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Timing the origin and expansion of the Mexican tropical dry forest.

Macroevolution examines the temporal patterns of biological diversity in deep time. When combined with biogeography, it can provide unique information about the historical changes in the distribution of communities and biomes. Here I document temporal and spatial changes of diversity in the genus Bursera and relate them to the origin and expansion of the tropical dry forests of Mexico. Bursera is very old, highly adapted to warm dry conditions, and a dominant member of the Mexican tropical dry forest. These characteristics make it a useful indicator of the history of this vegetation. I used a time-calibrated phylogeny to estimate Bursera's diversification rate at different times over the last 60 million years. I also reconstructed the geographic center and time of origin of all species and nodes from information on current distributions. Results show that between 30 and 20 million years ago, Bursera began a relatively rapid diversification. This suggests that conditions were favorable for its radiation and thus, very probably for the establishment of the dry forest as well. The oldest lineages diverged mostly in Western Mexico, whereas the more recent lineages diverged in the south-central part of the country. This suggests that the tropical dry forest probably first established in the west and then expanded south and east. The timing of the radiations in these areas corresponds to that suggested for formations of the mountainous systems in Western and Central Mexico, which have been previously recognized as critical for the persistence of the Mexican dry forest.

Biological Evolution↗

Increased pollen flow counteracts fragmentation in a tropical dry forest: an example from Swietenia humilis Zuccarini.

Habitat destruction and the resultant fragmentation of the remaining forest are a common phenomenon in the tropics. Most investigations emphasize the potential dangers of fragmentation in isolating patches of forest and exposing populations to loss of species diversity through founder effects, genetic drift, inbreeding, and restricted gene flow. However, a limited number of studies have shown that gene flow may be extensive in tropical trees, suggesting that it may occur between forest fragments and also "isolated" remnant trees. There is an urgent need to quantify pollen flow within and between forest fragments to test the veracity of such views and determine the genetic value of such fragments for in situ conservation. Microsatellite markers are used to genotype individuals of Swietenia humilis from a highly fragmented forest mosaic to directly quantify pollen-mediated gene flow. Distances of pollen flow more than 10 times greater than previously reported were detected. Our results show that some tropical angiosperm tree species may be much more adaptable and resilient to habitat destruction and fragmentation than previously considered. The description of many remnant trees as isolated or "living dead" may be more a conditioning of human perception than a true reflection of their potential conservation value.

Ecology↗

Nutrient additions to a tropical rain forest drive substantial soil carbon dioxide losses to the atmosphere.

Terrestrial biosphere-atmosphere carbon dioxide (CO(2)) exchange is dominated by tropical forests, where photosynthetic carbon (C) uptake is thought to be phosphorus (P)-limited. In P-poor tropical forests, P may also limit organic matter decomposition and soil C losses. We conducted a field-fertilization experiment to show that P fertilization stimulates soil respiration in a lowland tropical rain forest in Costa Rica. In the early wet season, when soluble organic matter inputs to soil are high, P fertilization drove large increases in soil respiration. Although the P-stimulated increase in soil respiration was largely confined to the dry-to-wet season transition, the seasonal increase was sufficient to drive an 18% annual increase in CO(2) efflux from the P-fertilized plots. Nitrogen (N) fertilization caused similar responses, and the net increases in soil respiration in response to the additions of N and P approached annual soil C fluxes in mid-latitude forests. Human activities are altering natural patterns of tropical soil N and P availability by land conversion and enhanced atmospheric deposition. Although our data suggest that the mechanisms driving the observed respiratory responses to increased N and P may be different, the large CO(2) losses stimulated by N and P fertilization suggest that knowledge of such patterns and their effects on soil CO(2) efflux is critical for understanding the role of tropical forests in a rapidly changing global C cycle.

Atmosphere↗

Insights on linking forests, trees, and people from the air, on the ground, and in the laboratory.

Governing natural resources sustainably is a continuing struggle. Major debates occur over what types of policy "interventions" best protect forests, with choices of property and land tenure systems being central issues. Herein, we provide an overview of findings from a long-term interdisciplinary, multiscale, international research program that analyzes the institutional factors affecting forests managed under a variety of tenure arrangements. This program analyzes satellite images, conducts social-ecological measurements on the ground, and tests the impact of structural variables on human decisions in experimental laboratories. Satellite images track the landscape dimensions of forest-cover change within different management regimes over time. On-the-ground social-ecological studies examine relationships between forest conditions and types of institutions. Behavioral studies under controlled laboratory conditions enhance our understanding of explicit changes in structure that affect relevant human decisions. Evidence from all three research methods challenges the presumption that a single governance arrangement will control overharvesting in all settings. When users are genuinely engaged in decisions regarding rules affecting their use, the likelihood of them following the rules and monitoring others is much greater than when an authority simply imposes rules. Our results support a frontier of research on the most effective institutional and tenure arrangements for protecting forests. They move the debate beyond the boundaries of protected areas into larger landscapes where government, community, and comanaged protected areas are embedded and help us understand when and why deforestation and regrowth occur in specific regions within these larger landscapes.

Community Participation↗

Rapid decay of tree-community composition in Amazonian forest fragments.

Forest fragmentation is considered a greater threat to vertebrates than to tree communities because individual trees are typically long-lived and require only small areas for survival. Here we show that forest fragmentation provokes surprisingly rapid and profound alterations in Amazonian tree-community composition. Results were derived from a 22-year study of exceptionally diverse tree communities in 40 1-ha plots in fragmented and intact forests, which were sampled repeatedly before and after fragment isolation. Within these plots, trajectories of change in abundance were assessed for 267 genera and 1,162 tree species. Abrupt shifts in floristic composition were driven by sharply accelerated tree mortality and recruitment within approximately 100 m of fragment margins, causing rapid species turnover and population declines or local extinctions of many large-seeded, slow-growing, and old-growth taxa; a striking increase in a smaller set of disturbance-adapted and abiotically dispersed species; and significant shifts in tree size distributions. Even among old-growth trees, species composition in fragments is being restructured substantially, with subcanopy species that rely on animal seed-dispersers and have obligate outbreeding being the most strongly disadvantaged. These diverse changes in tree communities are likely to have wide-ranging impacts on forest architecture, canopy-gap dynamics, plant-animal interactions, and forest carbon storage.

Ecosystem↗

Guatemalan forest synthesis after Pleistocene aridity.

Sediments from two lakes in the Peten Department, Guatemala, provide palynological evidence from Central America of late Pleistocene aridity and subsequent synthesis of mesic forests. Late Glacial vegetation consisted of marsh, savanna, and juniper scrub. An early Holocene temperate forest preceded a mesic tropical forest with Brosimum (ramon). Thus "primeval" rain forests of Guatemala are no older than 10,000 to 11,000 years and are considerably younger in the Peten due to Mayan disturbances. Among dated Neotropical sites, the Peten has the most mesic vegetation yet shown to have supplanted xeric vegetation present during the Pleistocene. The arid late Glacial-humid early Holocene transition appears to have been pantropical in the lowlands. The Peten was not a Pleistocene refugium for mesophytic taxa, as has been suggested. Thus genesis of extant rain forests in northern Central America and southern Mexico remains unexplained.

Journal Article↗

Diversity in soil fungi from undisturbed and disturbed Celtis tala and Scutia buxifolia forests in the eastern Buenos Aires province (Argentina).

The rhizospheric soil microfungi from a native forest (undisturbed and disturbed) were studied using soil dilution plate and soil washing methods. Fungi were isolated using slightly acid and alkaline culture media. 54 taxa were isolated: 49 from undisturbed forest soil and 37 from disturbed forest soil. Acremonium sp., Aspergillus ustus, Coemansia pectinata, Doratomyces stemonitis, Fusarium solani, F. oxysporum, Gliocladium roseum, Humicola fusco-atra, Mortierella sp., Penicillium lilacinum, Trichoderma harzianum, and T koningii, showed the highest frequency, in both, undisturbed and disturbed forests. In undisturbed soil forest the biodiversity index was 3.97 whereas in disturbed ones was 3.89.

Algorithms↗

Isotopic discrimination during litter decomposition and delta13C and delta15N soil profiles in a young artificial stand and in an old floodplain forest.

In the present study, rates of litter decomposition and microbial biomass nitrogen were monitored over an 8-month period in a young broadleaf plantation (18 y) and in an old floodplain forest. Moreover, delta13C and delta15N temporal variations within soil profiles were evaluated at both sites. Rates of litter decomposition were higher in spring and autumn than in summer, in both forests. At the end of the observation period the percentage of original litter remaining was not statistically different between the young and the old forest and accounted for 60-70% of the original amount. Microbial biomass nitrogen in the remaining litter and the percentage of litter mass lost during decomposition were positively correlated. The difference in litter quality affected the decomposition rate and also the changes in carbon isotopic composition during the decomposition process. In contrast, 15N isotopic signatures showed a similar trend in the litter of the two forests irrespective of the litter quality. Although delta13Csoil and delta15Nsoil showed considerable temporal variation they increased with depth in the soils of both sites but their seasonal changes did not reflect those of the decomposing litter. Within the same soil horizon, both delta13C and delta15N showed similar seasonal trends in the soils of the two forests, suggesting the involvement of environmental factors acting at regional level, such as soil temperature and rainfall variations, in regulating seasonal delta13C and delta15N soil variations.

Carbon Isotopes↗

Estimation of vectorial capacity of Anopheles dirus (Diptera: Culicidae) in a forest-fringed village of Assam (India).

To estimate the vectorial capacity of Anopheles dirus, the main vector of forest malaria in the northeastern region of India, in order to gain an understanding of entomological factors related to malaria transmission in forest-fringe areas of Assam, India, an isolated village in the tropical rain forest-fringed area in the district of Dibrugarh, Assam, under the influence of An. dirus alone was studied. Data on various entomological variables required for computation of the vectorial capacity were generated in each month from June 1999 to May 2000 in the field using standard techniques. Malaria prevalence was also studied during the same period in the study village and correlated with the estimated vectorial capacity of An. dirus. Vectorial capacity of An. dirus was highest, 0.779 for Plasmodium vivax (Pv) and 0.649 for Plasmodium falciparum (Pf), during the hot-monsoon season (June-September) and decreased to 0.08 (Pv) and 0.07(Pf) in the temperate postmonsoon season (October-November) before attaining zero values in the cool-dry season (December-February). With increasing temperature in the temperate premonsoon season (March-May), vectorial capacity recorded was 0.119 and 0.82 for Pv and Pf, respectively. Significant positive correlation was seen between the estimated vectorial capacity of An. dirus and the number of new Pf (r = 0.86, p < 0.001) and Pv (r = 0.69, p < 0.02) cases in the study village in different months. Thus, this study highlights the pattern of malaria transmission by An. dirus in a forest-fringe area of Assam that begins in March, peaks in July/August, subsides by November, and remains interrupted between December and February. Measures for controlling malaria in forest-fringe areas should be scheduled accordingly.

Animals↗

Physiological ecology of Mesozoic polar forests in a high CO2 environment.

Fossils show that coniferous forests extended into polar regions during the Mesozoic, a time when models and independent paleo-CO2 indicators suggest that the atmospheric CO2 concentration was at least double that of the present day. Consequently, such polar forests would have experienced high CO2 interacting with an extreme variation in light. Here we describe an experiment investigating this plant-environment interaction for extant tree species that were important components of polar forests, and give results from the first year of treatment. Specifically, we tested the hypotheses that growth in elevated CO2 (1) stimulates photosynthesis; (2) reduces photoinhibition during the polar summer; and (3) reduces respiration of above- and below-ground plant organs. Our results indicate that CO2 fertilization generally does not affect photosynthesis under continuous daylight characteristic of the polar summer but does increase it when the period of illumination is shorter. Growth in elevated CO2 did not alter the potential for photoinhibition. CO2 enrichment significantly reduced leaf and root respiration rates by 50 and 25 %, respectively, in a range of evergreen taxa. Incorporating these observed CO2 effects into numerical simulations using a process-based model of coniferous forest growth indicates that a high paleo-CO2 concentration would have increased the productivity of Cretaceous conifer forests in northern Alaska. This results from decreased respiratory costs that more than compensate for the absence of high CO2-high temperature interactions during the polar summer. The longer-term effects of CO2 enrichment on seasonal changes in the above- and below-ground carbon balance of trees are discussed.

Atmosphere↗

Effect of forest type on the distribution of Lutzomyia shannoni (Diptera: Psychodidae) and vesicular stomatitis virus on Ossabaw Island, Georgia.

We studied the effects of three forest types on multiple factors that are believed to influence the transmission of the New Jersey serotype of vesicular stomatitis (VSNJ) virus on Ossabaw Island, GA. These factors included availability of tree hole diurnal resting habitat for the presumed sand fly vector, Lutzomyia shannoni Dyar; relative abundance of L. shannoni; prevalence of VSNJ virus infection in sand flies; and prevalence of VSNJ virus antibodies in wild swine. Tree hole availability, sand fly abundance, and antibody prevalence in swine were significantly greater in maritime live oak forest than in other forest types. A single isolate of VSNJ virus was obtained from sand flies collected in maritime live oak forest. These data indicate that the relative abundance of adult L. shannoni is influenced significantly by the availability of tree holes and that VSNJ virus infection in wild swine is linked to forest type and is greatest in areas capable of supporting abundant populations of L. shannoni.

Animals↗

An analysis of sap flow in mountain ash (Eucalyptus regnans) forests of different age.

Diurnal measurements of sap velocity were made in 50-, 90-, 150- and 230-year-old mountain ash (Eucalyptus regnans F. Muell.) forests in the North Maroondah catchment (southeast Australia) over the periods January 8, 1990 to April 4, 1990 and October 29, 1990 to April 16, 1991. Over the two periods, daily mean sap velocities for the four forests, in order of increasing age, were 11.5, 11.4, 9.9 and 11.8 cm h(-1) respectively. Daily mean sap velocity did not differ significantly among the 50-, 90- and 230-year-old plots. However, in the 150-year-old trees it was significantly smaller by an average of 14%. Sap velocity varied diurnally and also between positions within individual trees and among trees both within and between stands. Despite this variability, the sampling intensity and duration were sufficient to establish that behavior was highly correlated among individuals within plots. There was a significant decline with age in the overstory sapwood conducting area of these forests. In order of increasing age, the values were 6.7, 6.1, 4.2 and 4.0 m(-2) ha(-1), respectively. When combined with daily mean sap velocity, these data allowed the calculation of overstory water use. Over the experimental period, water use of the overstory decreased with age ranging, on average, from 1.86 mm day(-1) for the 50-year-old plot to 0.81 mm day(-1) for the 230-year-old plot. Mean daily water use for the two intermediate-aged forests was 1.67 and 1.00 mm day(-1), respectively. Annual water use decreased with forest age from 679 mm for the 50-year-old stand to 296 mm for the 230-year-old stand. This difference corresponds to 3.8 x 10(3) m(3) ha(-1). The annual water use of the intermediate-aged stands was 610 and 365 mm for the 90- and 150-year-old stands, respectively.

Journal Article↗

A physiological basis for biosphere-atmosphere interactions in the boreal forest: an overview.

Interdisciplinary field experiments for global change research are large, intensive efforts that study the controls on fluxes of carbon, water, trace gases, and energy between terrestrial ecosystems and the atmosphere at a range of spatial scales. Forest ecophysiology can make significant contributions to such efforts by measuring, interpreting, and modeling these fluxes for the individual components of forest ecosystems and then integrating the results into holistic ecosystem process models. The Boreal Ecosystem-Atmosphere Study (BOREAS) was undertaken because of the importance of the boreal forest biome to various global change issues. The study was conducted from 1993 to 1996 at sites in Saskatchewan and Manitoba, Canada. Results have shown that physiological processes of plants in the boreal forest can have large-scale consequences. For example, the composition of tree species strongly influences flux rates, with deciduous species having much higher carbon and water fluxes than coniferous species. Additionally, physiological limitations to transpiration in boreal conifers, even when soil water is abundant, reduces latent heat flux and increases sensible heat flux over large regions. This physiological control of transpiration can increase the depth of the atmospheric boundary layer on warm spring days to a level similar to that found in desert biomes. This special issue features 10 articles that address various aspects of the physiological basis of biosphere-atmosphere interactions in the boreal forest. The articles emphasize the environmental controls on water flux, carbon flux, and ecosystem productivity.

Journal Article↗

Root mass, net primary production and turnover in aspen, jack pine and black spruce forests in Saskatchewan and Manitoba, Canada.

Root biomass, net primary production and turnover were studied in aspen, jack pine and black spruce forests in two contrasting climates. The climate of the Southern Study Area (SSA) near Prince Albert, Saskatchewan is warmer and drier in the summer and milder in the winter than the Northern Study Area (NSA) near Thompson, Manitoba, Canada. Ingrowth soil cores and minirhizotrons were used to quantify fine root net primary production (NPPFR). Average daily fine root growth (m m(-2) day(-1)) was positively correlated with soil temperature at 10-cm depth (r(2) = 0.83-0.93) for all three species, with black spruce showing the strongest temperature effect. At both study areas, fine root biomass (measured from soil cores) and fine root length (measured from minirhizotrons) were less for jack pine than for the other two species. Except for the aspen stands, estimates of NPPFR from minirhizotrons were significantly greater than estimates from ingrowth cores. The core method underestimated NPPFR because it does not account for simultaneous fine root growth and mortality. Minirhizotron NPPFR estimates ranged from 59 g m(-2) year(-1) for aspen stands at SSA to 235 g m(-2) year(-1) for black spruce at NSA. The ratio of NPPFR to total detritus production (aboveground litterfall + NPPFR) was greater for evergreen forests than for deciduous forests, suggesting that carbon allocation patterns differ between boreal evergreen and deciduous forests. In all stands, NPPFR consistently exceeded annual fine root turnover and the differences were larger for stands in the NSA than for stands in the SSA, whereas the difference between study areas was only significant for black spruce. The imbalance between NPPFR and fine root turnover is sufficient to explain the net accumulation of carbon in boreal forest soils.

Journal Article↗

Simulated nitrogen cycling response to elevated CO(2) in Pinus taeda and mixed deciduous forests.

Interactions between elevated CO(2) and N cycling were explored with a nutrient cycling model (NuCM, Johnson et al. 1993, 1995) for a Pinus taeda L. site at Duke University, North Carolina, and a mixed deciduous site at Walker Branch, Tennessee. The simulations tested whether N limitation would prevent growth increases in response to elevated CO(2), and whether growth responses to CO(2) in N-limited systems could be facilitated by increasing the biomass/N ratio (reducing N concentration) or increasing litter N mineralization, or both. Nitrogen limitation precluded additional growth when target growth rates and litterfall were increased (simulating potential response to elevated CO(2)) at the Duke University site. At the Walker Branch site, increasing target growth and litterfall caused a 7% increase in growth. Reducing foliar N concentrations reduced growth because of N limitation created by reduced litter quality (C:N ratio), reduced decomposition and increased N accumulation on the forest floor. These effects were most pronounced at the Duke University site, because the forest floor N turnover rate was lower than at the Walker Branch site. Reducing wood N concentration allowed prolonged increases in growth because of greater biomass/N; however, N uptake was reduced, allowing greater N immobilization on the forest floor and in soil. Increased N mineralization caused increased growth at the Duke University site, but not at the Walker Branch site. These simulations pose the counterintuitive hypothesis that increased biogeochemical cycling of N (as a result of increased litterfall N) causes reduced growth in an N-limited system because of increased accumulations of N on the forest floor and in soil. Translocation of N from senescing leaves before litterfall mitigates this response by allowing the trees to retain a greater proportion of N taken up rather than recycle it back to the forest floor and soil where it can be immobilized. Eliminating N translocation at Walker Branch changed the direction as well as the magnitude of the responses in three of the four scenarios simulated. Because the NuCM model does not currently allow translocation in coniferous species, the effects of translocation on N cycling in the Duke University simulations are not known.

Journal Article↗

Modeling daily gas exchange of a Douglas-fir forest: comparison of three stomatal conductance models with and without a soil water stress function.

Modeling stomatal conductance is a key element in predicting tree growth and water use at the stand scale. We compared three commonly used models of stomatal conductance, the Jarvis-Loustau, Ball-Berry and Leuning models, for their suitability for incorporating soil water stress into their formulation, and for their performance in modeling forest ecosystem fluxes. We optimized the parameters of each of the three models with sap flow and soil water content data. The optimized Ball-Berry model showed clear relationships with air temperature and soil water content, whereas the optimized Leuning and Jarvis-Loustau models only showed a relationship with soil water content. We conclude that use of relative humidity instead of vapor pressure deficit, as in the Ball-Berry model, is not suitable for modeling daily gas exchange in Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) in the Speulderbos forest near the village of Garderen, The Netherlands. Based on the calculated responses to soil water content, we linked a model of forest growth, FORGRO, with a model of soil water, SWIF, to obtain a forest water-balance model that satisfactorily simulated carbon and water (transpiration) fluxes and soil water contents in the Douglas-fir forest for 1995.

Journal Article↗

Carbon exchange of an old-growth eastern hemlock (Tsuga canadensis) forest in central New England.

Carbon (C) exchange of an approximately 200-year-old eastern hemlock (Tsuga canadensis L.) forest in central Massachusetts, USA, was estimated from mid-October 2000 through October 2001 based on eddy covariance measurements and statistical modeling from microclimatic data. Measurements were made in 68% of the hours during the year of study, with > 50% coverage in all months except December and August. Data were filtered by wind direction and atmospheric turbulence to remove invalid measurements. Analysis of filtered data showed that photosynthetically active radiation (PAR) was significant in predicting C exchange, except during the winter. Daily minimum air temperature affected C exchange in autumn and winter, whereas time of day, water vapor pressure deficit and air temperature had significant effects on C storage in spring, summer and fall. Most C storage in the stand occurred in April through July and in October 2001, with maximum rates in April and May. Persistent cold weather prevented C storage in December through March. In early spring 2001, C uptake was sensitive to nocturnal frost: daily minimum air temperatures below 0 degrees C reduced C fixation, and minima below -5 degrees C caused its virtual cessation. Soil temperature was a poor predictor of C balance during this period. In August, high soil and air temperatures (averaging 16.7 and 21.1 degrees C, respectively) drove high ecosystem respiration, which approximately balanced C uptake. These patterns show potential for stimulated C storage in hemlock forests in a warmer climate with fewer spring and autumn frosts, but reduced C storage during warmer summers. Estimated annual C storage was 3.0 Mg ha(-1), which is higher than for younger coniferous and deciduous forests during earlier years in the northeastern USA. Long-term data are needed to determine if the estimated high C storage in this hemlock forest is a result of interannual climate variation or an effect of forest composition.

Biomass↗

Relationships between net photosynthesis and foliar nitrogen concentrations in a loblolly pine forest ecosystem grown in elevated atmospheric carbon dioxide.

We examined the effects of elevated carbon dioxide concentration ([CO2]) on the relationship between light-saturated net photosynthesis (A(sat)) and area-based foliar nitrogen (N) concentration (N(a)) in the canopy of the Duke Forest FACE experiment. Measurements of A(sat) and N(a) were made on two tree species growing in the forest overstory and four tree species growing in the forest understory, in ambient and elevated [CO2] FACE rings, during early and late summer of 1999, 2001 and 2002, corresponding to years three, five and six of CO2 treatment. When measured at the growth [CO2], net photosynthetic rates of each species examined in the forest overstory and understory were stimulated by elevated [CO2] at each measurement date. We found no effect of elevated [CO2] on N(a) in any of the species. The slope of the A(sat)-N relationship was 81% greater in elevated [CO2] than in ambient [CO2] when averaged across all sample dates, reflecting a differential CO2 effect on photosynthesis at the top and bottom of the canopy. We compared A(sat)-N relationships in trees grown in ambient and elevated [CO2] at two common CO2 concentrations, during late summer 2001 and both early and late 2002, to determine if the stimulatory effect of elevated [CO2] on photosynthesis diminishes over time. At all three sample times, neither the slopes nor the y-intercepts of the A(sat)-N relationships of trees grown in ambient or elevated [CO2] differed when measured at common CO2 concentrations, indicating that the responses of photosynthesis to long-term elevated [CO2] did not differ from the responses to a short-term increase in [CO2]. This finding, together with the observation that N(a) was unaffected by growth in elevated [CO2], indicates that these overstory and understory trees growing at the Duke Forest FACE experiment continue to show a strong stimulation of photosynthesis by elevated [CO2].

Atmosphere↗