Ecology

Does the river continuum concept apply on a tropical island? Longitudinal variation in a Puerto Rican stream

Greathouse,Effie A.; Pringle, Catherine M. 2006. Does the river continuum concept apply on a tropical island? Longitudinal variation in a Puerto Rican stream.. Can. J. Fish. Aquat. Sci. 63: 134-152.

Abstract: 
We examined whether a tropical stream in Puerto Rico matched predictions of the river continuum concept (RCC) for macroinvertebrate functional feeding groups (FFGs). Sampling sites for macroinvertebrates, basal resources, and fishes ranged from headwaters to within 2.5 km of the fourth-order estuary. In a comparison with a model temperate system in which RCC predictions generally held, we used catchment area as a measure of stream size to examine truncated RCC predictions (i.e., cut off to correspond to the largest stream size sampled in Puerto Rico). Despite dominance of generalist freshwater shrimps, which use more than one feeding mode, RCC predictions held for scrapers, shredders, and predators. Collector–filterers showed a trend opposite to that predicted by the RCC, but patterns in basal resources suggest that this is consistent with the central RCC theme: longitudinal distributions of FFGs follow longitudinal patterns in basal resources. Alternatively, the filterer pattern may be explained by fish predation affecting distributions of filter-feeding shrimp. Our results indicate that the RCC generally applies to running waters on tropical islands. However, additional theoretical and field studies across a broad array of stream types should examine whether the RCC needs to be refined to reflect the potential influence of top-down trophic controls on FFG distributions.

INDIRECT UPSTREAM EFFECTS OF DAMS: CONSEQUENCES OF MIGRATORY CONSUMER EXTIRPATION IN PUERTO RICO

GREATHOUSE, EFFIE A.; PRINGLE, CATHERINE M.; MCDOWELL, WILLIAM H.; HOLMQUIST, JEFF G. 2006. INDIRECT UPSTREAM EFFECTS OF DAMS: CONSEQUENCES OF MIGRATORY CONSUMER EXTIRPATION IN PUERTO RICO. Ecological Applications, 16(1), :339-352.

Abstract: 
Large dams degrade the integrity of a wide variety of ecosystems, yet direct downstream effects of dams have received the most attention from ecosystem managers and researchers. We investigated indirect upstream effects of dams resulting from decimation of migratory freshwater shrimp and fish populations in Puerto Rico, USA, in both high- and low-gradient streams. In high-gradient streams above large dams, native shrimps and fishes were extremely rare, whereas similar sites without large dams had high abundances of native consumers. Losses of native fauna above dams dramatically altered their basal food resources and assemblages of invertebrate competitors and prey. Compared to pools in high-gradient streams with no large dams, pool epilithon above dams had nine times more algal biomass, 20 times more fine benthic organic matter (FBOM), 65 times more fine benthic inorganic matter (FBIM), 28 times more carbon, 19 times more nitrogen, and four times more non-decapod invertebrate biomass. High-gradient riffles upstream from large dams had five times more FBIM than did undammed riffles but showed no difference in algal abundance, FBOM, or non-decapod invertebrate biomass. For epilithon of lowgradient streams, differences in basal resources between pools above large dams vs. without large dams were considerably smaller in magnitude than those observed for pools in highgradient sites. These results match previous stream experiments in which the strength of native shrimp and fish effects increased with stream gradient. Our results demonstrate that dams can indirectly affect upstream free-flowing reaches by eliminating strong top-down effects of consumers. Migratory omnivorous shrimps and fishes occur throughout the tropics, and the consequences of their declines upstream from many tropical dams are likely to be similar to those in Puerto Rico. Thus, ecological effects of migratory fauna loss upstream from dams encompass a wider variety of species interactions and biomes than the bottom-up effects (i.e., elimination of salmonid nutrient subsidies) recognized for northern temperate systems.

Conservation and management of migratory fauna and dams in tropical streams of Puerto Rico

Greathouse, E. A., C. M. Pringle, and J. G. Holmquist.
2006. Conservation and management of migratory
fauna: dams in tropical streams of Puerto
Rico. Aquatic Conservation 16:695–712.

Abstract: 
1. Compared to most other tropical regions, Puerto Rico appears to have dammed its running waters decades earlier and to a greater degree. The island has more large dams per unit area than many countries in both tropical and temperate regions (e.g., 3x that of the U.S.), and the peak rate of large dam construction occurred two and three decades prior to reported peak rates in Latin America, Asia and Africa. 2. Puerto Rico is a potential window into the future of freshwater migratory fauna in tropical regions, given the island’s extent and magnitude of dam development and the available scientific information on ecology and management of the island’s migratory fauna. 3. We review ecology, management and conservation of migratory fauna in relation to dams in Puerto Rico. Our review includes a synthesis of recent and unpublished observations on upstream effects of large dams on migratory fauna and an analysis of patterns in free crest spillway discharge across Puerto Rican reservoirs. Analyses suggest that large dams with rare spillway discharge cause near, not complete, extirpation of upstream populations of migratory fauna. They also suggest several management and conservation issues in need of further research and consideration. These include research on the costs, benefits and effectiveness of simple fish/shrimp passage designs involving simulating spillway discharge and the appropriateness of establishing predatory fishes in reservoirs of historically fishless drainages.

Structure and composition of vegetation along an elevational gradient in Puerto Rico

Gould, W.A.; González, G.; Carrero Rivera, G. 2006. Structure and composition of vegetation along an elevational gradient in Puerto Rico.. Journal of Vegetation Science 17: 563-574, .

Abstract: 
Question: What are the composition, conservation status, and structural and environmental characteristics of eight mature tropical forest plant communities that occur along an elevational gradient. Location: Northeastern Puerto Rico. Methods: We quantified the species composition, diversity, conservation status, and ecological attributes of eight mature tropical forest plant communities in replicated plots located to sample representative components of important forest types occurring along an elevational gradient. A suite of environmental and vegetation characteristics were sampled at each plot and summarized to characterize communities and analyse trends along the elevational gradient. Results: The set of communities included 374 species; 92% were native, 14% endemic, and 4% critical elements (locally endangered) to the island. All communities, occurring within a wide range of patch sizes and degree of conservation protection, showed a high percentage of native species (> 89% per plot). The lowland moist forest communities, occurring within a matrix of urbanization, agriculture, and disturbance, had the highest degree of invasion by exotics. Community descriptions were nested within a variety of hierarchies to facilitate extrapolation of community characteristics to larger ecosystem units. Basal area, above-ground biomass, canopy heights, and mean species richness peaked at mid elevations. Conclusions: It is significant that all of these forest communities continue to be dominated by native species while existing in a matrix of human and natural disturbance, species invasion, and forest regeneration from widespread agriculture. The lowland moist and dry forest types represent a minority of the protected forested areas in Puerto Rico, serve as unique genetic reservoirs, and should be protected.

Earthworm communities along an elevation gradient in Northeastern Puerto Rico

Gonzalez, Grizelle; Garcia, Emerita; Cruz, Veronica; Borges, Sonia; Zalamea, Marcela; Rivera, Maria M. 2007. Earthworm communities along an elevation gradient in Northeastern Puerto Rico.. European Journal of Soil Biology 43 .

Abstract: 
In this study, we describe earthworm communities along an elevation gradient of eight forest types in Northeastern Puerto Rico, and determine whether their abundance, biomass and/or diversity is related to climatic, soil physical/chemical and/or biotic characteristics. We found that the density, biomass, and diversity of worms varied significantly among forest types. The density of earthworms was highest in the Pterocarpus forest. In terms of biomass, both elfin and the Pterocarpus forests had the highest values. The number of earthworm species significantly increased as elevation and annual rainfall increased and air temperature decreased. We conclude that differences in earthworm species richness along this elevation gradient may be due to a combination of biotic and soil physical and chemical factors. Soil pH and root length density are important predictors of number of worm species along this elevation gradient.

Hurricane Disturbance Alters Secondary Forest Recovery in Puerto Rico

Flynn DFB, Uriarte M, Crk T et al (2009) Hurricane disturbance
alters secondary forest recovery in Puerto Rico.
Biotropica 42:149–157

Abstract: 
Land-use history and large-scale disturbances interact to shape secondary forest structure and composition. How introduced species respond to disturbances such as hurricanes in post-agriculture forest recovery is of particular interest. To examine the effects of hurricane disturbance and previous land use on forest dynamics and composition, we revisited 37 secondary forest stands in former cattle pastures across Puerto Rico representing a range of exposure to the winds of Hurricane Georges in 1998. Stands ranged from 21 to480 yr since agricultural abandonment and were measured 9 yr posthurricane. Stem density decreased as stands aged, while basal area and species richness tended to increase. Hurricane disturbance exerted contrasting effects on stand structure, contingent on stand age. In older stands, the basal area of large trees fell, shifting to a stand structure characteristic of younger stands, while the basal area of large trees tended to rise in younger stands with increasing hurricane disturbance. These results demonstrate that large-scale natural disturbances can alter the successional trajectory of secondary forest stands recovering from human land use, but stand age, precipitation and soil series were better predictors of changes in stand structure across all study sites. Species composition changed substantially between census intervals, but neither age nor hurricane disturbance consistently predicted species composition change. However, exposure to hurricane winds tended to decrease the abundance of the introduced tree Spathodea campanulata, particularly in smaller size classes. In all sites the abundance of the introduced tree Syzygium jambos showed a declining trend, again most strongly in smaller size classes, suggesting natural thinning through succession.

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

Fetcher, Ned; Cordero, Roberto A.; Voltzow, Janice 2000. Lack of Ecotypic Differentiation: Plant Response to Elevation, Population Origin, and Wind in the Luquillo Mountains, Puerto Rico. BIOTROPICA 32(2) :225-234 .

Abstract: 
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, Prestoea 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 (Amax) 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 Rico

Growth Rings, Phenology, Hurricane Disturbance and Climate in Cyrilla racemiflora L., a Rain Forest Tree of the Luquillo Mountains, Puerto Rico
Allan P. Drew
Biotropica
Vol. 30, No. 1 (Mar., 1998), pp. 35-49

Abstract: 
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.

Biological Nitrogen Fixation in Two Tropical Forests: Ecosystem-Level Patterns and Effects of Nitrogen Fertilization

Cusack DF, Silver W, McDowell WH (2009b) Biological nitrogen fixation
in two tropical forests: ecosystem-level patterns and effects of nitrogen
fertilization. Ecosystems, 12, 1299–1315.

Abstract: 
Humid tropical forests are often characterized by large nitrogen (N) pools, and are known to have large potential N losses. Although rarely measured, tropical forests likely maintain considerable biological N fixation (BNF) to balance N losses. We estimated inputs of N via BNF by free-living microbes for two tropical forests in Puerto Rico, and assessed the response to increased N availability using an on-going N fertilization experiment. Nitrogenase activity was measured across forest strata, including the soil, forest floor, mosses, canopy epiphylls, and lichens using acetylene (C2H2) reduction assays. BNF varied significantly among ecosystem compartments in both forests. Mosses had the highest rates of nitrogenase activity per gram of sample, with 11 ± 6 nmol C2H2 reduced/g dry weight/h (mean ± SE) in a lower elevation forest, and 6 ± 1 nmol C2H2/g/h in an upper elevation forest. We calculated potential N fluxes via BNF to each forest compartment using surveys of standing stocks. Soils and mosses provided the largest potential inputs of N via BNF to these ecosystems. Summing all components, total background BNF inputs were 120 ± 29 lg N/m2/h in the lower elevation forest, and 95 ± 15 lg N/m2/h in the upper elevation forest, with added N significantly suppressing BNF in soils and forest floor. Moisture content was significantly positively correlated with BNF rates for soils and the forest floor. We conclude that BNF is an active biological process across forest strata for these tropical forests, and is likely to be sensitive to increases in N deposition in tropical regions.

A Canopy Trimming Experiment in Puerto Rico: The Response of Litter Invertebrate Communities to Canopy Loss and Debris Deposition in a Tropical Forest Subject to Hurricanes

Richardson, Barbara A.; Richardson, Michael J.; Gonzalez, Grizelle; Shiels, Aaron B.; Srivastava, Diane S. 2010. A canopy trimming experiment in Puerto Rico: the response of litter invertebrate communities to canopy loss and debris deposition in a tropical forest subject to hurricanes. Ecosystems. 13: 286-301.

Abstract: 
Hurricanes cause canopy removal and deposition of pulses of litter to the forest floor. A Canopy Trimming Experiment (CTE) was designed to decouple these two factors, and to investigate the separate abiotic and biotic consequences of hurricane-type damage and monitor recovery processes. As part of this experiment, effects on forest floor invertebrate communities were studied using litterbags. Canopy opening resulted in increased throughfall, soil moisture and light levels, but decreased litter moisture. Of these, only throughfall and soil moisture had returned to control levels 9 months after trimming. Canopy opening was the major determinant of adverse changes in forest floor invertebrate litter communities, by reducing diversity and biomass, irrespective of debris deposition, which played a secondary role. Plots subjected to the most disturbance, with canopy removed and debris added, had the lowest diversity and biomass. These two parameters were higher than control levels when debris was added to plots with an intact canopy, demonstrating that increased nutrient potential or habitat complexity can have a beneficial effect, but only if the abiotic conditions are suitable. Animal abundance remained similar over all treatments, because individual taxa responded differentlyto canopy trimming. Mites, Collembola, and Psocoptera, all microbiovores feeding mainly on fungal hyphae and spores, responded positively, with higher abundance in trimmed plots, whereas all other taxa, particularly predators and larger detritivores, declined in relative abundance. Litterbag mesh size and litter type had only minor effects on communities, and canopy trimming and debris deposition explained most variation between sites. Effects of trimming on diversity, biomass, and abundance of some invertebrate taxa were still seen when observations finished and canopy closure was complete at 19 months. This suggests that disturbance has a long-lasting effect on litter communities and may, therefore, delay detrital processing, depending on the severity of canopy damage and rate of regrowth.
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