Puerto Rico

Immigration history of amphidromous species on a Greater Antillean island

Cook, Benjamin D.; Pringle, Catherine M.; Hughes, Jane M. 2010. Immigration history of amphidromous species on a Greater Antillean island. Journal of Biogeography. 37: 270-277.

Aim To use molecular data to test for dispersal structuring in the immigration history of an amphidromous community on an island. Location The Caribbean island of Puerto Rico. Methods Mitochondrial DNA sequences were obtained from 11 amphidromous species, including shrimps, fish and a gastropod, sampled from throughout the island. The timing of population expansion (TE) in each species was calculated using nucleotide variation and molecular clock dating methods. The order of species accumulation was then reconstructed (oldest to most recent estimate for TE), and groups of species with non-overlapping estimates for TE were identified. The temporal span and average immigration rate for each group were calculated and compared with expectations of two previously published models of island immigration [the ‘dispersal-structured model of island recolonization’ (Whittaker & Jones, Oikos, 1994, 69, 524–529), which predicts short phases of rapid immigration followed by extended phases with relatively slow immigration rates; and the ‘colonization window hypothesis’ (Carine, Taxon, 2005, 54, 895–903), which suggests that opportunities for island colonization are temporally constrained to discrete waves of colonization]. Results The molecular data indicated the immigration history of Puerto Rican amphidromous fauna from the late Pleistocene through the Holocene and identified two groups of species with non-overlapping estimates for TE and one group that overlapped with the other two groups. The temporal span, average immigration rate and lack of discreteness between all three groups indicated a continuum of immigration rather than distinct phases of species arrivals. Main conclusions This study did not support the expectations of the immigration models and suggested that amphidromous species from Puerto Rico comprise a single class of marine-based dispersers. The immigration sequence we report probably reflects a recolonization chronology in this community, in keeping with the notion of species turnover through time. Four areas of future research into the immigration history of amphidromous species on islands are identified, and indicated the possibility that equilibrium processes govern long-term community change in amphidromous biota on islands

Effects of land-use change on channel morphology in northeastern Puerto Rico

Clark, J. J., and P. R. Wilcock (2000), Effects of land-use change on
channel morphology in northeastern Puerto Rico, Geol. Soc. Am. Bull.,
112(12), 1763– 1777.

Between 1830 and 1950 much of northeastern Puerto Rico was cleared for agriculture. Runoff increased by 50% and sediment supply to the river channels increased by more than an order of magnitude. Much of the land clearance extended to steep valley slopes, resulting in widespread gullying and landslides and a large load of coarse sediments delivered to the stream channels. A shift from agriculture to industrial and residential land uses over the past 50 yr has maintained the elevated runoff while sediment supply has decreased, allowing the rivers to begin removing coarse sediment stored within their channels. The size, abundance, and stratigraphic elevation of in-channel gravel bar deposits increases, channel depth decreases, and the frequency of overbank flooding increases downstream along these channels. This is presumed to be a transient state and continued transport will lead to degradation of the bed in downstream sections as the channel adjusts to the modern supply of water and sediment. A downstream decrease in channel size is contrary to the expected geometry of self-adjusted channels, but is consistent with the presence of partially evacuated sediment remaining from the earlier agricultural period. Reverse (downstream decreasing) channel morphology is not often cited in the literature, although consistent observations are available from areas with similar land-use history. Identification of reverse channel morphology along individual watercourses may be obscured in multiwatershed compilations in which other factors produce a consistent, but scattered downstream trend. Identification of reverse channel morphology along individual streams in areas with similar land-use history would be useful for identifying channel disequilibrium and anticipating future channel adjustments.


Boose, E.R., Serrano, M.I. & Foster, D.R. (2004) Landscape
and regional impacts of hurricanes in Puerto Rico. Ecological
Monographs, 74, 335–352.

Puerto Rico is subject to frequent and severe impacts from hurricanes, whose long-term ecological role must be assessed on a scale of centuries. In this study we applied a method for reconstructing hurricane disturbance regimes developed in an earlier study of hurricanes in New England. Patterns of actual wind damage from historical records were analyzed for 85 hurricanes since European settlement in 1508. A simple meteorological model (HURRECON) was used to reconstruct the impacts of 43 hurricanes since 1851. Long-term effects of topography on a landscape scale in the Luquillo Experimental Forest (LEF) were simulated with a simple topographic exposure model (EXPOS). Average return intervals across Puerto Rico for F0 damage (loss of leaves and branches) and F1 damage (scattered blowdowns, small gaps) on the Fujita scale were 4 and 6 years, respectively. At higher damage levels, a gradient was created by the direction of the storm tracks and the weakening of hurricanes over the interior mountains. Average return intervals for F2 damage (extensive blowdowns) and F3 damage (forests leveled) ranged from 15 to 33 years and 50 to 150 years, respectively, from east to west. In the LEF, the combination of steep topography and constrained peak wind directions created a complex mosaic of topographic exposure and protection, with average return intervals for F3 damage ranging from 50 years to .150 years. Actual forest damage was strongly dependent on land-use history and the effects of recent hurricanes. Annual and decadal timing of hurricanes varied widely. There was no clear centennial-scale trend in the number of major hurricanes over the historical period.

Biotic and abiotic controls on the ecosystem significance of consumer excretion in two contrasting tropical streams

Biotic and abiotic controls on the ecosystem significance of consumer excretion in two contrasting tropical streams


Article first published online: 14 JUN 2010

DOI: 10.1111/j.1365-2427.2010.02461.x

1. Excretion of nitrogen (N) and phosphorus (P) is a direct and potentially important role for aquatic consumers in nutrient cycling that has recently garnered increased attention. The ecosystem-level significance of excreted nutrients depends on a suite of abiotic and biotic factors, however, and few studies have coupled measurements of excretion with consideration of its likely importance for whole-system nutrient fluxes. 2. We measured rates and ratios of N and P excretion by shrimps (Xiphocaris elongata and Atya spp.) in two tropical streams that differed strongly in shrimp biomass because a waterfall excluded predatory fish from one site. We also made measurements of shrimp and basal resource carbon (C), N and P content and estimated shrimp densities and ecosystem-level N and P excretion and uptake. Finally, we used a 3-year record of discharge and NH4-N concentration in the high-biomass stream to estimate temporal variation in the distance required for excretion to turn over the ambient NH4-N pool. 3. Per cent C, N, and P body content of Xiphocaris was significantly higher than that of Atya. Only per cent P body content showed significant negative relationships with body mass. C:N of Atya increased significantly with body mass and was higher than that of Xiphocaris. N : P of Xiphocaris was significantly higher than that of Atya. 4. Excretion rates ranged from 0.16–3.80 lmol NH4-N shrimp)1 h)1, 0.23–5.76 lmol total dissolved nitrogen (TDN) shrimp)1 h)1 and 0.002–0.186 lmol total dissolved phosphorus (TDP) shrimp)1 h)1. Body size was generally a strong predictor of excretion rates in both taxa, differing between Xiphocaris and Atya for TDP but not NH4-N and TDN. Excretion rates showed statistically significant but weak relationships with body content stoichiometry. 5. Large between-stream differences in shrimp biomass drove differences in total excretion by the two shrimp communities (22.3 versus 0.20 lmol NH4-N m)2 h)1, 37.5 versus 0.26 lmol TDN m)2 h)1 and 1.1 versus 0.015 lmol TDP m)2 h)1), equivalent to 21% and 0.5% of NH4-N uptake and 5% and <0.1% of P uptake measured in the high- and lowbiomass stream, respectively. Distances required for excretion to turn over the ambient NH4-N pool varied more than a hundredfold over the 3-year record in the high-shrimp stream, driven by variability in discharge and NH4-N concentration. 6. Our results underscore the importance of both biotic and abiotic factors in controlling consumer excretion and its significance for nutrient cycling in aquatic ecosystems. Differences in community-level excretion rates were related to spatial patterns in shrimp biomass dictated by geomorphology and the presence of predators. Abiotic factors also had important effects through temporal patterns in discharge and nutrient concentrations. Future excretion studies that focus on nutrient cycling should consider both biotic and abiotic factors in assessing the significance of consumer excretion in aquatic ecosystems.

Correlation between earthworms and plant litter decomposition in a tropical wet forest of Puerto Rico

Dechainea, Jennifer; Ruanb, Honghua; Sanchez-de Leon, Yaniria; Zou, Xiaoming, 2005. Correlation between earthworms and plant litter decomposition in a tropical wet forest of Puerto Rico.. Pedobiologia 49 :601-607.

Earthworms are recognized to play an important role in the decomposition of organic materials. To test the use of earthworms as an indicator of plant litter decomposition, we examined the abundance and biomass of earthworms in relation to plant litter decomposition in a tropical wet forest of Puerto Rico. We collected earthworms at 0–0.1m and 0.1–0.25m soil depths from upland and riparian sites that represent the natural variation in soils and decomposition rates within the forest. Earthworms were hand-sorted and weighed for both fresh and dry biomass. Earthworms were dominated by the exotic endogeic species Pontoscolex corethrurus Mu¨ller; they were more abundant, and had higher biomasses in the upland than in riparian sites of the forest. Plant leaf litter decomposed faster in the upland than riparian sites. We found that earthworm abundance in the upper 0.1m of the soil profile positively correlated with decomposition rate of plant leaf litter. Ground litter removal had no effect on the abundance or biomass of endogeic earthworms. Our data suggest that earthworms can be used to predict decomposition rates of plant litter in the tropical wet forest, and that the decomposition of aboveground plant litter has little influence on the abundance and biomass of endogeic earthworms.

Variations in Belowground Carbon Storage and Soil CO2 Flux Rates along a Wet Tropical Climate Gradient

McGroddy, Megan; Silver, Whendee L. 2000. Variations in Belowground Carbon Storage and Soil CO2 Flux Rates along a Wet Tropical Climate Gradient. BIOTROPICA 32(4a): 614-624 .

We used a humid tropical elevation gradient to examine the relationships among climate, edaphic conditions, belowground carbon storage, and soil respiration rates. We also compared open and closed canopy sites to increase the range of microclimate conditions sampled along the gradient, and determine the effects of canopy openings on C and P storage, and C dynamics. Total soil C, the light C fraction, and all of the component fractions of the P pool were significantly related to soil moisture, and all but total soil C were also significantly related to temperature. Both labile and recalcitrant soil P fractions were negatively correlated with the light C fraction, while the dilute HCl-extractable P pool, generally thought of as intermediate in availability, was positively correlated with light C, suggesting that P may play an important role in C cycling within these systems. Total fine root biomass was greatest at 1000 m elevation and lowest at 150 m, and was strongly and positively correlated with soil moisture content. Soil respiration rates were significantly and negatively correlated with fine root biomass and the light C fraction. In forested sites, soil respiration rates were strongly and negatively correlated with total belowground C pools (soils 1 roots 1 forest floor). Belowground C pools did not follow the expected increasing trend with decreases in temperature along the gradient. Our results indicated that in humid tropical forests, the relationships among soil C and nutrient pools, soil respiration rates, and climate are complex. We suggest that frequent and prolonged anaerobic events could be important features of these environments that may explain the observed trends.


Exotic Earthworms Accelerate Plant Litter Decomposition in a Puerto Rican Pasture and a Wet Forest
Z. G. Liu and X. M. Zou
Ecological Applications
Vol. 12, No. 5 (Oct., 2002), pp. 1406-1417

Tropical land-use changes can have profound influence on earthworms that play important roles in regulating soil processes. Converting tropical forests to pastures often drastically increases the abundance of exotic earthworm populations such as Pontoscolex corethrurus. We initiated this study to examine the influence of exotic earthworms on the decomposition of plant leaves and roots in a tropical pasture and a wet forest of Puerto Rico. We employed two treatments: control with natural earthworm population, and earthworm reduction using an electroshocking technique. Decomposition rates of plant leaves on the ground surface and root materials within the surface mineral soil were estimated using a litterbag technique. To understand the role that exotic earthworms play in altering plant litter decomposition, we also compared soil CO2 evolution rates, soil microbial biomass, and physical and chemical soil properties between the controls and earthwormreduced plots during a one-year period. Earthworm populations in the electroshocked enclosures were reduced by 85% and 87% as compared with pasture and forest controls by the end of the experiment. Earthworm reduction significantly decreased the annual decay rates of plant leaves but had no effects on those of plant roots in both pasture and forest sites. Although the control plots had less mass remaining on every litterbag collecting date, significant treatment effects on leaf decomposition occurred only after 240 d in both sites. The decay rates were greater when organic materials had low carbon to nitrogen or phosphorus ratios. Soil respiration was also decreased in the earthworm-reduced plots. In contrast, soil microbial biomass C was not affected by earthworm reduction. Furthermore, there were no significant differences between the two treatments in soil bulk density, moisture content, pH, or temperature at either site. Our results suggest that exotic earthworms may accelerate leaf litter decomposition by elevating rates of litter consumption/digestion or microbial activity, rather than by improving soil physical/chemical conditions or altering microbial biomass.


Fletcher RC, Brantley SL. 2010. Reduction of bedrock blocks as corestones in the weathering profile: observations
and model. Am. J. Sci. 310:131–64

the Espiritu Santo and Mameyes rivers within the Luquillo Experimental Forest (Puerto Rico) are interpreted as corestones, reduced from initial joint-bounded bedrock blocks by subsurface weathering. Maximum corestone size, expressed as the geometric mean of the three dimensions, S 3 abc, shows a smooth envelope when plotted against elevation. We postulate that, for each catchment, they represent in situ corestones within a stratified weathering profile, many tens of meters in thickness, that has been subsequently exhumed by younger erosion. We formulate a simplified one-dimensional model for reduction in corestone size within a steady-state weathering profile that incorporates: (i) vertical fluid transport of the reactant and the soluble products of chemical weathering; (ii) linear kinetics of corestone reduction; and, subsequently, (iii) erosion. The rate of advance of a steady-state weathering profile is a statement of the mass balance between entering reactants and weathering components, here idealized as H and albite. The mathematical relations, tie the laboratory-determined rate constant for dissolution of albite (k) to a generalized kinetic constant for the rate of decrease (K) in corestone diameter to the advance rate of the weathering profile (V ). The last parentheses contain an effective roughness at the scale of the weathering profile, where S0 is the maximum size of initial bedrock blocks, inferred to be set by initial bedrock fracture spacing, and 3L* is the profile thickness. The laboratory scale roughness value, , is the ratio of the surface area accessed by BET analysis to that of the corestone grain scale. In the model, erosion is not coupled with weathering, although the presence of corestones of finite size, SE>0, exiting at the erosional surface may be postulated to affect the erosional flux. The thickness of the corestone weathering profile derived for the model for the distance between bedrock and a corestone-free saprolite cap is approximately This expression is the product of the effective pH buffering-adjusted input reactant flux per unit area times a stoichiometeric factor linking this to net albite dissolution, divided by the rate of corestone size reduction at the input concentration of protons. Further, the profile thickness scales with the input “particle” size, S0. The model fit, which yields the ratio is consistent with a rate constant for albite dissolution that lies between laboratorymeasured and field-estimated values. Sensitivity to the reaction order of albite dissolution with respect to H, N, is small, except near the base of the profile. This model yields insights into the relationship between fracture spacing and the evolution of particle size and chemistry in weathering profiles.

The Frequency and Distribution of Recent Landslides in three MontaneTropical Regions of Puerto Rico

Larsen, Matthew C.; Torres-Sanchez, Angel J. 1998. The frequency and distribution of recent landslides in three montane tropical regions of Puerto Rico. Geomorphology 24 :309-331.

Landslides are common in steep mountainous areas of Puerto Rico where mean annual rainfall and the frequency of intense storms are high. Each year, landslides cause extensive damage to property and occasionally result in loss of life. Average population density is high, 422 peoplerkm2, and is increasing. This increase in population density is accompanied by growing stress on the natural environment and physical infrastructure. As a result, human populations are more vulnerable to landslide hazards. The Blanco, Cibuco, and Coamo study areas range in surface area from 276 to 350 km2 and represent the climatologic, geographic, and geologic conditions that typify Puerto Rico. Maps of recent landslides developed from 1:20,000-scale aerial photographs, in combination with a computerized geographic information system, were used to evaluate the frequency and distribution of shallow landslides in these areas. Several types of landslides were documented— rainfall-triggered debris flows, shallow soil slips, and slumps were most abundant. Hillslopes in the study area that have been anthropogenically modified, exceed 128 in gradient, are greater than 300 m in elevation, and face the east-northeast, are most prone to landsliding. A set of simplified matrices representing geographic conditions in the three study areas was developed and provides a basis for the estimation of the spatial controls on the frequency of landslides in Puerto Rico. This approach is an example of an analysis of the frequency of landslides that is computationally simple, and therefore, may be easily transferable to other settings.


Larsen, M.C., Torres-Sánchez, A.J., and Concepción, I.M., 1998, Slopewash, surface runoff, and fine-litter transport in forest and landslide scars in humid-tropical steeplands, Luquillo Experimental Forest, Puerto Rico [abs] EOS, Transactions American Geophysical Union, vol. 80.

Rainfall, slopewash (the erosion of soil particles), surface runoff, and fine-litter transport at humid-tropical steepland sites in the Luquillo Experimental Forest, Puerto Rico (18? 20' N, 65? 45' W) were measured from 1991 to 1995. Hillslopes underlain by: 1) Cretaceous tuffaceous sandstone and siltstone in subtropical rain (tabonuco) forest with vegetation recovering from Hurricane Hugo (1989); and underlain by 2) Tertiary quartz diorite in subtropical lower montane wet (colorado and dwarf) forest with undisturbed forest canopy were compared to recent landslide scars. Monthly surface runoff on these very steep hillslopes (24? to 43?) was only 0.2 to 0.5 percent of monthly rainfall. Slopewash was higher in sandy loam soils whose parent material is quartz diorite (averaging 46 g m-2 a-1) than in silty-clay loam soils derived from tuffaceous sandstone and siltstone where the average was 9 g m-2 a-1. Annual slopewash of 100 to 349 g m-2 on the surfaces of two recent, small landslide scars was measured initially but slopewash decreased to only 3 to 4 g m-2 a-1 by the end of the study. The mean annual mass of fine litter (mainly leaves and twigs) transported downslope at the forested sites ranged from 5 to 8 g m-2 and was lower at the tabonuco forest site, where post-Hurricane Hugo recovery is still in progress. Mean annual fine-litter transport was 2.5 g m-2 on the two landslide scars.
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