Effects of nutrient additions on ecosystem carbon cycle in a Puerto Rican tropical wet forest

LI, YIQING; XU, MING; ZOU, XIAOMING 2006. Effects of nutrient additions on ecosystem carbon cycle in a Puerto Rican tropical wet forest. Global Change Biology 11, :1-10,.

Wet tropical forests play a critical role in global ecosystem carbon (C) cycle, but C allocation and the response of different C pools to nutrient addition in these forests remain poorly understood. We measured soil organic carbon (SOC), litterfall, root biomass, microbial biomass and soil physical and chemical properties in a wet tropical forest from May 1996 to July 1997 following a 7-year continuous fertilization. We found that although there was no significant difference in total SOC in the top 0–10cm of the soils between the fertilization plots (5.42  0.18 kgm2) and the control plots (5.27  0.22 kgm2), the proportion of the heavy-fraction organic C in the total SOC was significantly higher in the fertilized plots (59%) than in the control plots (46%) (Po0.05). The annual decomposition rate of fertilized leaf litter was 13% higher than that of the control leaf litter.We also found that fertilization significantly increased microbial biomass (fungi1bacteria) with 952  48mgkg1soil in the fertilized plots and 755  37mgkg1soil in the control plots. Our results suggest that fertilization in tropical forests may enhance long-term C sequestration in the soils of tropical wet forests.

carbon isotope characterization of vegetation and soil organic matter in subtropical forests in luquillo, puerto rico

Carbon Isotope Characterization of Vegetation and Soil Organic Matter in Subtropical Forests in Luquillo, Puerto Rico
Joseph C. von Fischer and Larry L. Tieszen
Vol. 27, No. 2 (Jun., 1995), pp. 138-148

We examined natural abundances of 13C in vegetation and soil organic matter (SOM) of subtropical wet and rain forests to characterize the isotopic enrichment through decomposition that has been reported for temperate forests. Soil cores and vegetative samples from the decomposition continuum (leaves, new litter, old litter, wood, and roots) were taken from each of four forest types in the Luquillo Experimental Forest, Puerto Rico. SOM δ13C was enriched 1.6% relative to aboveground litter. We found no further enrichment within the soil profile. The carbon isotope ratios of vegetation varied among forests, ranging from -28.2% in the Colorado forest to -26.9% in the Palm forest. Isotope ratios of SOM differed between forests primarily in the top 20 cm where the Colorado forest was again most negative at -28.0%, and the Palm forest was most positive at -26.5%. The isotopic differences between forests are likely attributable to differences in light regimes due to canopy density variation, soil moisture regimes, and/or recycling of CO2. Our data suggest that recalcitrant SOM is not derived directly from plant lignin since plant lignin is even more 13C depleted than the bulk vegetation. We hypothesize that the anthropogenic isotopic depletion of atmospheric CO2 (ca 1.5% in the last 150 years) accounts for some of the enrichment observed in the SOM relative to the more modern vegetation in this study and others. This study also supports other observations that under wet or anaerobic soil environments there is no isotopic enrichment during decomposition or with depth in the active profile.

Effects of different types of conditioning on rates of leaf-litter shredding by Xiphocaris elongata, a Neotropical freshwater shrimp

Crowl, Todd A.; Welsh, Vanessa; Heartsill-Scalley, Tamara 2006. Effects of different types of conditioning on rates of leaf-litter shredding by Xiphocaris elongata, a Neotropical freshwater shrimp.. J. N. Am. Benthol. Soc., 25(1):198-208.

Temperate headwater streams with closed canopies rely on inputs of terrestrially derived organic matter to provide the major energy basis for their food webs. Microbial colonization, or conditioning, makes leaf litter more nutritional and palatable to stream detritivores, but few studies have investigated the relative importance of litter source to macroshredders in tropical streams. We determined the source (terrestrial, aquatic, or aerial), quantity, and species composition of allochthonous inputs into the Quebrada Prieta, a tropical headwater stream in Puerto Rico, as a first step toward understanding the importance of conditioning history to rates of tropical leaf-litter processing by decapod consumers. Fresh leaves of 4 common species of leaves were treated by exposing them to different conditions for 2 wk. These exposure treatments (conditioning histories) represented routes by which leaves might enter streams and included submersion (aquatic input), incubation on the streambank soil (terrestrial input), and suspension above the ground (aerial input). Conditioned leaves were placed in small experimental microcosms with or without shrimp (Xiphocaris elongata) for 20 d. Shrimp significantly increased the rate of decomposition of all leaf species independent of conditioning history. Conditioning history had little effect on breakdown rates independent of the presence of shrimp. One species (Rourea surinamensis) had faster mass loss when the leaves were conditioned as aquatic inputs rather than as terrestrial or aerial inputs. Our results indicate that conditioning history has little effect on the ability of some macroconsumers to alter detrital foodweb dynamics in tropical streams. Tropical stream ecosystems may function differently from temperate ecosystems because of the dominance of large detritivores such as shrimps.


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.

Comparing soil organic carbon dynamics in plantation and secondary forest in wet tropics in Puerto Rico

YIQING, LI; XU, MING; ZOU XIAOMING; SHI§, PEIJUN; ZHANG, YAOQI 2005. Comparing soil organic carbon dynamics in plantation and secondary forest in wet tropics in Puerto Rico. Global Change Biology 11,: 239–248, doi: 10.1111/j.1365-2486.2005.00896.x.

We compared the soil carbon dynamics between a pine plantation and a secondary forest, both of which originated from the same farmland abandoned in 1976 with the same cropping history and soil conditions, in the wet tropics in Puerto Rico from July 1996 to June 1997. We found that the secondary forest accumulated the heavy-fraction organic carbon (HF-OC) measured by the density fractionation technique, more efficiently than the tree plantation did. Although there was no significant difference in total soil organic carbon (SOC) between the plantation (5.59  0.09 kgm2) and the secondary forest (5.68  0.16 kgm2), the proportion of HF-OC carbon to the total SOC was significantly higher in the secondary forest (61%) than in the plantation (45%) (Po0.05). Forest floor mass and aboveground litterfall in the plantation were 168% and 22.8% greater than those in the secondary forest, respectively, while the decomposition rate of leaf litter in the plantation was 23.3% lower than that in the secondary forest. The annual mean soil respirations in the plantation and the secondary forest were 2.32  0.15 and 2.65  0.18 gCm2 day1, respectively, with a consistently higher rate in the secondary forest than in the plantation throughout the year. Microbial biomass measured by fumigation–incubation method demonstrated a strong seasonal variation in the secondary forest with 804mgkg1 in the wet season and 460mgkg1 in the dry season. However, the seasonal change of microbial biomass in the plantation was less significant. Our results suggested that secondary forests could stock more long-term SOC than the plantations in the wet tropics because the naturally generated secondary forest accumulated more HF-OC than the managed plantation.

Short-Term Disappearance of Foliar Litter in Three Species Before and After a Hurricane'

Short-Term Disappearance of Foliar Litter in Three Species before and after a Hurricane
Neal H. Sullivan, William B. Bowden and William H. McDowell
Vol. 31, No. 3 (Sep., 1999), pp. 382-393

Litter disappearance was examined before (1989) and after (1990) Hurricane Hugo in the Luquillo Experimental Forest, Puerto Rico using mesh litterbags containing abscised Cyrilla racemiflora or Dacryodes excelsa leaves or fresh Prestoea montana leaves. Biomass and nitrogen dynamics were compared among: (i) species; (ii) mid- and high elevation forest types; (iii) riparian and upland sites; and (iv) pre- and post-hurricane disturbed environments. Biomass disappearance was compared using multiple regression and negative exponential models in which the slopes were estimates of the decomposition rates subsequent to apparent leaching losses and the y-intercepts were indices of initial mass losses (leaching). Cyrilla racemiflora leaves with low nitrogen (0.39%) and high lignin (22.1%) content decayed at a low rate and immobilized available nitrogen. Dacryodes excelsa leaves had moderate nitrogen (0.67%) and lignin (16.6%) content, decayed at moderate rates, and maintained the initial nitrogen mass. Prestoea montana foliage had high nitrogen (1.76%) and moderate lignin (16.7%) content and rapidly lost both mass and nitrogen. There were no significant differences in litter disappearance and nitrogen dynamics among forest types and slope positions. Initial mass loss of C. racemiflora leaves was lower in 1990 but the subsequent decomposition rate did not change. Initial mass losses and the overall decomposition rates were lower in 1990 than in 1989 for Dacryodes excelsa. Dacryodes excelsa and C. racemiflora litter immobilized nitrogen in 1990 but released 10-15 percent of their initial nitrogen in 1989, whereas P. montana released nitrogen in both years (25-40%). Observed differences in litter disappearance rates between years may have been due to differences in the timing of precipitation. Foliar litter inputs during post-hurricane recovery of vegetation in Puerto Rico may serve to immobilize and conserve site nitrogen.


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.

Decay rate and substrate quality of fine roots and foliage of two tropical tree species in the Luquillo Experimental Forest, Puerto Rico

Bloomfield, J., K. A. Vogt, and D. J. Vogt. 1993. Decay rate and substrate quality of fine roots and foliage of two tropical tree species in the Luquillo Experimental Forest, Puerto Rico. Plant and Soil 150:233-245.

Decomposition rates, initial chemical composition, and the relationship between initial chemistry and mass loss of fine roots and foliage were determined for two woody tropical species, Prestoea montana and Dacryodes excelsa, over a gradient of sites in two watersheds in the Luquillo Experimental Forest, Puerto Rico. At all locations, fine roots decayed significantly more slowly than foliage during the initial 6 months. Substrate quality of the initial tissue showed marked differences between roots and foliage when using cell wall chemistry, secondary chemistry and total elemental analysis as indices. Quantity of acid detergent fiber (ADF) (non-digestible cell wall fiber) and lignin content were higher for roots than leaves: D. excelsa roots had 55.3% ADF and 28.7% lignin while leaves had 36.2% ADF and 11.8% lignin; P. montana roots had 68.0% ADF and 26.8% lignin while leaves had 48.5% ADF and 16.1% lignin. Aluminum concentrations were higher in fine roots (843 mg kg -x in D. excelsa, 1500 mg kg -x in P. montana) than leaves (244mg kg -x in D. excelsa, 422mgkg -1 in P. montana), while calcium concentrations were higher in foliage (5.5 mg g-1 in D. excelsa, 7.8 mg g-X in P. montana) than roots (3.4mgg -1 in D. excelsa, 3.1 mgg -x in P. montana). Nitrogen did not show any trend with tissue or species type. A linear model between mass remaining after 6 months and initial tissue chemistry could be developed only for calcium (r 2= 0.64).

Top-down effects of a terrestrial frog on forest nutrient dynamics

Beard,Karen H.; Vogt, Kristiina A.; Kulmatiski,Andrew 2002. Top-down effects of a terrestrial frog on forest nutrient dynamics.. Oecologia 133 :583 593.

Many studies have found top-down effects of predators on prey, but few studies have linked top-down effects of vertebrate predators to nutrient cycling rates in terrestrial systems. In this study, large and significant effects of a terrestrial frog, Eleutherodactylus coqui (coqu), were recorded on nutrient concentrations and fluxes in a subtropical wet forest. In a manipulative experiment, coqus at natural densities were contained in or excluded from 1 m3 enclosures for 4 months. Chemistry of leaf wash (throughfall), foliage, and decomposed leaf litter in the enclosures were measured as indicators of coqu effects on nutrient cycling. Coqu exclusion decreased elemental concentrations in leaf washes by 83% for dissolved organic C, 71% for NH4 +, 33% for NO3 –, 60% for dissolved organic N, and between 60 and 100% for Ca, Fe, Mg, Mn, P, K, and Zn. Coqu exclusion had no effect on foliar chemistry of plants transplanted into the enclosures. However, coqu exclusion decreased nutrient availability in decomposing mixed leaf litter by 12% and 14% for K and P, respectively, and increased C:N ratios by 13%. Changes in nutrient concentrations that occurred with coqu exclusion appear to be due to concentrations of nutrients in coqu waste products and population turnover. The results supported our hypothesis that coqus have an observable effect on nutrient dynamics in this forest. We suggest that the primary mechanism through which they have this effect is through the

The effects of the frog Eleutherodactylus coqui on invertebrates and ecosystem processes at two scales in the Luquillo Experimental Forest, Puerto Rico

The Effects of the Frog Eleutherodactylus coqui on Invertebrates and Ecosystem Processes at Two Scales in the Luquillo Experimental Forest, Puerto Rico
Karen H. Beard, Anne K. Eschtruth, Kristiina A. Vogt, Daniel J. Vogt and Frederick N. Scatena
Journal of Tropical Ecology
Vol. 19, No. 6 (Nov., 2003), pp. 607-617

Determining the ubiquity of top-down control effects of predators on their prey and ecosystem processes is important for understanding community and ecosystem-level consequences that may result from predator loss. We conducted experiments at two spatial scales to investigate the effects of terrestrial frogs (Eleutherodactylus coqui) on aerial and litter invertebrates, plant growth and herbivory, and litter decomposition. At both scales, frogs reduced aerial invertebrates and leaf herbivory, but had no effect on litter invertebrates. At the smaller scale, frogs increased foliage production rates, measured as the number of new leaves and new leaf area produced, by 80% and decomposition rates by 20%. The influence of E. coqui on increasing primary productivity and decomposition rates at the smaller scale appeared to be a result of elimination and excretion rather than of controlling prey. While the results provide evidence for frogs controlling herbivorous prey at both scales, species effects on ecosystem processes were only detectable at the smaller scale. The results highlight the difficulties in conducting experiments at large spatial scales. The findings from this study imply that the loss of amphibians and other species of higher trophic levels may affect nutrient cycling rates in tropical forests.
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