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Calibration and assessment of seasonal changes in leaf area index of a tropical dry forest in different stages of succession.

A simple measure of the amount of foliage present in a forest is leaf area index (LAI; the amount of foliage per unit ground surface area), which can be determined by optical estimation (gap fraction method) with an instrument such as the Li-Cor LAI-2000 Plant Canopy Analyzer. However, optical instruments such as the LAI-2000 cannot directly differentiate between foliage and woody components of the canopy. Studies investigating LAI and its calibration (extracting foliar LAI from optical estimates) in tropical forests are rare. We calibrated optical estimates of LAI from the LAI-2000 with leaf litter data for a tropical dry forest. We also developed a robust method for determining LAI from leaf litter data in a tropical dry forest environment. We found that, depending on the successional stage of the canopy and the season, the LAI-2000 may underestimate LAI by 17% to over 40%. In the dry season, the instrument overestimated LAI by the contribution of the woody area index. Examination of the seasonal variation in LAI for three successional stages in a tropical dry forest indicated differences in timing of leaf fall according to successional stage and functional group (i.e., lianas and trees). We conclude that when calculating LAI from optical estimates, it is necessary to account for the differences between values obtained from optical and semi-direct techniques. In addition, to calculate LAI from litter collected in traps, specific leaf area must be calculated for each species rather than from a mean value for multiple species.

Calibration↗

Foliar temperature tolerance of temperate and tropical evergreen rain forest trees of Australia.

Australian rain forests extend from tropical climates in the north to temperate climates in the south, providing an opportunity to investigate physiological responses to temperature of both temperate and tropical species within the same forest type. Eight, rain forest canopy tree species were selected to cover the 33 degrees latitudinal range of rain forests in eastern Australia. Temperature tolerance was measured in 6-year-old plants grown in a common environment, by exposing leaves to a series of high temperatures during late summer and a series of freezing temperatures during midwinter. Damage was evaluated based on chlorophyll fluorescence measurements made 2 h after exposure and by visual assessment of leaf damage made a week after exposure. Leaves of the tropical species were more heat tolerant and less frost tolerant than leaves of the temperate species, which is consistent with their climate distributions. In contrast, the temperature tolerance of the photosynthetic apparatus was unrelated to climate in a species' native habitat. However, the tropical species underwent significant photoinhibition during winter. All species maintained the integrity of the photosynthetic apparatus and avoided tissue damage over a similar span of temperatures (about 60 degrees C), reflecting the similar annual temperature ranges in Australia's temperate and tropical rain forests. Chlorophyll fluorescence measurements and visual assessment of leaf damage provided different estimates of the absolute and relative temperature tolerances of the species, thus emphasizing the importance of a direct assessment of tissue damage for determining a species' temperature tolerance.

Australia↗

Assessment of oak forest condition based on leaf biochemical variables and chlorophyll fluorescence.

Pedunculate oak forests (Quercus robur L.) in the Ticino Regional Park, Italy, are declining as a result of insect attacks, summer droughts and air pollution. The assessment and monitoring of forest condition can provide a basis for managing and conserving forest ecosystems and thereby avoid loss of valuable natural resources. Currently, most forest assessments are limited to ground-based visual evaluations that are local and subjective. It is therefore difficult to compare data collected by different crews or to define reliable trends over years. We examined vegetation variables that can be quantitatively estimated by remote observations and, thus, are suitable for objective monitoring over extended forested areas. We found that total chlorophyll (Chl) concentration is the most suitable variable for assessing pedunculate oak decline. It is highly correlated with visual assessments of discoloration. Furthermore, Chl concentration can be accurately estimated from leaf optical properties, making it feasible to map Chl concentration at the canopy level from satellite and airborne remote observations.

Chlorophyll↗

Contrasting seasonal leaf habits of canopy trees between tropical dry-deciduous and evergreen forests in Thailand.

We compared differences in leaf properties, leaf gas exchange and photochemical properties between drought-deciduous and evergreen trees in tropical dry forests, where soil nutrients differed but rainfall was similar. Three canopy trees (Shorea siamensis Miq., Xylia xylocarpa (Roxb.) W. Theob. and Vitex peduncularis Wall. ex Schauer) in a drought-deciduous forest and a canopy tree (Hopea ferrea Lanessan) in an evergreen forest were selected. Soil nutrient availability is lower in the evergreen forest than in the deciduous forest. Compared with the evergreen tree, the deciduous trees had shorter leaf life spans, lower leaf masses per area, higher leaf mass-based nitrogen (N) contents, higher leaf mass-based photosynthetic rates (mass-based P(n)), higher leaf N-based P(n), higher daily maximum stomatal conductance (g(s)) and wider conduits in wood xylem. Mass-based P(n) decreased from the wet to the dry season for all species. Following onset of the dry season, daily maximum g(s) and sensitivity of g(s) to leaf-to-air vapor pressure deficit remained relatively unchanged in the deciduous trees, whereas both properties decreased in the evergreen tree during the dry season. Photochemical capacity and non-photochemical quenching (NPQ) of photosystem II (PSII) also remained relatively unchanged in the deciduous trees even after the onset of the dry season. In contrast, photochemical capacity decreased and NPQ increased in the evergreen tree during the dry season, indicating that the leaves coped with prolonged drought by down-regulating PSII. Thus, the drought-avoidant deciduous species were characterized by high N allocation for leaf carbon assimilation, high water use and photoinhibition avoidance, whereas the drought-tolerant evergreen was characterized by low N allocation for leaf carbon assimilation, conservative water use and photoinhibition tolerance.

Biomass↗

Vertical variation in canopy structure and CO(2) exchange of oak-maple forests: influence of ozone, nitrogen, and other factors on simulated canopy carbon gain.

Stand-level and physiological measurements were made for oak and maple species common in Wisconsin forests. Scaling relationships were identified to allow the development of a model for estimating net carbon exchange at the levels of a leaf, canopy stratum, and whole canopy. Functional relationships were determined between tissue gas exchange rates and perceived controlling variables. Vertical variation in leaf properties and in the distribution of foliage by weight, area, and species were characterized for several closed canopy forests. Forest canopies were divided into four horizontal strata to develop predictive models for canopy gas exchange. Leaf and canopy layer carbon dioxide exchange rates were predicted using leaf nitrogen concentration, leaf mass per area, ozone exposure, predawn leaf water potential, photosynthetically active radiation, and vapor pressure deficit as driving variables. Direct measurements of leaf gas exchange were used to validate the components (subroutines) of the model. Net carbon dioxide exchange was simulated for canopy layers at 5-min intervals over a diurnal time course. Simulations of canopy CO(2) exchange were made for a 30-m tall, mixed oak-maple forest under hypothetical ambient and greater-than-ambient ozone pollution regimes. Daily canopy net CO(2) exchange was predicted for seven forest stands and compared with estimates of aboveground net primary production, N availability, leaf area index, and canopy N.

Journal Article↗

Considerations in modeling change in temperate forest nitrogen cycles.

The general features of nitrogen (N) cycles in temperate forests and the important processes to consider when modeling change in these cycles include atmospheric inputs, N fixation, litter (root and aboveground) transfers and decomposition, soil processes, N uptake and effects on productivity and litter quality, and N outputs. Nitrogen cycling is closely linked with the carbon (C) and water cycles. Thus models of N cycling must include aspects of these other cycles. Although much is known about individual processes, development of a generic model of forest N cycling is not possible at present because the links and interactions among the individual processes are not well understood. The weakest links with respect to the N cycle are: quantification of atmospheric (especially dry) deposition rates in polluted environments, controls on C and N allocation in vegetation, controls on N turnover in fine roots, controls on decomposition of the older components of soil organic matter, and feedbacks among N availability, litter "quality" and subsequent N mineralization rates. To examine possible effects of long-term change in climate or atmospheric chemistry on the storage of C and other elements in forest ecosystems, we need to model in detail the effects of these factors on complex soil processes such as organic matter decomposition. Some promising models have been developed, but they need to be validated across a range of forest types before they can be used with confidence for long-term prediction. Mean annual leaf litter N concentration offers potential as a simple index of annual N uptake in forests.

Journal Article↗

Forest fragmentation is associated with primary brood sex ratio in the treecreeper (Certhia familiaris).

We studied the primary brood sex ratio of an old-growth forest passerine, the Eurasian treecreeper (Certhia familiaris), along a gradient of forest fragmentation. We found evidence that male nestlings were more costly to produce, since they suffered twofold higher nestling mortality and were larger in body size than females. Furthermore, the proportion of males in the brood was positively associated with the provisioning rate and the amount of food delivered to the nestlings. During the first broods, a high edge density and a high proportion of pine forests around the nests were related to a decreased production of males. The densities of spiders, the main food of the treecreeper, were 38% higher on spruce trunks than on pine trunks. This suggests that pine-dominated territories with female-biased broods may have contained less food during the first broods. The observation was further supported by the fact that the feeding frequencies were lower in territories with high proportions of pines. In the second broods, territories with a high forest patch density produced female-biased broods, whereas high-quality territories with a large amount of deciduous trees and mixed forests produced male-biased broods. Our results suggest that habitat quality as measured by habitat characteristics is associated with sex allocation in free-living birds.

Animals↗

Can sunspot activity and ultraviolet-B radiation explain cyclic outbreaks of forest moth pest species?

Cyclic outbreaks of forest moth pest species have long remained a puzzle for foresters and ecologists. This paper presents time-series exhibiting a strong negative relationship between sunspot numbers and population indices of autumnal and winter moths, both in a mountain birch forest in central Norway and in a mixed lowland forest in southern Norway. In the latter area, also the population level of a moth species feeding entirely on lichens was negatively related to sunspot numbers. Low sunspot activity leads to a thinner ozone layer and thus higher surface ultraviolet (UV)-B radiation. As winter moth larvae prefer leaves subjected to enhanced UV-B radiation, we suggest that the causal relationship between sunspots and moths is that the metabolic costs of producing UV-B-protective pigments during periods of low sunspot activity reduce trees' and lichens' resistance to herbivores, and thus increase the survival of moth larvae. Higher peak densities of moth cycles in mountain forests could be explained by the general higher UV-B radiation at higher altitudes.

Animals↗

Increasing biomass in Amazonian forest plots.

A previous study by Phillips et al. of changes in the biomass of permanent sample plots in Amazonian forests was used to infer the presence of a regional carbon sink. However, these results generated a vigorous debate about sampling and methodological issues. Therefore we present a new analysis of biomass change in old-growth Amazonian forest plots using updated inventory data. We find that across 59 sites, the above-ground dry biomass in trees that are more than 10 cm in diameter (AGB) has increased since plot establishment by 1.22 +/- 0.43 Mg per hectare per year (ha(-1) yr(-1), where 1 ha = 10(4) m2), or 0.98 +/- 0.38 Mg ha(-1) yr(-1) if individual plot values are weighted by the number of hectare years of monitoring. This significant increase is neither confounded by spatial or temporal variation in wood specific gravity, nor dependent on the allometric equation used to estimate AGB. The conclusion is also robust to uncertainty about diameter measurements for problematic trees: for 34 plots in western Amazon forests a significant increase in AGB is found even with a conservative assumption of zero growth for all trees where diameter measurements were made using optical methods and/or growth rates needed to be estimated following fieldwork. Overall, our results suggest a slightly greater rate of net stand-level change than was reported by Phillips et al. Considering the spatial and temporal scale of sampling and associated studies showing increases in forest growth and stem turnover, the results presented here suggest that the total biomass of these plots has on average increased and that there has been a regional-scale carbon sink in old-growth Amazonian forests during the previous two decades.

Biomass↗

Some aspects of ecophysiological and biogeochemical responses of tropical forests to atmospheric change.

Atmospheric changes that may affect physiological and biogeochemical processes in old-growth tropical forests include: (i) rising atmospheric CO2 concentration; (ii) an increase in land surface temperature; (iii) changes in precipitation and ecosystem moisture status; and (iv) altered disturbance regimes. Elevated CO2 is likely to directly influence numerous leaf-level physiological processes, but whether these changes are ultimately reflected in altered ecosystem carbon storage is unclear. The net primary productivity (NPP) response of old-growth tropical forests to elevated CO2 is unknown, but unlikely to exceed the maximum experimentally measured 25% increase in NPP with a doubling of atmospheric CO2 from pre-industrial levels. In addition, evolutionary constraints exhibited by tropical plants adapted to low CO2 levels during most of the Late Pleistocene, may result in little response to increased carbon availability. To set a maximum potential response for a Central Amazon forest, using an individual-tree-based carbon cycling model, a modelling experiment was performed constituting a 25% increase in tree growth rate, linked to the known and expected increase in atmospheric CO2. Results demonstrated a maximum carbon sequestration rate of ca. 0.2 Mg C per hectare per year (ha(-1) yr(-1), where 1 ha = 10(4) m2), and a sequestration rate of only 0.05 Mg C ha(-1) yr(-1) for an interval centred on calendar years 1980-2020. This low rate results from slow growing trees and the long residence time of carbon in woody tissues. By contrast, changes in disturbance frequency, precipitation patterns and other environmental factors can cause marked and relatively rapid shifts in ecosystem carbon storage. It is our view that observed changes in tropical forest inventory plots over the past few decades is more probably being driven by changes in disturbance or other environmental factors, than by a response to elevated CO2. Whether these observed changes in tropical forests are the beginning of long-term permanent shifts or a transient response is uncertain and remains an important research priority.

Atmosphere↗

Sources or sinks? The responses of tropical forests to current and future climate and atmospheric composition.

How tropical rainforests are responding to the ongoing global changes in atmospheric composition and climate is little studied and poorly understood. Although rising atmospheric carbon dioxide (CO2) could enhance forest productivity, increased temperatures and drought are likely to diminish it. The limited field data have produced conflicting views of the net impacts of these changes so far. One set of studies has seemed to point to enhanced carbon uptake; however, questions have arisen about these findings, and recent experiments with tropical forest trees indicate carbon saturation of canopy leaves and no biomass increase under enhanced CO2. Other field observations indicate decreased forest productivity and increased tree mortality in recent years of peak temperatures and drought (strong El Niño episodes). To determine current climatic responses of forests around the world tropics will require careful annual monitoring of ecosystem performance in representative forests. To develop the necessary process-level understanding of these responses will require intensified experimentation at the whole-tree and stand levels. Finally, a more complete understanding of tropical rainforest carbon cycling is needed for determining whether these ecosystems are carbon sinks or sources now, and how this status might change during the next century.

Atmosphere↗

Sampling ozone exposure of Canadian forests at different scales: some case studies.

The use of passive samplers in extensive monitoring, such as that used in national forest health monitoring plots, indicates that these devices are able to determine both spatial and temporal differences in ozone exposure of the plots. This allows for categorisation of the plots and the potential for cause-effect analysis of certain forest health responses. Forest exposure along a gradient of air pollution deposition demonstrates large variation in accumulated exposures. The efficacy of using passive samplers for in situ monitoring of forest canopy exposure was also demonstrated. The sampler data produced weak relationships with ozone values from the nearest "continuous" monitor, even though data from colocated samplers showed strong relationships. This spatial variation and the apparent effect of elevation on ozone exposure demonstrate the importance of topography and tree canopy characteristics in plant exposure on a regional scale. In addition, passive sampling may identify the effects of local pollutant gases, such as NO, which may scavenge ozone locally only to increase the production of this secondary pollutant downwind, as atmospheric reactions redress the equilibrium between concentrations of this precursor and those of the generated ozone. The use of passive samplers at the stand level is able to resolve vertical profiles within the stand and edge effects that are important in exposure of understorey and ground flora. Recent case studies using passive samplers to determine forest exposure to ozone indicate a great potential for the development of spatial models on a regional, landscape, and stand level scale.

Canada↗

Regional comparison of nitrogen export to Japanese forest streams.

Nitrogen (N) emissions in Asian countries are predicted to increase over the next several decades. An understanding of the mechanisms that control temporal and spatial fluctuation of N export to forest streams is important not only to quantify critical loads of N, N saturation status, and soil acidification N dynamics and budgets in Japanese forested watersheds is not clear due to the lack of regional comparative studies on stream N chemistry. To address the lack of comparative studies, we measured inorganic N (nitrate and ammonium) concentrations from June 2000 to May 2001 in streams in 18 experimental forests located throughout the Japanese archipelago and belonging to the Japanese Union of University Forests. N concentrations in stream water during base flow and high flow periods were monitored, and N mineralization potential in soil was measured using batch incubation experiments. Higher nitrate concentrations in stream water were present in central Japan, an area that receives high rates of atmospheric N deposition. In northern Japan, snowmelt resulted in increased nitrate concentrations in stream water. The potential net N mineralization rate was higher in surface soil than in subsurface soil, and the high potential for N mineralization in the surface soil partly contributed to the increase in nitrate concentration in stream water during a storm event. Regional differences in the atmospheric N deposition and seasonality of precipitation and high discharge are principal controls on the concentrations and variations of nitrates in stream water in forested watersheds of Japan.

Ammonia↗

Soil N fluxes in three contrasting dry tropical forests.

A comparative study of N fluxes in soil among a dry dipterocarp forest (DDF), a dry evergreen forest (DEF), and a hill evergreen forest (HEF) in Thailand was done. N fluxes in soil were estimated using an ion exchange resin core method and a buried bag method. Soil C and N pools were 38 C Mg/ha/30 cm and 2.5 N Mg/ha/30 cm in DDF, 82 C Mg/ha/30 cm and 6.2 N Mg/ha/30 cm in DEF, and 167 C Mg/ha/30 cm and 9.3 N Mg/ha/30 cm in HEF. Low C concentration in the DDF and DEF sites was compensated by high fine soil content. In the highly weathered tropical soil, fine soil content seemed to be important for C accumulation. Temporal and vertical fluctuations of N fluxes were different among the sites. The highest N flux was exhibited at the onset of the wet season in DDF, whereas inorganic N production and estimated uptake of N were relatively stable during the wet season in DEF and HEF. It is suggested that N cycling in soil becomes stable in dry tropical forests to intermediate in temperate forests. N deposition may result in large changes of N cycling in the DDF and DEF due to low accumulations of C and N.

Carbon↗

Respiratory carbon losses and the carbon-use efficiency of a northern hardwood forest, 1999-2003.

Quantitative assessment of carbon (C) storage by forests requires an understanding of climatic controls over respiratory C loss. Ecosystem respiration can be estimated biometrically as the sum (R Sigma) of soil (Rs), leaf (Rl) and wood (Rw) respiration, and meteorologically by measuring above-canopy nocturnal CO2 fluxes (Fcn). Here we estimated R Sigma over 5 yr in a forest in Michigan, USA, and compared R Sigma and Fcn on turbulent nights. We also evaluated forest carbon-use efficiency (Ec = P(NP)/P(GP)) using biometric estimates of net primary production (P(NP)) and R Sigma and Fcn-derived estimates of gross primary production (P(GP)). Interannual variation in R Sigma was modest (142 g C m(-2) yr(-1)). Mean annual R Sigma was 1425 g C m(-2) yr(-1); 71% from Rs, 18% from Rl, and 11% from Rw. Hourly R Sigma was well correlated with Fcn, but 11 to 58% greater depending on the time of year. Greater R Sigma compared with Fcn resulted in higher estimated annual P(GP) and lower annual Ec (0.42 vs 0.54) using biometric and meteorological data, respectively. Our results provide one of the first multiyear estimates of R Sigma in a forested ecosystem, and document the responses of component respiratory C losses to major climatic drivers. They also provide the first assessment of Ec in a deciduous forest using independent estimates of P(GP).

Carbon↗

Conservation of the northern spotted owl under the Northwest Forest Plan.

Development of the Northwest Forest Plan (NWFP) was motivated by concerns about the over-harvest of late-seral forests and the effects of intensive forest management on the long-term viability of the Northern Spotted Owl (Strix occidentalis caurina). Following several years of intense political and legal debates, the final NWFP was approved in 1994. Even though the plan evolved with a broad ecosystem perspective, it remained anchored in the Spotted Owl reserve design proposed in 1990. Based on a criterion of stable or increasing populations, a decade later it remains unclear whether the enactment of the NWFP has improved the conservation status of Spotted Owls. The results of intensive monitoring of several Spotted Owl populations for over a decade suggest a continuing range-wide decline even though rates of timber harvest have declined dramatically on federal lands. The cause of the decline is difficult to determine because the research needed to establish cause and effect relations has not been done. One plausible hypothesis is that the owl's life history greatly constrains its rate of population growth even when habitat is no longer limiting. Since enactment of the NWFP, new threats have arisen, including the movement of Barred Owls (S. varia) into the range of the Spotted Owl, political pressure to increase levels of timber harvest, and recent changes to forest laws that eliminate the requirement to assess the viability of wildlife populations on U.S. Department of Agriculture Forest Service lands. At this time is appears that Spotted Owl conservation rests critically on continued implementation of the protections afforded by the NWFP and the U.S. Endangered Species Act.

Animals↗

Salvage logging in the montane ash eucalypt forests of the Central Highlands of Victoria and its potential impacts on biodiversity.

The two major forms of disturbance in the montane ash eucalypt forests of the Central Highlands of Victoria (southeastern Australia) are clearfell logging and unplanned wildfires. Since the 1930s wildfire has been followed by intensive and extensive salvage-logging operations, which may proceed for many years after a wildfire has occurred. Although applied widely, the potential effects of salvage logging on native flora and fauna have been poorly studied. Our data indicate that the abundance of large trees with hollows is significantly reduced in forests subject to salvage harvesting. This has implications for thepersistence of an array of such cavity-using vertebrates as the endangered arboreal marsupial, Leadbeater's possum (Gymnobelidues leadbeateri). Salvage logging also reduces the prevalence of multiaged montane ash forests--places that typically support the highest diversity of arboreal marsupials and forest birds. Limited research has been conducted on the effects of salvage logging on plants; thus, we constructed hypotheses about potential impacts for further testing based on known responses to clearfell logging and key life history attributes. We predict many species, such as vegetatively resprouting tree ferns, will decline, as they do after clearfelling. We also suggest that seed regenerators, which typically regenerate well after fire or conventional clearfelling, will decline after salvage logging because the stimulation for germination (fire) takes place prior to mechanical disturbance (logging). Understoryplant communities in salvage-logged areas will be dominated by a smaller suite of species, and those that are wind dispersed, have viable soil-stored seed remaining after salvage logging, or have deep rhizomes are likely to be advantaged. We recommend the following improvements to salvage-logging policies that may better incorporate conservation needs in Victorian montane ash forests: (1) exemption of salvage logging from some areas (e.g., old-growth stands and places subject to only partial stand damage); (2) increased retention of biological legacies on burned areas through variations in the intensity of salvage logging; and (3) reduction in the levels of physical disturbance on salvage-logged areas, especially through limited seedbed preparation and mechanical disturbance.

Animals↗

Influence of forest fragmentation on community structure of frogs and lizards in northeastern Costa Rica.

To better understand responses of reptiles and amphibians to forest fragmentation in the lowland Neotropics, we examined community and population structure of frogs and lizards in the fragmented landscape surrounding La Selva Biological Station in the Sarapiquí region of northeastern Costa Rica. We used diurnal quadrats and nocturnal transects to sample frogs and lizards in nine forest fragments (1-7 ha each) and La Selva (1100 ha). Species richness in all fragments combined was 85% of that found in La Selva with comparable sampling effort. Richness varied from 10 to 24 species among forest fragments, compared with 36 species at La Selva. Lizard density was higher and frog density was lower in forest fragments than in La Selva. Community composition varied among sites and by fragment size class, and species occurrence was nested with respect to fragment area. Isolation and habitat variables did not significantly affect species richness, composition, or nestedness. We classified 34% of species as fragmentation sensitive because they were absent or occurred at low densities in fragments. Nevertheless, the relatively high diversity observed in the entire set of fragments indicates that preserving a network of small forest patches may be of considerable conservation value to the amphibians and reptiles of this region.

Animals↗