Puerto Rico


Controls of Primary Productivity: Lessons from the Luquillo Mountains in Puerto Rico
Robert B. Waide, Jess K. Zimmerman and F. N. Scatena
Vol. 79, No. 1 (Jan., 1998), pp. 31-37

The Luquillo Mountains of eastern Puerto Rico are used as a case study to evaluate possible single- or multiple-factor controls of productivity in montane forests. A review of published studies from the Luquillo Mountains revealed that canopy height, productivity, and species richness decline while stem density increases with elevation, as is typical of other montane forests. A mid-elevation floodplain palm stand with high levels of productivity provides a notable exception to this pattern. Previous basic and applied studies of productivity in the Luquillo Mountains have consistently considered the overall gradient in productivity to be important in understanding forest structure and function. Recent observational and experimental studies have determined that disturbance of all types is an important factor mediating productivity in both low- and high-elevation (cloud) forests. For example, low-elevation forest recovers more quickly from hurricane disturbance and is more responsive to nutrient additions than is cloud forest. All of the factors proposed for limiting productivity are supported in one way or another by research in the Luquillo Mountains. What is critically lacking is both an appreciation for the way that these factors interact and experiments appropriate to evaluate multiple controls acting simultaneously.


This paper compares aboveground forest structure and macronutrient stoichiometry over 5 15 years of hurricane induced secondary succession by species, life history groups, community 6 species composition, and geomorphic setting. Stem density continually increased after the 7 impact of the Category 4 hurricane Hugo and 15 years later, it was greater than pre-hurricane. 8 There were significant spatial and temporal differences in the number of species, the diversity 9 index per plot, forest structure, and biomass. The greatest compositional differences occurred 10 between the post-Hugo and the 15-year census. Prior to hurricane Hugo most plots had very 11 similar species composition and abundances, and thus occupied a small area in non-metric 12 multidimensional species space. Following the hurricane new species combinations occurred 13 and the location of plots was spread in multidimensional space. Diversity indices were 14 significantly different among geomorphic settings before and immediately after hurricane Hugo. 15 However, these differences were not observed again until the 15-year census where they returned 16 to pre-hurricane levels. Plant associations based on abundance, life history traits, and landscape 17 position had measurable differences in their structure, composition, aboveground nutrient 18 storage, and stoichiometry. However, these differences were reflected in a variety of ways at 19 different spatial scales. At the species level differences in macronutrient tissue concentrations 20 were apparent when comparing co-existing primary forests dominants, early successional 21 dominants, high-light and low-light species, and species whose stem densities are negatively 22 correlated. Community level differences were greater for forest structure and total nutrient 23 storage compared to the mass weighted concentrations of macronutrients. The largest differences observed were in Mg and can be attributed to the succession of pioneer species 2 following the hurricane. Over the entire 15-year period, the watershed average aboveground 3 stoichiometry was relatively consistent and this is linked to the biomass dominance of a few 4 species. The successional history recorded here also suggests that community level differences 5 in species composition, structure, and stoichiometry were well established after 10 to 15 years of 6 secondary succession.


Multivariate analysis of water quality and physical
characteristics of selected watersheds in Puerto Rico.
Journal of the American Water Resources Association 39:

Multivariate analyses were used to develop equations that could predict certain water quality (WQ) conditions for unmonitored watersheds in Puerto Rico based on their physical characteristics. Long term WQ data were used to represent the WQ of 15 watersheds in Puerto Rico. A factor analysis (FA) was performed to reduce the number of chemical constituents. Cluster analysis (CA) was used to group watersheds with similar WQ characteristics. Finally, a discriminant analysis (DA) was performed to relate the WQ clusters to different physical parameters and generate predicting equations. The FA identified six factors (77 percent of variation explained): nutrients, dissolved ions, sodium and chloride, silicacious geology, redox conditions, and discharge. From the FA, specific conductance, sodium, phosphorous, silica, and dissolved oxygen were selected to represent the WQ characteristics in the CA. The CA determined five groups of watersheds (forested, urban polluted, mixed urban/rural, forested plutonic, and limestone) with similar WQ properties. From the five WQ clusters, two categories can be observed: forested and urban watersheds. The DA found that changes in forest cover, percent of limestone, mean annual rainfall, and watershed shape factor were the most important physical features affecting the WQ of watersheds in Puerto Rico.

Effects of nutrient availability and other elevational changes on bromeliad populations and their invertebrate communities in a humid tropical forest in Puerto Rico

RICHARDSON, BARBARA A.; RICHARDSON,M. J.; SCATENA, F. N.; MCDOWELL, W. H. 2000. Effects of nutrient availability and other elevational changes on romeliad populations and their invertebrate communities in a humid tropical forest in Puerto Rico. Journal of Tropical Ecology 16:167±188.

Nutrient inputs into tank bromeliads were studied in relation to growth and productivity, and the abundance, diversity and biomass of their animal inhabitants, in three forest types along an elevational gradient. Concentrations of phosphorus, potassium and calcium in canopy-derived debris, and nitrogen and phosphorus in phytotelm water, declined with increasing elevation. Dwarf forest bromeliads contained the smallest amounts of debris/plant and lowest concentrations of nutrients in plant tissue. Their leaf turnover rate and productivity were highest and, because of high plant density, they comprised 12.8% of forest net primary productivity (0.47 t ha-1 y-1), and contained 3.3 t ha-1 of water. Annual nutrient budgets indicated that these microcosms were nutrient-abundant and accumulated < 5% of most nutrients passing through them. Exceptions were K and P in the dwarf forest, where accumulation was c. 25% of inputs. Animal and bromeliad biomass/plant peaked in the intermediate elevation forest, and were positively correlated with the debris content/bromeliad across all forest types. Animal species richness showed a signi®cant mid-elevational peak, whereas abundance was independent of species richness and debris quantities, and declined with elevation as forest net primary productivity declined. The unimodal pattern of species richness was not correlated with nutrient concentrations, and relationships among faunal abundance, species richness, nutrient inputs and environment are too complex to warrant simple generalizations about nutrient resources and diversity, even in apparently simple microhabitats.

Estimating soil turnover rate from tree uprooting during hurricanes in Puerto Rico

Lenart, Melanie T.; Falk, D.A.; Scatena, F.N.; Osterkamp, W.R. 2010. Estimating soil turnover rate from tree uprooting during hurricanes in Puerto Rico. Forest Ecology and Management. 259:1076-1084.

Soil turnover by tree uprooting in primary and secondary forests on the island of Puerto Rico was measured in 42 study plots in the months immediately after the passage of a Category 3 hurricane. Trunk basal area explained 61% of the variability of mound volume and 53% of the variability of mound area. The proportion of uprooted trees, the number of uprooted trees, or the proportion of uprooted basal area explained 84–85% of the variation in hurricane-created mound area. These same variables explain 79–85% of the variation in mound volume. The study indicates that the soil turnover period from tree uprooting by Puerto Rican hurricanes is between 1600 and 4800 years. These rates are faster than soil turnover by landslides and background treefall in the same area and provide a useful age constraint on soil profile development and soil carbon sequestration in these dynamic landscapes.

Helping HELP with limited resources: the Luquillo experience

Scatena, F.N.; Ortiz-Zayas, JR; Blanco-Libreros, J.F. 2008. Helping HELP with limited resources: the Luquillo experience. Water SA. 34(4 special HELP edition): 497-508.

By definition the HELP approach involves the active participation of individuals from a wide range of disciplines and backgrounds, including representatives of industry, academics, natural resource managers, and local officials and community leaders. While there is considerable enthusiasm and support for the integrated HELP approach, a central problem for all HELP basins is how to effectively engage individuals and groups with few, if any financial resources. In the Luquillo HELP project we have managed this issue by focusing our efforts on holding small, public meetings and workshops with technocrats and managers who are engaged in local water resource management. To date several forums have been organised, including: technical meetings with the directors of natural resource agencies; presentations and panel discussions at the meetings of local professional societies, including the societies of Civil Engineers and Architects, the Commonwealth Association of Tourism, the Association of Builders and Developers, and the Puerto Rican Association of Lawyers. During these forums HELP specialists gave presentations and led discussions on how integrated watershed management can help resolve local problems. Because the audience are directly involved with these issues, they are quite responsive to these discussions and have often provided unique solutions to common problems. Technical workshops are co-sponsored by local municipalities – these day-long workshops are hosted by a municipality and include managers from other municipalities, the local water authority, and local community leaders. Additional activities include: technical advice on water infrastructure projects is given; there are educational exchanges between local and international students, scientists, natural resource managers, and community leaders; and synthesis publications relevant to integrated water resource management are produced. Other activities have included compiling oral environmental histories and organising watershed restoration activities. This paper describes these activities and discusses the benefits and costs of each approach.

An EMERGY Evaluation of Puerto Rico and the Luquillo Experimental Forest

Scatena, F.N.; Doherty, S.J.; Odum, H.T.; Kharecha, P. 2002. An EMERGY
evaluation of Puerto Rico and the Luquillo Experimental Forest. Gen. Tech. Rep.
IITF-GTR-9. Río Piedras, PR: U.S. Department of Agriculture, Forest Service,
International Institute of Tropical Forestry. 79 p.

The many functions of Puerto Rico and the Luquillo Experimental Forest (the Forest) were evaluated in units of solar EMERGY, an energy-based measure of resource contribution and influence, defined as the energy of one type required to produce a flow or storage of another type. Rainfall and tectonic uplift are the largest environmental inputs into the Forest. The interaction of these inputs results in an erosional landscape where the EMERGY of biological processes is less than the EMERGY associated with the physical and chemical sculpturing of the landscape. The environmental work that built the natural capital of these forests is 9 to 50 times their current dollar market values. Of the investments evaluated in this study, the effects associated with water extraction are the largest. Tectonic inputs and the hydrologic cycle also provide most of the environmental EMERGY flows in the island of Puerto Rico. The ratio of societal inputs to environmental inputs, however, is 45 for Puerto Rico and 3.5 for the Forest. Per capita EMERGY- use is typical of moderately developed economies, but the island has one of the most investment-intensive, least self-sufficient economies known and an EMERGY signature that resembles a city-state.

Water Withdrawn From the Luquillo Experimental Forest, 2004

Crook, Kelly E.; Scatena, Fred N.; Pringle, Catherine M. 2007. Water Withdrawn From the Luquillo Experimental Forest, 2004. U.S. Department of Agriculture, Forest Service, International Institute of Tropical Forestry. Gen. Tech. Rep. IITF-GTR-34.

This study quantifies the amount of water withdrawn from the Luqillo Experimental Forest (LEF) in 2004. Spatially averaged mean monthly water budgets were generated for watersheds draining the LEF by combining long-term data from various government agencies with estimated extraction data. Results suggest that, on a typical day, 70 percent of water generated within the forest is diverted before reaching the ocean. This is up from an estimated 54 percent in 1994. Analysis showed that up to 63 percent of average monthly stream runoff is diverted from individual watersheds during drier months. Watersheds with large water intakes have the most dramatic decrease in streamflow, particularly the Río Espiritu Santo watershed, where 82 percent of median flow is diverted.


BLANCO,JUAN F.;SCATENA,FREDERICK N. 2007. The spatial arrangement of neritina virginea (gastropoda: neritidae) during upstream migration in a split-channel reach.. River Res. Applic. 23: 235-245.

This paper relates differences in flow hydraulics between a main channel (MC) and a side channel (SC) of a river to patterns of upstream migration by Neritina virginea (Neritidae: Gastropoda), a dominant diadromous snail in streams of Puerto Rico (Greater Antilles). Near-bed water velocity, snail density and shell size were measured on a weekly basis between August and December 2000 along cross-sections in a main channel (MC) and an adjacent channel (SC) under a bridge crossing of the Rio Mameyes of Northeastern Puerto Rico. Near-bed velocity and water depth were used to compute Reynolds (Re) and Froude (Fr) numbers, and to classify flows within each channel. During base flow conditions (<2m3 s1), flow was chaotic and supercritical (Fr>1) in the MC, and non-chaotic and subcritical (Fr<1) in the SC. Higher mean densities (>100 indm2) of relatively small snails (mean s.d., 6.3 2.8 mm) were consistently recorded in the MC. Conversely, the SC had lower mean densities(<20 indm2) and significantly larger snails (7.6 2.4 mm). Within the MC, migratory groups preferred near-bed velocities>0.8ms1. Within the SC, they preferred the channel thalweg and depths>30 cm. The spatial arrangement that was observed between and within the channels may be related to food resources, predation pressure or biomechanics. Characteristics of preferred upstream migration pathways of N. virginea must be accounted when building road crossings incoastal streams with diadromous fauna.

Bivergent thrust wedges surrounding oceanic island arcs: Insight from observations and sandbox models of the northeastern Caribbean plate

ten Brink U, Marshak S, Granja JL (2009) Bivergent thrust wedges
surrounding oceanic island arcs: insights from observations and
sandbox models in the north-eastern Caribbean plate. Geol Soc
Am Bull 121:1522–1536

At several localities around the world, thrust belts have developed on both sides of oceanic island arcs (e.g., Java-Timor, Panama, Vanuatu, and the northeastern Caribbean). In these localities, the overall vergence of the backarc thrust belt is opposite to that of the forearc thrust belt. For example, in the northeastern Caribbean, a north-verging accretionary prism lies to the north of the Eastern Greater Antilles arc (Hispaniola and Puerto Rico), whereas a south-verging thrust belt called the Muertos thrust belt lies to the south. Researchers have attributed such bivergent geometry to several processes, including: reversal of subduction polarity; subduction-driven mantle fl ow; stress transmission across the arc; gravitational spreading of the arc; and magmatic infl ation within the arc. New observations of deformational features in the Muertos thrust belt and of fault geometries produced in sandbox kinematic models, along with examination of published studies of island arcs, lead to the conclusion that the bivergence of thrusting in island arcs can develop without reversal of subduction polarity, without subarc mantle fl ow, and without magmatic infl ation. We suggest that the Eastern Greater Antilles arc and comparable arcs are simply crustalscale bivergent (or “doubly vergent”) thrust wedges formed during unidirectional subduction. Sandbox kinematic modeling suggests, in addition, that a broad retrowedge containing an imbricate fan of thrusts develops only where the arc behaves relatively rigidly. In such cases, the arc acts as a backstop that transmits compressive stress into the backarc region. Further, modeling shows that when arcs behave as rigid blocks, the strike-slip component of oblique convergence is accommodated entirely within the prowedge and the arc—the retrowedge hosts only dip-slip faulting (“frontal thrusting”). The existence of large retrowedges and the distribution of faulting in an island arc may, therefore, be evidence that the arc is relatively rigid. The rigidity of an island arc may arise from its mafi c composition and has implications for seismic-hazard analysis.
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