Rapid Cycling of Organic Nitrogen in Taiga Forest Ecosystems

ABSTRACT We examined the dynamics of organic nitrogen (N) turnover in situ across a primary successional sequence in interior Alaska, USA, in an attempt to understand the magnitude of these fluxes in cold, seasonally frozen soils. Through a combination of soil extraction procedures and measurements of ¹³C-enriched CO₂ efflux from soils amended in the field with ¹³C-labeled amino acids, we were able to trace the fate of this N form. Amino acid turnover in situ at soil temperatures of 10°C or below show that amino acids represent a highly dynamic soil N pool with turnover times of approximately 3–6 h. The rapid turnover of free amino acids is associated with high soil proteolytic activity, which in turn is tightly correlated with soil protein concentration. Moreover, these estimates of soil amino acid turnover in the field correspond well with measurements of amino acid turnover under equivalent temperatures in the laboratory. The gross flux of amino acid-N over the growing season greatly exceeded the annual vegetation N requirement, suggesting that microbial biomass represent a significant sink for this organic N. Depending on the strength of this sink, N flow via free soil amino acids can potentially account for the entire N demand of vegetation in the absence of net N mineralization. These relationships underscore the important biogeochemical role of labile DON fractions in high-latitude forest ecosystems.     

Carbon Quality and Stocks in Organic Horizons in Boreal Forest Soils

We investigated the mechanisms that determine the quality and quantity of organic carbon (C) stocks in boreal forest soils by analyzing both qualitative and quantitative changes in the organic fractions in the soil organic matter (OM) in a vertical gradient in the decomposition continuum of the organic horizon [litter layer (L), fermentation layer (F), and humus layer (H)] in forest soils using a sequential fractionation method at two forest types along a climatic gradient in Finland. We predicted that the concentrations of water-soluble (WSE) and non-polar (NPE) extractives should decrease and those of the acid-soluble (AS) fraction and acid-insoluble residue (AIR) should increase from the L to the F, and from the F to the H layers, but the C/N ratio of soil OM should stay constant after reaching the critical quotient. We also predicted that the AIR concentrations should be higher in the south than north boreal, and in sub-xeric than mesic forests. Consistent with our hypothesis, the concentrations of WSE and NPE fractions decreased and concentrations of AIR increased in the vertical soil gradient. The highest concentrations of the AS fraction were found in the F layer. The C/N ratio was lowest in the F layer, and the highest in the H layer, indicating that soil OM is depleted in N in relation to C along the vertical soil gradient. Concentrations of WSE and NPE were lower, and concentrations of AIR were higher in the south than in north boreal forests, which is in agreement with our hypothesis that higher soil temperatures may enhance accumulation of slowly decomposable OM in the soil. The concentrations of AIR were higher in the sub-xeric than mesic forests. Contrary to our expectations, however, the differences in the chemical quality in soil OM between the site types were amplified from the L to the H layer. The size of the C storage was significantly larger in south than north boreal sites, and larger in the mesic than in the sub-xeric sites.     

Spatial Variation of Bird Species in Landscapes Dominated by Old-growth Forests in Northern Boreal Finland

An important issue in conservation planning is to study the distribution and abundance patterns of species in natural landscapes. Information of concordant and clumped species distribution pattertns can enhance attempts to focus conservation efforts in sites/areas preferred by various species. We compared bird species abundance, community composition and species distribution patterns in three large old-growth forest areas (size 40–120 km²) in northern boreal Finland on the basis of quantitative bird censuses. Total bird density and species composition, and mean density and variances of the most abundant species were highly similar between the three areas. Most of the bird species were distributed randomly within the areas and species preferring old-growth forests showed compensatory density variation without preference for any sub-area. Mean density of species preferring old-growth forests was significantly higher in the studied large old-growth forests than in smaller old-growth forests and was about threefold higher than regional density of species in a predominantly managed landscape matrix. Bird species preferring old-growth forests are probably not dependent on a certain specific part of a virgin boreal forest landscape but rather on the overall size of the high-quality and diverse old-growth forest area.     

Carbon and Nitrogen Dynamics of Boreal Jack Pine Stands With and Without a Green Alder Understory

We compared the species composition, structure and selected components of the carbon (C) and nitrogen (N) budgets of similar-aged, mature boreal jack pine (Pinus banksiana Lamb.) forests with and without green alder [Alnus crispa (Ait.) Pursh.] in two different boreal environments. The C and N content of the overstory biomass components (for example, stem, branch, and foliage), total vegetation, forest floor, and mineral soil were greater (P= 0.05 to P= 0.10) for jack pine with alder (JPA) stands than for jack pine without alder (JP) stands at both study areas. Jack pine foliage N isotopic discrimination (δ¹⁵N) and annual litterfall N content were significantly greater (P < 0.05) in the JPA than the JP stands at both study areas, suggesting that alder was fixing N and that N availability was greater in the JPA than the JP stands. The greater leaf area index (LAI) and overstory C accumulation in the JPA than the JP stands (P < 0.05) is likely because of the greater N availability in the JPA stands, but the effect of soil texture discontinuity on water availability in the JPA stands can not be dismissed. Percent ground cover by feathermoss varied among the jack pine communities and was positively correlated with overstory LAI (r ²= 0.83, P< 0.05). One index of N-use efficiency (NUE), defined as aboveground net primary productivity (ANPP) per litterfall N, was significantly greater (P < 0.05) for the JP than the JPA stands, but a second index of NUE, ANPP/N uptake, did not differ between the two jack pine communities. Jack pine trees growing without alder produced more organic matter per unit of N, but percent N retranslocation from senescing foliage and N mean residence time in the overstory did not differ between the JPA and the JP stands. A conceptual model is presented that illustrates the potential influence of alder on the species composition, structure, and function of boreal jack pine forests. Received 6 January 1998; accepted 15 April 1998.     

Base Cation Cycling in a Pristine Watershed of the Canadian Boreal Forest

In forest ecosystems the single largest respiratory flux influencing net ecosystem productivity (NEP) is the total soil CO₂ efflux; however, it is difficult to make measurements of this flux that are accurate at the ecosystem scale. We examined patterns of soil CO₂ efflux using five different methods: auto-chambers, portable gas analyzers, eddy covariance along and two models parameterized with the observed data. The relation between soil temperature and soil moisture with soil CO₂ effluxes are also investigated, both inter-annually and seasonally, using these observations/results. Soil respiration rates (R _soil) are greatest during the growing season when soil temperatures are between 15 and 25 °C, but some soil CO₂ efflux occurs throughout the year. Measured soil respiration was sensitive to soil temperature, particularly during the spring and fall. All measurement methods produced similar annual estimates. Depending on the time of the year, the eddy covariance (flux tower) estimate for ecosystem respiration is similar to or slightly lower than estimates of annual soil CO₂ efflux from the other methods. As the eddy covariance estimate includes foliar and stem respiration which the other methods do not; it was expected to be larger (perhaps 15–30%). The auto-chamber system continuously measuring soil CO₂ efflux rates provides a level of temporal resolution that permits investigation of short- to longer term influences of factors on these efflux rates. The expense of building and maintaining an auto chamber system may not be necessary for those researchers interested in estimating R _soil annually, but auto-chambers do allow the capture of data from all seasons needed for model parameterization.     

Mass-Balance Analyses of Boreal Forest Population Cycles: Merging Demographic and Ecosystem Approaches

UsingEcopath, a trophic mass-balance modeling framework, we developed six models of a Canadian boreal forest food web centered around snowshoe hares, which have conspicuous 10-year population cycles. Detailed models of four phases of the cycle were parameterized with long-term population data for 12 vertebrate taxa. We also developed five other models that, instead of observed data, used parameter values derived from standard assumptions. Specifically, in the basic model, production was assumed to equal adult mortality, feeding rates were assumed to be allometric, and biomass was assumed to be constant. In the actual production, functional response, and biomass change models, each of these assumed values from the basic model was replaced individually by field data. Finally, constant biomass models included actual production by all species and functional responses of mammalian predators and revealed the proportion of herbivore production used by species at higher trophic levels. By comparing these models, we show that detailed information on densities and demographics was crucial to constructing models that captured dynamic aspects of the food web. These detailed models reinforced an emerging picture of the causes and consequences of the snowshoe hare cycle. The snowshoe hare decline and low phases were coincident with times when per capita production was relatively low and predation pressure high. At these times, ecotrophic efficiencies (EE) suggest there was little production that remained unconsumed by predators. The importance of both production and consumption implies that bottom–up and top–down factors interacted to cause the cycle. EEs of other herbivores (ground squirrels, red squirrels, small mammals, small birds, grouse) were generally low, suggesting weak top–down effects. Predation rates on these “alternative” prey, except ground squirrels, were highest when predators were abundant, not when hares were rare; consequently, any top–down effects reflected predator biomass and were not a function of diet composition or functional responses. Finally, several predators (lynx, coyotes, great-horned owls) showed clear bottom–up regulation, reproducing only when prey exceeded threshold densities. Taken altogether, these results demonstrate that ecosystem models parameterized by population data can describe the dynamics of nonequilibrial systems, but only when detailed information is available for the species modeled. Received 30 November 2000; Accepted 6 September 2001.     

Effect of Bat Exclusion on Patterns of Seed Rain in Tropical Rain Forest in French Guiana

To assess the impact of bats on seed dispersal in a tropical mature forest (Nouragues, French Guiana), we conducted a bat exclusion experiment and tested the hypotheses that an artificial reduction in the abundance of bats would result in: (1) a decrease in seed species diversity, at the community level; and (2) an increase in seed limitation (a failure of seeds to reach all suitable sites for germination) at the species level. Seed rain was sampled in two contiguous forest plots for a total of 120 d, using 49 seed traps (1 m²) arranged in 7 × 7 grids and spaced at 5-m intervals. Using mist nets, bat activity was reduced in one forest plot for a total of 60 nights. Thirty-nine plant species, or species groups, likely to be consumed and dispersed by bats, were collected within a total sample of 50,063 seeds. The overall seed rain was dominated by epiphytic Araceae and Cyclanthaceae (83.3%) and tree species within the genera Cecropia and Ficus (16.0%). Seeds from bat-dispersed shrubs and treelets ( Piper , Solanum , and Vismia ) were relatively rare (0.7%). The bat exclusion resulted in a 30.5 percent reduction in seed species richness and increased seed limitation for most of the species sampled. Seed limitation was caused mainly by a reduced seed rain (seed source limitation) rather than a decrease in seed dispersal uniformity (seed dispersal limitation). Therefore, bat-dispersed plants with low seed production are likely to be particularly affected by a decline in bat abundance, as a result of anthropogenic change.     

Diversity and Community Structure of Primary Wood-Inhabiting Bacteria in Boreal Forest

DNA-based pyrosequencing analysis of the V1- V3 16S rRNA gene region was used to identify bacteria community and shift during early stages of wood colonization in boreal forest soils. The dataset comprised 142,447 sequences and was affiliated to 11 bacteria phyla, 25 classes and 233 genera. The dominant groups across all samples were Proteobacteria, followed by Bacteroidetes, Acidobacteria, Actinobacteria, Amatimonadetes, Planctomycetes and TM7 group. The community structure of the primary wood-inhabiting bacteria differed between types of forest soils and the composition of bacteria remained stable over prolonged incubation time. The results suggest that variations in soil bacterial community composition have an influence on the wood-inhabiting bacterial structure.     

Impacts of Elevated Carbon Dioxide and Temperature on a Boreal Forest Ecosystem (CLIMEX Project)

To evaluate the effects of climate change on boreal forest ecosystems, both atmospheric CO₂ (to 560 ppmv) and air temperature (by 3°–5°C above ambient) were increased at a forested headwater catchment in southern Norway. The entire catchment (860 m²) is enclosed within a transparent greenhouse, and the upper 20% of the catchment area is partitioned such that it receives no climate treatment and serves as an untreated control. Both the control and treatment areas inside the greenhouse receive deacidified rain. Within 3 years, soil nitrogen (N) mineralization has increased and the growing season has been prolonged relative to the control area. This has helped to sustain an increase in plant growth relative to the control and has also promoted increased N export in stream water. Photosynthetic capacity and carbon–nitrogen ratio of new leaves of most plant species did not change. While the ecosystem now loses N, the long-term fate of soil N is a key uncertainty in predicting the future response of boreal ecosystems to climate change. Received 18 November 1997; accepted 13 April 1998.     

Differences in Englemann Spruce Forest Biogeochemistry East and West of the Continental Divide in Colorado, USA

We compared Englemann spruce biogeochemical processes in forest stands east and west of the Continental Divide in the Colorado Front Range. The divide forms a natural barrier for air pollutants such that nitrogen (N) emissions from the agricultural and urban areas of the South Platte River Basin are transported via upslope winds to high elevations on the east side but rarely cross over to the west side. Because there are far fewer emissions sources to the west, atmospheric N deposition is 1–2 kg N ha⁻¹ y⁻¹ on the west side, as compared with 3–5 kg N ha⁻¹ y⁻¹ on the east side. Species composition, elevation, aspect, parent material, site history, and climate were matched as closely as possible across six east and six west side old-growth forest stands. Higher N deposition sites had significantly lower organic horizon C:N and lignin:N ratios, lower foliar C:N ratios, as well as greater %N, higher N:Ca, N:Mg, and N:P ratios, and higher potential net mineralization rates. When C:N ratios dropped below 29, as they did in east-side organic horizon soils, mineralization rates increased linearly. Our results are comparable to those from studies of the northeastern United States and Europe that have found changes in forest biogeochemistry in response to N deposition inputs between 3 and 60 kg ha⁻¹ y⁻¹. Though they are low by comparison with more densely populated and agricultural regions, current levels of N deposition, have caused measurable changes in Englemann spruce forest biogeochemistry east of the Continental Divide in Colorado. Received 22 January 2001; accepted 11 June 2001.     

Plant Effects on Spatial and Temporal Patterns of Nitrogen Cycling in Shortgrass Steppe

Because of the water-limited nature and discontinuous plant cover of shortgrass steppe, spatial patterns in ecosystem properties are influenced more by the presence or absence of plants than by plant type. However, plant type may influence temporal patterns of nutrient cycling between plant and soil. Plants having the carbon-3 (C₃) or carbon-4 (C₄) photosynthetic pathway differ in phenology as well as other attributes that affect nitrogen (N) cycling. We estimated net N mineralization rates and traced nitrogen-15 (¹⁵N) additions among plant and soil components during May, July, and September of 1995 in native plots of C₃ plants, C₄ plants, or mixtures of C₃ and C₄. Net N mineralization was significantly greater in C₃ plots than in C₄ plots during both July and September. C₃ plots retained significantly more ¹⁵N in May than did mixed and C₄ plots; these differences in ¹⁵N retention were due to greater ¹⁵N uptake by C₃ plants than by C₄ plants during May. There were no significant differences in total ¹⁵N retention among plant communities for July and September. Soil ¹⁵N was influenced more by presence or absence of plants than by type of plant; greater quantities of ¹⁵N remained in soil interspaces between plants than in soil directly under plants for July and September. Our results indicate that plant functional type (C₃ versus C₄) can affect both the spatial and the temporal patterns of N cycling in shortgrass steppe. Further research is necessary to determine how these intraseasonal differences translate to longer-term and coarser-scale effects of plants on N cycling, retention, and storage. Received 8 December 1997; accepted 6 May 1998.     

Fine‐scale Spatial Genetic Structure of Ten Dipterocarp Tree Species in a Bornean Rain Forest

Fine‐scale spatial genetic structure is increasingly recognized as an important factor in the studies of tropical forest trees as it influences genetic diversity of local populations. The biologic mechanisms that generate fine‐scale spatial genetic structure are not fully understood. We studied fine‐scale spatial genetic structure in ten coexisting dipterocarp tree species in a Bornean rain forest using microsatellite markers. Six of the ten species showed statistically significant fine‐scale spatial genetic structure. Fine‐scale spatial genetic structure was stronger at smaller spatial scales (≤ 100 m) than at larger spatial scales (> 100 m) for each species. Multiple regression analysis suggested that seed dispersal distance was important at the smaller spatial scale. At the larger scale (> 100 m) and over the entire sample range (0–1000 m), pollinators and spatial distribution of adult trees were more important determinants of fine‐scale spatial genetic structure. Fine‐scale spatial genetic structure was stronger in species pollinated by less mobile small beetles than in species pollinated by the more mobile giant honeybee (Apis dorsata). It was also stronger in species where adult tree distributions were more clumped. The hypothesized mechanisms underlying the negative correlation between clump size and fine‐scale spatial genetic structure were a large overlap among seed shadows and genetic drift within clumped species.     

Land-Use Changes in Southern Appalachian Landscapes: Spatial Analysis and Forecast Evaluation

Understanding human disturbance regimes is crucial for developing effective conservation and ecosystem management plans and for targeting ecological research to areas that define scarce ecosystem services. We evaluate and develop a forecasting model for land-use change in the Southern Appalachians. We extend previous efforts by (a) addressing the spatial diffusion of human populations, approximated by building density, (b) examining a long time period (40 years, which is epochal in economic terms), and (c) explicitly testing the forecasting power of the models. The resulting model, defined by linking a negative binomial regression model of building density with a logit model of land cover, was fit using spatially referenced data from four study sites in the Southern Appalachians. All fitted equations were significant, and coefficient estimates indicated that topographic features as well as location significantly shape population diffusion and land use across these landscapes. This is especially evident in the study sites that have experienced development pressure over the last 40 years. Model estimates also indicate significant spatial autocorrelation in land-use observations. Forecast performance of the models was evaluated by using a separate validation data set for each study area. Depending on the land-use classification scheme, the models correctly predicted between 68% and 89% of observed land uses. Tests based on information theory reject the hypothesis that the models have no explanatory power, and measures of entropy and information gain indicate that the estimated models explain between 47% and 66% of uncertainty regarding land-use classification. Overall, these results indicate that modeling land-cover change alone may not be useful over the long run, because changing land cover reflects the outcomes of more than one human process (for example, agricultural decline and population growth). Here, additional information was gained by addressing the spatial spread of human populations. Furthermore, coarse-scale measures of the human drivers of landscape change (for example, population growth measured at the county level) appear to be poor predictors of changes realized at finer scales. Simulations demonstrate how this type of approach might be used to target scarce resources for conservation and research efforts into ecosystem effects. Received 13 March 1998; accepted 30 September 1998     

Analysis of Large-Scale Spatial Heterogeneity in Vegetation Indices among North American Landscapes

We analyzed the spatial heterogeneity in vegetation indices among 13 North American landscapes by using full Landsat Thematic Mapper images. Landscapes varied broadly in the statistical distribution of vegetation indices, but were successfully ordinated by using a measure of central tendency (the mean) and a measure of dispersion (the standard deviation or the coefficient of variation). Differences in heterogeneity among landscapes were explained by their topographic relief and their land cover. Landscape heterogeneity (standard deviation of the Normalized Difference Vegetation Index, NDVI) tended to increase linearly with topographic relief (standard deviation of elevation), but landscapes with low relief were much more heterogeneous than expected from this relationship. The latter were characterized by a large proportion of agricultural land. Percent agriculture, in turn, was inversely related to topographic relief. The strength of these relationships was evaluated against changes in image spatial resolution (grain size). Aggregation of NDVI images to coarser grain size resulted in steady decline of their standard deviation. Although the relationship between landscape heterogeneity and explanatory variables was generally preserved, rates of decrease in heterogeneity with grain size differed among landscapes. A spatial autocorrelation analysis showed that rates of decrease were related to the scale at which pattern is manifested. On one end of the spectrum are agricultural, low-relief landscapes with low spatial autocorrelation and small-scale heterogeneity associated with fields; their heterogeneity decreased sharply as grain size increased. At the other end, desert landscapes were characterized by low small-scale heterogeneity, high spatial autocorrelation, and almost no change in heterogeneity as grain sized was increased—their heterogeneity, associated with land forms, was present at a large scale. Received 1 October 1997; accepted 11 February 1998.     

Ecospace: Prediction of Mesoscale Spatial Patterns in Trophic Relationships of Exploited Ecosystems, with Emphasis on the Impacts of Marine Protected Areas

Growing disillusion with the predictive capability of single species fisheries assessment methods and the realization that the management approaches they imply will always fail to protect bycatch species has led to growing interest in the potential of marine protected areas (MPAs) as a tool for protecting such species and allowing for rebuilding populations of target species and damaged habitat. Ecospace is a spatially explicit model for policy evaluation that allows for considering the impact of MPAs in an ecosystem (that is, trophic) context, and that relies on the Ecopath mass-balance approach for most of its parameterization. Additional inputs are movement rates used to compute exchanges between grid cells, estimates of the importance of trophic interactions (top-down vs bottom up control), and habitat preferences for each of the functional groups included in the model. An application example, including the effect of an MPA, and validation against trawl survey data is presented in the form of a color map illustrating Ecospace predictions of biomass patterns on the shelf of Brunei Darussalam, Southeast Asia. A key general prediction of Ecospace is spatial “cascade” effects, wherein prey densities are low where predators are abundant, for example, in protected areas or areas where fishing costs are high. Ecospace also shows that the potential benefits of local protection can be easily negated by high movement rates, and especially by concentration of fishing effort at the edge of the MPAs, where cascade effects generate prey gradients that attract predators out of the protected areas. Despite various limitations (for example, no explicit consideration of seasonal changes or directed migration), the outward simplicity of Ecospace and the information-rich graphs it generates, coupled with the increasingly global availability of the required Ecopath files, will likely ensure a wide use for this approach, both for generating hypotheses about ecosystem function and evaluating policy choices. Received 24 February 1999; accepted 16 June 1999.     

Changes in Production and Nutrient Cycling across a Wetness Gradient within a Floodplain Forest

Floodplain forest ecosystems are highly valuable to society because of their potential for water quality improvement and vegetation productivity, among many other functions. Previous studies have indicated that hydrology influences productivity but that the relationship between hydroperiod and productivity is a complex one. Consequently, we compared multiple indexes of productivity, nutrient circulation, and hydroperiod among three communities on the Flint River floodplain, Georgia, that differed in terms of inundation frequency. We hypothesized that (a) the wettest community would have the lowest total net primary production (NPP) values because of saturated soil conditions; (b) as wetness increases, nutrient circulation in litterfall would decrease because of the hypothesized lower productivity in the wetter community; and (c) as wetness increases, internal translocation would become more efficient. The study site was partitioned into three wetness types—somewhat poorly drained (SPD), intermediate (I) and poorly drained (PD). We found that belowground biomass was greatest on the SPD, litterfall was similar for all three sites, and that woody biomass current annual increment (CAI) was greatest in the PD community. However, when the three variables were totaled for each site, the PD had the greatest NPP, thus disproving hypothesis (a). For hypothesis (b), we observed that P content in litterfall, although not significant, followed the predicted trend; nitrogen (N) content displayed the opposite pattern (PD > I > SPD). As wetness increased, internal translocation became more efficient for phosphorus (support for hypothesis [c]), but the SPD community was more efficient at retranslocating N (contradiction of hypothesis [c]). Received 19 June 2000; accepted 19 October 2000.     

Patterns of nutrient flow in the pygmy forest region of northern California

Summary Vegetation of the Mendocino coastal region forms a gradient from coastal redwood (Sequoia sempervirens) stands, through Bishop pine (Pinus muricata), to endemic pygmy conifers (Pinus contorta ssp.bolanderi, Cupresses pygmaca) on terrace flat/podzols. Along this gradient nutrient stocks and net uptake decrease, and strategies for rapid nutrient circulation within the ecosystems are increasingly in evidence. Within the pygmy conifer forest, the dominant plant species exhibit differing characteristics in their pattern of net uptake, partition and turnover or recirculation of each essential element. The pygmy forest nutrient pool has apparently been built up over centuries by soil weathering, N-fixation by lichens, and inputs from precipitation. Such an ecosystem would be likely to show weak resilience to a perturbation that removed standing vegetattion or the litter layer. Across a wide range of terrestrial ecosystems, the ratio of plant nutrient stocks to phytomass tends to decrease with increase in phytomass and net primary productivity. This trend is reflected on a smaller scale by the decrease in nutrient phytomass ratio from pygmy forest, through Bishop pine, to redwood stands (0.013, 0.009, 0.006 respectively). Nevertheless, terrestrial communities in the temperate and subtropical regions of moderate rainfall and non-saline soils appear to share a relatively constant nutrient/phytomass concentration (mean ±s.e.=0.101±0.0003; n=9).Nomenclature follows Munz, 1959: A Californian Flora. Univ. California Press, Berkeley.This research formed part of my doctoral dissertation at Cornell University. I am grateful for the extensive guidance of R.H. Whittaker throughout this work, and for his support under National Science Foundation grant GB-8095X. I thank Hans Jenny for permission to include unpublished information from his studies, and T. Wurzburger (Univ. Calif., Santa Barbara), H. Gauch, Jr. (Cornell Univ.) and A. Wallace (Univ. Calif., Los Angeles) for assistance in the analysis of tissue samples.     

The Environmental Control of Near-Surface Thermoclines in Boreal Lakes

We report on the effect of lake size, water transparency, and wind on the frequency of transient near-surface thermoclines in 39 boreal lakes from the Experimental Lakes Area (ELA) and Northwest Ontario Lake Size Series (NOLSS). This study was based on more than 3000 archived temperature profiles amassed over a 25-year period for lakes ranging from 2 ha to 8 million ha in surface area. The incidence of transient thermoclines decreased with increasing lake size from 90% of all summer days in small lakes (less than 4 ha) to 40% or less in the larger NOLSS lakes (up to 34,700 ha). No transient near-surface thermoclines were detected in Lake Superior. Forest fires and climatic variability were also found to affect the frequency of near-surface thermoclines. Long-term trends indicate an increase in average annual wind velocity in the area, possibly as the result of extensive forest fires and clearcutting. The subsequent decrease in the frequency of shallow secondary thermoclines in aquatic ecosystems has possible consequences for the lake biota, as the result of changes in radiation, turbulence, and the nutrient regime. Received 8 November 2000; accepted 30 April 2001.     

Spatial and Temporal Patterns of Frugivorous Hornbill Movements in Central Africa and their Implications for Rain Forest Conservation

Tropical forest conservation and restoration require an understanding of the movements and habitat preferences of important seed dispersers. With forests now being altered at an unprecedented rate, avian frugivores are becoming increasingly vital for forest regeneration. Seed movement, however, is highly dependent on the behavioral characteristics of their dispersers. Here, we examined the movements, habitat preferences, and range sizes of two African frugivores: the Black‐casqued (Ceratogymna atrata) and the White‐thighed (Bycanistes albotibialis) Hornbill, in the lowland rain forests of southern Cameroon. Using satellite transmitters, we tracked eight hornbills for 3 yr to characterize their movements and relate them to environmental landscape features. Hornbill movements differed significantly, with B. albotibialis ranging over larger areas (mean = 20,274 ha) than C. atrata (mean = 5604 ha), and females of both species covering over 15 times the area of males. Evidence suggests that movements are irruptive during particular periods, perhaps driven by low resource availability. In addition, hornbills often returned to the same localities within a year, although movements were not characterized as migratory. Both species displayed significant differences in habitat preference, with B. albotibialis utilizing disturbed habitat more frequently than C. atrata (t = ?22.04, P = 2.2 × 10^?16). Major roads were found to act as barriers for C. atrata, but not for B. albotibialis. The ability of both hornbill species to move large distances suggests hornbills will play a vital role in the maintenance and regeneration of rain forests in Central Africa as forest fragmentation increases and terrestrial vertebrates decline in numbers.