfall fluxes (4 out of 12) and all volume weighted throughfall enrichment ratios calculated for the 15-year period were greater than one. However, median weekly enrichment ratios were less than 1 for sea salts and dissolved organic carbon, between 1 and 2 for Mg, Ca, SiO2 and SO4–S, and greater than 10 for NH4–N, PO4–P, and K. Droughts tended to reduce enrichment ratios of cations and sea-salts, but increased enrichment ratios of NH4–N, PO4–P, and K. In the weeks following hurricanes and tropical storms, relative throughfall tended to be higher and enrichment ratios tended to be lower. Saharan dust and the activity of Caribbean volcanoes can also be detected in the time series. Nevertheless, the impacts of particular events are variable and modified by the magnitude of the event, the preand post-event rainfall, and the time since the previous event. Rainfall, throughfall, rainfall pH, and rainfall fluxes of seven constituents had decreasing trends over the 15-year period. However, these decreases were small, less than inter-annual and annual varia-tions, and not considered to be ecologically significant. These long-term observations indicate that physical and biological processes associated with water passing through the canopy act to buffer internal nutrient cycles from inter-annual and seasonal variations in rainfall inputs.

The effect of human activity on the structure and composition of a tropical forest in Puerto Rico

Garcia-Montiel, D.C.; Scatena, F.N. 1994. The effect of human activity on the structure and composition of a tropical forest in Puerto Rico. Forest Ecology and Management, 63 :57-78.

From European settlement to the 1940s, the Bisley watersheds of the Luquilio Experimental Forest, Puerto Rico, were used for agroforestry, selective logging, charcoal production, and timber management. Each of these activities affected different parts of the landscape in different ways and at different times. After nearly 50 years of unhindered regeneration, six impacts remain apparent: ( 1 ) shifts in the dominance and age structure of canopy species; (2) immigration of subcanopy crop species and the establishment of banana as a riparian dominant; (3) increases in the importance of canopy species used for coffee shade; (4) the impoverishment of certain commercial timber species; (5) an increase in the density of palms around abandoned charcoal kilns; (6) a reduction in the regeneration of canopy species around abandoned charcoal kilns. Changes in the above-ground nutrient pool may also have occurred. Human disturbances in the study site were progressive rather than discrete events, had adverse impacts on forest regeneration, and increased the spatial heterogeneity of the forest.

Rainfall, Runoff and Elevation Relationships in the Luquillo Mountains of Puerto Rico

Garcia, A.R. Warner, G.S. Scatena, F. and Civco, D.L. 2002. Bisley Rainfall and
Throughfall Rainfall, Runoff and Elevation Relationships in the Luquillo
mountains of Puerto Rico. Caribbean Journal of Science. 2002 (In press).
Published as Scientific Contribution No. 1642 of the Storrs Agricultural
Experiment Station.

Long-terrn rainfall and discharge data from the Luquillo Experimental Forest (LEF) were analysed to develop relationships between rainfall, stream-runoff and elevation. These relationships were then used with a Geographic Information System (GIS) to determine spatially-averaged, mean annual hydrologic budgets for watersheds and forest types within the study area. A significant relationship exists between 1) elevation and mean annual rainfall; 2) elevation and the average number of days per year without rainfall; 3) annual stream runoff and the weighted mean elevation of a watershed; and 4) annual stream runoff and the elevation of the gaging station. A comparison of rainfall patterns between a high and a low elevation station indicated that annual and seasonal variations in rainfall are similiar along the elevational gradient. However, the upper elevation station had greater annual mean rainfall (4436 mm/yr compared to 3524 mn/yr) while the lower station had a greater variation in daily, monthly, and annual totals. Model estimates indicate that a total of 3864 mm/yr (444 hm3) of rainfall falls on the forest in an average year. The Tabonuco, Colorado, Palm, and Dwarf forest types receive an estimated annual rainfall of 3537, 4191, 4167, and 4849 mm/yr, respectively. Of the average annual rainfall input, 65% (2526 mm/yr) is converted to runoff and the remainding 35% (1338 mm/yr) is lost from the system by evapotranspiration and other abstractions. In comparsion to other tropical forests, the LEF as a whole has more evapotranspiration than many tropical montane forests but less than many lowland tropical forests.


Beard, Karen H., Kristiina A. Vogt, Daniel J. Vogt, Frederick N. Scatena, Alan P. Covich, Ragnhildur Sigurdardottir, Thomas G. Siccama, and Todd A. Crowl. 2005. STRUCTURAL AND FUNCTIONAL RESPONSES OF A SUBTROPICAL FOREST TO 10 YEARS OF HURRICANES AND DROUGHTS. Ecological Monographs 75:345–361. [doi:10.1890/04-1114]

Little is known about ecosystem-level responses to multiple, climatic disturbance events. In the subtropical forests of Puerto Rico, the major natural disturbances are hurricanes and droughts. We tested the ecosystem-level effects of these disturbances in sites with different land use histories. From 1989 to 1992, data were collected to determine the effects of Hurricane Hugo and two droughts on litterfall inputs, fine-root biomass, and decomposition rates in three topographic locations (stream, riparian, upslope) within two watersheds. From 1994 to 1998, we added a third watershed and an experiment in which coarse-wood levels were manipulated to simulate hurricane inputs. Data were collected on tree and palm growth rates, litterfall inputs, fine-root biomass, and decomposition rates. From 1994 to 1998, four hurricanes and three droughts were recorded. Measured parameters had unique responses and recovery rates to hurricanes and droughts. Litterfall inputs returned to long-term mean rates within one month following droughts and small-to-moderate hurricanes but required five years to recover after an intense hurricane. In contrast, fine-root biomass recovered seven months after an intense hurricane but failed to recover after five years following a severe drought. Despite the dramatic effects of these weather events on some ecosystem parameters, we found that aboveground measures of tree and palm growth were more affected by preexisting site conditions (e.g., nitrogen availability due to past land use activities) than hurricanes or droughts. The addition of coarse woody debris increased tree and palm growth, fine-root biomass, and litter production; however, in the case of tree and palm growth, this effect was least measurable in the sites with the highest productivity. We found that decomposition rates were more controlled by litter quality than weather conditions. In conclusion, we found that certain ecosystem structures (e.g., canopy structure and fine-root biomass) generally recovered more slowly from disturbance events than certain ecosystem processes (e.g., plant growth rates, decomposition rates). We also found that past land use activities and disturbance legacies were important in determining the responses and recovery rates of the ecosystem to disturbance.

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

Cox, S. B.; Willig, M. R.; Scatena,F. N.; 2002. Variation in nutrient characteristics of surface soils from the Luquillo Experimental Forest of Puerto Rico: A multivariate perspective.. Plant and Soil 247 : 189-198.

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.

Effects of extreme low flows on freshwater shrimps in a perennial tropical stream

COVICH, A.P.; CROWL, T.A.; SCATENA, F.N. 2003. Effects of extreme low flows on freshwater shrimps in a perennial tropical stream.. Freshwater Biology 48, 1199-1206.

1. Long-term data on rainfall suggests that perennial rainforest streams rarely are subject to drying of riffles or pools in the wet, non-seasonal Caribbean climate of Puerto Rico. Unusually low rainfall in 1994 caused some headwater riffles to dry out completely, resulting in isolated pools, reduced pool volumes and loss of access to microhabitats by benthic invertebrates. 2. From 1992 to 1998, shrimp populations were sampled bimonthly using baited traps in six pools along 1200 m (from 305 to 480 m in altitude) of Quebrada Prieta, a second-order headwater stream in the Luquillo Experimental Forest (Caribbean National Forest). 3. Following contraction of the smaller and shallower pools in the most upstream sectionof the stream, mean densities of the dominant shrimp (Atya lanipes) increased from 22 to 75shrimp m)2 of pool area during the 1994 drought year. A second common species(Xiphocaris elongata) increased from 5 to 14 shrimp m)2. A smaller percentage of adults of both species was gravid during the drought. 4. Following the 1994 drought (1995–1998), densities of both shrimp species and reproductive activity of Atya returned to predrought (1990–1993) levels. However, the reproductive activity of Xiphocaris remained lower than in the predrought period. 5. It is suggested that prolonged droughts, even in tropical rainforest biomes, may significantly alter aquatic communities through localised crowding effects resulting from habitat contraction, and lead to prolonged decreases in reproductive output. Consequently, major alterations in aquatic populations and communities would be predicted by current climate change scenarios of decreased total rainfall and increased variability.

An Annotated List of the Flora of the Bisley Area, Luquillo Experimental Forest, Puerto Rico 1987 to 1992

Chinea, J. D., R. J. Beymer, C. Rivera, I. Sastre de Jesu´ s, and F.
N. Scatena. 1993. An annotated list of the flora of the Bisley
Area, Luquillo Experimental Forest, Puerto Rico 1987 to 1992.
General Technical Report SO-94:1–12. U.S. Department of Agriculture,
Southern Forest Experiment Station, New Orleans.

Known species of plants, including bryophytes and ferns, are listed for the area of the Bisley experimental watershed area, a subtropical wet forest in the Luquillo Mountains of northeastern Puerto Rico.

Effects of an invasive tree on community structure and diversity in a tropical forest in Puerto Rico

Brown, K. A. ; Scatena, F. N., and Gurevitch, J. 2006. Effects of an invasive tree on community structure and diversity in a tropical forest in Puerto Rico. . Forest Ecology and Management . 2006; 226:145-152.

We report the effects of an invasive tree (Syzygium jambos, Myrtaceace) on species composition, plant diversity patterns, and forest regeneration in primary and secondary forest in the Luquillo Mountains of northeastern Puerto Rico, including the area in and around the Caribbean National Forest (CNF) and the Luquillo Long Term Ecological Research site (Luquillo LTER). Land use history was reconstructed using aerial photographs from 1936 to 1989 and study sites were categorized into four groups that corresponded to their status in 1936: unforested, young secondary, mature secondary, and primary forests. In randomly selected forest stands in each forest type, we measured the abundance of invasive and native tree species, seedling recruitment for S. jambos as well as soil nutrient pools and tested for the effects of land use history on S. jambos density and diversity. A partial Mantel test was used to control for historical and elevational differences across study sites. The results indicate that S. jambos density was highest in habitats classified in 1936 as unforested, young, or mature secondary forests. Compared to all other forest classes, species diversity was significantly higher in primary forests. However, there was no statistically significant difference between observed and estimated species richness across the four forest types. S. jambos density and species diversity were strongly negatively correlated, even after controlling for land use history and elevation. There was significantly higher S. jambos seedling recruitment in areas that were either unforested or had young secondary forests in 1936. The results also indicate that S. jambos is able to establish viable populations in habitats with different soil nutrient status. S. jambos has also altered vegetation composition and diversity patterns in habitats where it is the dominant tree species. After nearly 185 years since its introduction to the island, S. jambos is not only well established within 30 m of stream channels, its presence does not appear to be limited by topographic, soil nutrient, or elevational conditions. This study suggests that land use change and subsequent plant invasions have produced a new vegetation assemblage that has led to potentially long-term changes in community structure, species composition, and successional trajectory in regenerating secondary forests in the Luquillo Mountains.
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