Zimmerman J.K.

The Ecological Consequences of Socioeconomic and Land-Use Changes in Postagriculture Puerto Rico

Grau, H. Ricardo; Aide, T. Mitchell; Zimmerman, Jess K.; Thomlinson, John R.; Helmer, Eileen; Zou, Xioming 2003. The ecological consequences of socioeconomic and land-use changes in post agriculture Puerto Rico. BioScience. Vol. 53, no. 12 (Dec. 2003): Pages 1159-1168.

Contrary to the general trend in the tropics, forests have recovered in Puerto Rico from less than 10% of the landscape in the late 1940s to more than 40% in the present. The recent Puerto Rican history of forest recovery provides the opportunity to study the ecological consequences of economic globalization, reflected in a shift from agriculture to manufacturing and in human migration from rural to urban areas. Forest structure rapidly recovers through secondary succession, reaching mature forest levels of local biodiversity and biomass in approximately 40 years. Despite the rapid structural recovery, the legacy of pre-abandonment land use, including widespread abundance of exotic species and broadscale floristic homogenization, is likely to persist for centuries.

Trends and scenarios of the carbon budget in postagricultural Puerto Rico (1936–2060)

Grau, H. R., T. M. Aide, J. K. Zimmerman, and J. R.
Thomlinson. 2004. Trends and scenarios of the carbon
budget in post-agricultural Puerto Rico (1936–2060). Global
Change Biology 10:1163–1179.

Contrary to the general trend in the tropics, Puerto Rico underwent a process of agriculture abandonment during the second half of the 20th century as a consequence of socioeconomic changes toward urbanization and industrialization. Using data on landuse change, biomass accumulation in secondary forests, and ratios between gross domestic product (GDP) and carbon emissions, we developed a model of the carbon budget for Puerto Rico between 1936 and 2060. As a consequence of land abandonment, forests have expanded rapidly since 1950, achieving the highest sequestration rates between 1980 and 1990. Regardless of future scenarios of demography and land use, sequestration rates will decrease in the future because biomass accumulation decreases with forest age and there is little agricultural land remaining to be abandoned. Due to high per-capita consumption and population density, carbon emissions of Puerto Rico have increased dramatically and exceeded carbon sequestration during the second half of the 20th century. Although Puerto Rico had the highest percent of reforestation for a tropical country, emissions during the period 1950–2000 were approximately 3.5 times higher than sequestration, and current annual emission is almost nine times the rate of sequestration. Additionally, while sequestration will decrease over the next six decades, current socioeconomic trends suggest increasing emissions unless there are significant changes in energy technology or consumption patterns. In conclusion, socioeconomic changes leading to urbanization and industrialization in tropical countries may promote high rates of carbon sequestration during the decades following land abandonment. Initial high rates of carbon sequestration can balance emissions of developing countries with low emission/GDP ratio. In Puerto Rico, the socioeconomic changes that promoted reforestation also promoted high-energy consumption, and resulted in a net increase in carbon emissions.

Variation in Susceptibility to Hurricane Damage as a Function of Storm Intensity in Puerto Rican Tree Species

Canham, Charles D.; Thompson, Jill; Zimmerman, Jess K.; Uriarte, Maria. 2010 Variation in Susceptibility to Hurricane Damage as a Function of Storm Intensity in Puerto Rican Tree Species. Biotropica, 42 (1). 87-94. 10.1111/j.1744-7429.2009.00545.x

One of the most significant challenges in developing a predictive understanding of the long-term effects of hurricanes on tropical forests is the development of quantitative models of the relationships between variation in storm intensity and the resulting severity of tree damage and mortality. There have been many comparative studies of interspecific variation in resistance of trees to wind damage based on aggregate responses to individual storms. We use a new approach, based on ordinal logistic regression, to fit quantitative models of the susceptibility of a tree species to different levels of damage across an explicit range of hurricane intensity. Our approach simultaneously estimates both the local intensity of the storm within a plot and the susceptibility to storm damage of different tree species within plots. Using the spatial variation of storm intensity embedded in two hurricanes (Hugo in 1989 and Georges in 1998) that struck the 16 ha Luquillo Forest Dynamics Plot in eastern Puerto Rico, we show that variation in susceptibility to storm damage is an important aspect of life history differentiation. Pioneers such as Cecropia schreberiana are highly susceptible to stem damage, while the late successional species Dacryodes excelsa suffered very little stem damage but significant crown damage. There was a surprisingly weak relationship between tree diameter and the susceptibility to damage for most of the 12 species examined. This may be due to the effects of repeated storms and trade winds on the architecture of trees and forest stands in this Puerto Rican subtropical wet forest.

Forest Regeneration in a chronosequence of tropical abandoned pastures: implications for restoration ecology

Forest Regeneration in a chronosequence of tropical abandoned pastures: implications for restoration ecology

During the mid-1900s, most of the island of Puerto Rico was deforested, but a shift in the economy from agriculture to small industry beginning in the 1950s resulted in the abandonment of agricultural lands and recovery of secondary forest. This unique history provides an excellent opportunity to study secondary forest succession and suggest strategies for tropical forest restoration. To determine the pattern of secondary succession, we describe the woody vegetation in 71 abandoned pastures and forest sites in four regions of Puerto Rico. The density, basal area, aboveground biomass, and species richness of the secondary forest sites were similar to those of the old growth forest sites (>80 yr) after approximately 40 years. The dominant species that colonized recently abandoned pastures occurred over a broad elevational range and are widespread in the neotropics. The species richness of Puerto Rican secondary forests recovered rapidly, but the species composition was quite different in comparison with old growth forest sites, suggesting that enrichment planting will be necessary to restore the original composition. Exotic species were some of the most abundant species in the secondary forest, but their long-term impact depended on life history characteristics of each species. These data demonstrate that one restoration strategy for tropical forest in abandoned pastures is simply to protect the areas from fire, and allow natural regeneration to produce secondary forest. This strategy will be most effective if remnant forest (i.e., seed sources) still exist in the landscape and soils have not been highly degraded. Patterns of forest recovery also suggest strategies for accelerating natural recovery by planting a suite of generalist species that are common in recently abandoned pastures in Puerto Rico and throughout much of the neotropics.

Nitrogen Immobilization by Decomposing Woody Debris and the Recovery of Tropical Wet Forest from Hurricane Damage

Nitrogen Immobilization by Decomposing Woody Debris and the Recovery of Tropical Wet Forest from Hurricane Damage
J. K. Zimmerman, W. M. Pulliam, D. J. Lodge, V. Quiñones-Orfila, N. Fetcher, S. Guzmán-Grajales, J. A. Parrotta, C. E. Asbury, L. R. Walker and R. B. Waide
Vol. 72, No. 3 (Apr., 1995), pp. 314-322

Following damage caused by Hurricane Hugo (September 1989) we monitored inorganic nitrogen availability in soil twice in 1990, leaf area index in 1991 and 1993, and litter production from 1990 through 1992 in subtropical wet forest of eastern Puerto Rico. Experimental removal of litter and woody debris generated by the hurricane (plus any standing stocks present before the hurricane) increased soil nitrogen availability and above-ground productivity by as much as 40% compared to unmanipulated control plots. These increases were similar to those created by quarterly fertilization with inorganic nutrients. Approximately 85% of hurricane-generated debris was woody debris >5 cm diameter. Thus, it appeared that woody debris stimulated nutrient immobilization, resulting in depression of soil nitrogen availability and productivity in control plots. This was further suggested by simulations of an ecosystem model (CENTURY) calibrated for our site that indicated that only the large wood component of hurricane-generated debris was of sufficiently low quality and of great enough mass to cause the observed effects on productivity. The model predicted that nutrient immobilization by decaying wood should suppress net primary productivity for 13 yr and total live biomass for almost 30 yr following the hurricane. Our findings emphasize the substantial influence that woody debris has upon nutrient cycling and productivity in forest ecosystems through its effects on the activity of decomposers. We suggest that the manner in which woody debris regulates ecosystem function in different forests is significantly affected by disturbance regime.

Acclimation of tropical tree species to hurricane disturbance: ontogenetic differences

Wen, S.Y., Fetcher, N. & Zimmerman, J.K. (2008) Acclimation of tropical tree
species to hurricane disturbance: ontogenetic differences. Tree Physiology,
28, 935–946.

We investigated acclimation responses of seedlings and saplings of the pioneer species Cecropia schreberiana Miq. and three non-pioneer species, Dacryodes excelsa Vahl, Prestoea acuminata (Willdenow) H.E. Moore var. montana (Graham) Henderson and Galeano, and Sloanea berteriana Choisy ex DC, following a hurricane disturbance in a lower montane wet forest in Puerto Rico. Measurements were made, shortly after passage of the hurricane, on leaves expanded before the hurricane (pre-hurricane leaves) and, at a later time, on recently matured leaves that developed after the hurricane (post-hurricane leaves) from both seedlings and saplings at sites that were severely damaged by the hurricane (disturbed sites) and at sites with little disturbance (undisturbed sites). Pre-hurricane leaves of the non-pioneer species had relatively low light-saturated photosynthetic rates (Amax) and stomatal conductance (gs); neither Amax nor gs responded greatly to the increase in irradiance that resulted from the disturbance, and there were few significant differences between seedlings and saplings. Pre-hurricane leaves of plants at undisturbed sites had low dark respiration rates per unit area (Rd) and light compensation points (LCP), whereas pre-hurricane leaves of plants at disturbed sites had significantly higher Rd and LCP. Post-hurricane leaves of plants at disturbed sites had significantly higher Amax and Rd than plants at undisturbed sites. Compared with seedlings, saplings had higher Amax and Rd and showed greater acclimation to the increase in irradiance that followed the disturbance. Post-hurricane leaves of the non-pioneer species had significantly lower Amax and were less responsive to changes in irradiance than the pioneer species C. schreberiana. Variation in Amax across light environments and stages was strongly related to differences in leaf mass per unit area (LMA), especially in the non-pioneer species. As indicated by Vcmax or Jmax per unit nitrogen, light acclimation of Amax was determined by leaf morphology (LMA) for the nonpioneer species and by both leaf morphology and leaf biochemistry for C. schreberiana. Ontogenetic changes in Amax were attributable to changes in leaf morphology. The ontogenetic component of variation in Amax across light environments and stages differed among species, ranging from 36 to 59% for the non-pioneer species (D. excelsa, 59.3%; P. acuminata var. montana, 44.7%; and S. berteriana, 36.3%) compared with only 17% in the pioneer species C. schreberiana.

Natural disturbance and human land use as determinants of tropical forest dynamics: results from a forest simulator

Uriarte, M., C. D. Canham, J. Thompson, J. K. Zimmerman,
L. Murphy, A. M. Sabat, N. Fetcher, and B. L.
Haines. 2009. Natural disturbance and human land
use as determinants of tropical forest dynamics:
results from a forest simulator. Ecological Monographs

Forests are often subject to multiple, compounded disturbances, representing both natural and human-induced processes. Predicting forest dynamics requires that we consider how these disturbances interact to affect species demography. Here we present results of an individual-based, spatially explicit forest simulator that we developed to analyze the compounded effects of hurricane disturbance and land use legacies on the dynamics of a subtropical forest. We used data from the 16-ha Luquillo Forest Dynamics Plot in Puerto Rico, together with a reconstruction of historical wind damage, to parameterize the simulator. We used the model to ask two questions. (1) What are the implications of variation in hurricane frequency and severity for the long-term dynamics of forest composition, diversity, and structure? Both storm severity and frequency had striking effects on forest dynamics, composition, and structure. The periodicity of disturbance also played an important role, with periods of high hurricane activity fostering the establishment of species that may become rare in the absence of severe storms and quiescent periods allowing these species to reach reproductive size. Species responses to hurricane disturbance could not be predicted from their life history attributes. However, species perceived to be primary forest species exhibited low temporal variation in abundance through the simulations. (2) How do hurricanes and legacies from human land use interact to determine community structure and composition? Our results suggest that, over time, regardless of the storm regime, land use legacies will become less apparent but will lead to a forest community that contains a mixture of secondary and primary forest species formerly dominant in areas of different land use. In the long term, hurricane disturbance generated two communities with slightly greater similarity than those not subject to storms. Thus, the inclusion of hurricane disturbance does not alter the prediction that land use legacies in this tropical forest will diminish over time. Our simulations also highlight the contingent effects of human legacies on subsequent community dynamics, including the response to hurricane disturbance, therefore supporting the notion that compounded disturbances can interact in ways that cannot be predicted by the study of single disturbances. The widespread importance of land use as a large-scale disturbance makes it imperative that it be addressed as a fundamental ecological process.

Land Use History, Environment, and Tree Composition in a Tropical Forest

Thompson, Jill; Brokaw, Nicholas; Zimmerman, Jess K.; Waide, Robert B.; Everham, Edwin M. III; Lodge, D. Jean; Taylor, Charlotte M.; Garcia-Montiel, Diana; Fluet, Marcheterre 2002. Land use history, environment, and tree composition in a tropical forest. Ecological applications. Vol. 12, no. 5 (2002): pages 1344-1363.

The effects of historical land use on tropical forest must be examined to understand present forest characteristics and to plan conservation strategies. We compared the effects of past land use, topography, soil type, and other environmental variables on tree species composition in a subtropical wet forest in the Luquillo Mountains, Puerto Rico. The study involved stems > 10 cm diameter measured at 130 cm above the ground, within the 16-ha Luquillo Forest Dynamics Plot (LFDP), and represents the forest at the time Hurricane Hugo struck in 1989. Topography in the plot is rugged, and soils are variable. Historical documents and local residents described past land uses such as clear-felling and selective logging followed by farming, fruit and coffee production, and timber stand improvement in the forest area that now includes the LFDP. These uses ceased 40-60 yr before the study, but their impacts could be differentiated by percent canopy cover seen in aerial photographs from 1936. Using these photographs, we defined four historic cover classes within the LFDP. These ranged from cover class 1, the least tree-covered area in 1936, to cover class 4, with the least intensive historic land use (selective logging and timber stand improvement). In 1989, cover class 1 had the lowest stem density and proportion of large stems, whereas cover class 4 had the highest basal area, species richness, and number of rare and endemic species. Ordination of tree species composition (89 species, 13 167 stems) produced arrays that primarily corresponded to the four cover classes (i.e., historic land uses). The ordination arrays corresponded secondarily to soil characteristics and topography. Natural disturbances (hurricanes, landslides, and local treefalls) affected tree composition, but these effects did not correlate with the major patterns of species distributions on the plot. Thus, it appears that forest development and natural disturbance have not masked the effects of historical land use in this tropical forest, and that past land use was the major influence on the patterns of tree composition in the plot in 1989. The least disturbed stand harbors more rare and endemic species, and such stands should be protected.

Plant responses to simulated hurricane impacts in a subtropical wet forest, Puerto Rico

Shiels, Aaron B.; Zimmerman, Jess K.; García-Montiel, Diana C.; Jonckheere, Inge; Holm, Jennifer; Horton, David; Brokaw, Nicholas. 2010. Plant responses to simulated hurricane impacts in a subtropical wet forest, Puerto Rico. Journal of Ecology. doi: 10.1111/j.1365-2745.2010.01646.x.

1. We simulated two key components of severe hurricane disturbance, canopy openness and detritus deposition, to determine the independent and interactive effects of these components on woody plant recruitment and forest structure. 2. We increased canopy openness by trimming branches and added or subtracted canopy detritus in a factorial design. Plant responses were measured during the 4-year study, which followed at least 1 year of pre-manipulation monitoring. 3. The physical conditions of canopy openness and detritus deposition in our experiment resembled the responses to Hurricane Hugo, a severe category 4 hurricane that struck this forest in 1989. 4. Canopy detritus deposition killed existing woody seedlings and provided a mechanical barrier that suppressed seedling recruitment. The increase in understorey light caused by canopy trimming stimulated germination from the seed bank and increased seedling recruitment and density of pioneer species several hundred-fold when hurricane debris was absent. Many significant interactions between trimming and detritus deposition were evident from the manner in which seedling density, recruitment and mortality changed over time, and subsequently influenced the composition of woody stems (individuals ‡ 1 cmd.b.h.). 5. When the canopy was trimmed, stem densities increased> 2-fold and rates of recruitment into the stem size class increased> 25-fold. Trimming had no significant effect on stem mortality. The two dominant species that flourished following canopy trimming were the pioneer species Cecropia schreberiana and Psychotria berteriana. Deposition of canopy detritus had little effect on stems, although basal area increased slightly when detritus was added. There were no evident effects of the interactions between canopy trimming and detritus deposition on stems. 6. Synthesis. The separate and interactive effects of canopy openness and detritus deposition result in variable short-term trajectories of forest recovery. However, the short interval of increased canopy openness due to hurricane impacts and its influence on the recruitment of pioneer trees is the dominant factor that drives short-termrecovery and may alter long-term structure and composition of the forest.

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

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.
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