Impact of Future Climate on Radial Growth of Four Major Boreal Tree Species in the Eastern Canadian Boreal Forest

Immediate phenotypic variation and the lagged effect of evolutionary adaptation to climate change appear to be two key processes in tree responses to climate warming. This study examines these components in two types of growth models for predicting the 2010–2099 diameter growth change of four major boreal species Betula papyrifera, Pinus banksiana, Picea mariana, and Populus tremuloides along a broad latitudinal gradient in eastern Canada under future climate projections. Climate-growth response models for 34 stands over nine latitudes were calibrated and cross-validated. An adaptive response model (A-model), in which the climate-growth relationship varies over time, and a fixed response model (F-model), in which the relationship is constant over time, were constructed to predict future growth. For the former, we examined how future growth of stands in northern latitudes could be forecasted using growth-climate equations derived from stands currently growing in southern latitudes assuming that current climate in southern locations provide an analogue for future conditions in the north. For the latter, we tested if future growth of stands would be maximally predicted using the growth-climate equation obtained from the given local stand assuming a lagged response to climate due to genetic constraints. Both models predicted a large growth increase in northern stands due to more benign temperatures, whereas there was a minimal growth change in southern stands due to potentially warm-temperature induced drought-stress. The A-model demonstrates a changing environment whereas the F-model highlights a constant growth response to future warming. As time elapses we can predict a gradual transition between a response to climate associated with the current conditions (F-model) to a more adapted response to future climate (A-model). Our modeling approach provides a template to predict tree growth response to climate warming at mid-high latitudes of the Northern Hemisphere.     

Generation of Spatial Patterns in Boreal Forest Landscapes

Boreal forests are composed of a few plant species with contrasting traits with respect to ecosystem functioning and spatial patterning. Early successional deciduous species, such as birch and aspen, disperse seeds widely, do not tolerate low light and nitrogen availabilities, have rapidly decaying litter, and are highly preferred by herbivores. These later succeed to conifers, such as spruce and fir, which disperse seeds locally, tolerate low light levels and low nitrogen availability, have litter that decays slowly, and are unpalatable to most mammalian herbivores. Although there are also early successional conifers, such as jack pine and Scots pine, the aspen-birch-spruce-fir successional sequence is the most common over much of North America, and (without fir) in Fennoscandia and Siberia. The course of succession in these forests is controlled partly by seed dispersal and selective foraging by mammalian herbivores. Both of these processes are spatially dynamic, but little is known about how their spatial dynamics may affect ecosystem processes, such as nitrogen cycling or productivity. We present spatially explicit models that demonstrate the following: (a) Spatially explicit seed dispersal results in more clumped distribution of tree species and persistence of greater paper birch biomass than uniform seed rain across the landscape. Such results are consistent with current spatially explicit population models of dispersal and coexistence. (b) With localized seed dispersal, the concentrations of available soil nitrogen are distributed in larger patches with sharp transitions from low to high nitrogen availability near patch edges. In contrast, with a uniform seed rain, the distribution of soil nitrogen availability was more uniform and “hotspots” were more localized. Thus, the spatial pattern of an ecosystem process (nitrogen cycling) is determined by seed dispersal and competition for light among competing populations. (c) A dispersing herbivore, such as moose, that selectively forages on early successional deciduous species with high quality litter, such as aspen or birch, and discriminates against late successional conifers, such as spruce or fir, imposes higher-order repeated patterns of plant species and biomass distribution on the landscape. Thus, seed dispersal and herbivore foraging correlate properties in adjacent patches but in different ways, and different spatial patterns emerge. Other processes, such as insect outbreaks, fire, and water flow, also may correlate properties between adjacent patches and result in additional patterns. Received 8 February 1999; accepted 28 May 1999.     

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.     

Effects of Soil Burn Severity on Post-Fire Tree Recruitment in Boreal Forest

Fire, which is the dominant disturbance in the boreal forest, creates substantial heterogeneity in soil burn severity at patch and landscape scales. We present results from five field experiments in Yukon Territory, Canada, and Alaska, USA that document the effects of soil burn severity on the germination and establishment of four common boreal trees: Picea glauca, Picea mariana, Pinus contorta subsp. latifolia, and Populus tremuloides. Burn severity had strong positive effects on seed germination and net seedling establishment after 3 years. Growth of transplanted seedlings was also significantly higher on severely burned soils. Our data and a synthesis of the literature indicated a consistent, steep decline in conifer establishment on organic soils at depths greater than 2.5 cm. A meta-analysis of seedling responses found no difference in the magnitude of severity effects on germination versus net establishment. There were, however, significant differences in establishment but not germination responses among deciduous trees, spruce, and pine, suggesting that small-seeded species experience greater mortality on lightly burned, organic soils than large-seeded species. Together, our analyses indicate that variations in burn severity can influence multiple aspects of forest stand structure, by affecting the density and composition of tree seedlings that establish after fire. These effects are predicted to be most important in moderately-drained forest stands, where a high potential variability in soil burn severity is coupled with strong severity effects on tree recruitment.     

Archaeal Communities in Boreal Forest Tree Rhizospheres Respond to Changing Soil Temperatures

Temperature has generally great effects on both the activity and composition of microbial communities in different soils. We tested the impact of soil temperature and three different boreal forest tree species on the archaeal populations in the bulk soil, rhizosphere, and mycorrhizosphere. Scots pine, silver birch, and Norway spruce seedlings were grown in forest humus microcosms at three different temperatures, 7–11.5°C (night–day temperature), 12–16°C, and 16–22°C, of which 12–16°C represents the typical mid-summer soil temperature in Finnish forests. RNA and DNA were extracted from indigenous ectomycorrhiza, non-mycorrhizal long roots, and boreal forest humus and tested for the presence of archaea by nested PCR of the archaeal 16S rRNA gene followed by denaturing gradient gel electrophoresis (DGGE) profiling and sequencing. Methanogenic Euryarchaeota belonging to Methanolobus sp. and Methanosaeta sp. were detected on the roots and mycorrhiza. The most commonly detected archaeal 16S rRNA gene sequences belonged to group I.1c Crenarchaeota, which are typically found in boreal and alpine forest soils. Interestingly, also one sequence belonging to group I.1b Crenarchaeota was detected from Scots pine mycorrhiza although sequences of this group are usually found in agricultural and forest soils in temperate areas. Tree- and temperature-related shifts in the archaeal population structure were observed. A clear decrease in crenarchaeotal DGGE band number was seen with increasing temperature, and correspondingly, the number of euryarchaeotal DGGE bands, mostly methanogens, increased. The greatest diversity of archaeal DGGE bands was detected in Scots pine roots and mycorrhizas. No archaea were detected from humus samples from microcosms without tree seedling, indicating that the archaea found in the mycorrhizosphere and root systems were dependent on the plant host. The detection of archaeal 16S rRNA gene sequences from both RNA and DNA extractions show that the archaeal populations were living and that they may have significant contribution to the methane cycle in boreal forest soil, especially when soil temperatures rise.     

Patterns of DNA Barcode Variation in Canadian Marine Molluscs

Background

Molluscs are the most diverse marine phylum and this high diversity has resulted in considerable taxonomic problems. Because the number of species in Canadian oceans remains uncertain, there is a need to incorporate molecular methods into species identifications. A 648 base pair segment of the cytochrome c oxidase subunit I gene has proven useful for the identification and discovery of species in many animal lineages. While the utility of DNA barcoding in molluscs has been demonstrated in other studies, this is the first effort to construct a DNA barcode registry for marine molluscs across such a large geographic area.

Methodology/Principal Findings

This study examines patterns of DNA barcode variation in 227 species of Canadian marine molluscs. Intraspecific sequence divergences ranged from 0–26.4% and a barcode gap existed for most taxa. Eleven cases of relatively deep (>2%) intraspecific divergence were detected, suggesting the possible presence of overlooked species. Structural variation was detected in COI with indels found in 37 species, mostly bivalves. Some indels were present in divergent lineages, primarily in the region of the first external loop, suggesting certain areas are hotspots for change. Lastly, mean GC content varied substantially among orders (24.5%–46.5%), and showed a significant positive correlation with nearest neighbour distances.

Conclusions/Significance

DNA barcoding is an effective tool for the identification of Canadian marine molluscs and for revealing possible cases of overlooked species. Some species with deep intraspecific divergence showed a biogeographic partition between lineages on the Atlantic, Arctic and Pacific coasts, suggesting the role of Pleistocene glaciations in the subdivision of their populations. Indels were prevalent in the barcode region of the COI gene in bivalves and gastropods. This study highlights the efficacy of DNA barcoding for providing insights into sequence variation across a broad taxonomic group on a large geographic scale.     

Effect of Tree Species and Mycorrhizal Colonization on the Archaeal Population of Boreal Forest Rhizospheres

Group 1.1c Crenarchaeota are the predominating archaeal group in acidic boreal forest soils. In this study, we show that the detection frequency of 1.1c crenarchaeotal 16S rRNA genes in the rhizospheres of the boreal forest trees increased following colonization by the ectomycorrhizal fungus Paxillus involutus. This effect was very clear in the fine roots of Pinus sylvestris, Picea abies, and Betula pendula, the most common forest trees in Finland. The nonmycorrhizal fine roots had a clearly different composition of archaeal 16S rRNA genes in comparison to the mycorrhizal fine roots. In the phylogenetic analysis, the 1.1c crenarchaeotal 16S rRNA gene sequences obtained from the fine roots formed a well-defined cluster separate from the mycorrhizal ones. Alnus glutinosa differed from the other trees by having high diversity and detection levels of Crenarchaeota both on fine roots and on mycorrhizas as well as by harboring a distinct archaeal flora. The similarity of the archaeal populations in rhizospheres of the different tree species was increased upon colonization by the ectomycorrhizal fungus. A minority of the sequences obtained from the mycorrhizas belonged to Euryarchaeota (order Halobacteriales).     

Variation in Seedling Density and Seed Predation Indicators for the Emergent Tree Dipteryx panamensis in Continuous and Fragmented Rain Forest1

Seedling density and the condition of stony endocarps of the tree Dipteryx panamensis were assessed in protected continuous forest and two forest fragments exposed to hunting and selective logging. Seedling density was higher in forest fragments than in continuous forest, while more whole endocarps and fewer chewed and half endocarps were found in fragments, indicating lower seed predation at fragment sites. These findings appear to contradict two earlier D. panamensis studies and we discuss methodological differences that could account for our disparate results. Hunting and fragmentation effects on mammal populations are suggested as a cause for the altered recruitment pattern in fragments.     

Developmental Patterns of Tree Dimensions in a Neotropical Deciduous Forest1

For 26 tree species in very dry tropical forest in Mexico, the developmental trends of relationships among trunk diameter, tree height, and crown diameter were inferred from a one‐time measurement of dispersed individuals across the size range from saplings to large, mature trees. On hillside sites in this high diversity forest, maximum dimensions were usually <10‐m height, 4‐m crown diameter, and 0.3‐m trunk diameter. The relationship of height to trunk diameter was characterized by an asymptotic, three‐parameter model. Crown diameter was a linear function of trunk diameter. The parameter values for both models varied widely among the species. In general, the dispersion among species of the height–crown diameter relationship increased linearly with trunk diameter (up to 0.2 m). Arborescent cacti were distant from other species at all sizes, although they were well modeled using the same equations. Empirical and theoretical features and limitations of the present and previous models, including mechanical buckling and water‐stress theories, are considered.     

Tree Mortality Following Boreal Forest Fires Reveals Scale-Dependant Interactions Between Community Structure and Fire Intensity

Fire disturbance patterns influence forest communities at a range of spatial scales. Forest community structure may also influence fire disturbance patterns, because tree species vary in their fuel value and in their tolerance to fire damage. However, the influence of community structure on fire disturbance likely depends on latent ecological differences between fires and on the spatial scale at which patterns are observed. Using data on fire intensity, community structure, and post-fire tree survival in four systematically sampled boreal forest fires, we tested the hypotheses that: (1) patterns in post-fire tree survival reflect interactions between fire intensity and community structure; (2) these relationships change with the spatial scale of observation. To test the first hypothesis, we used information theoretic methods to compare eight generalized linear mixed effects models describing the influence of community structure and fire intensity on tree survival in a 500 m² sample plot, accounting for latent fire-to-fire differences in response. To test the scaling hypothesis, we reaveraged the data at nine successively larger spatial resolutions up to approximately 2 km², at each resolution tracking the parameter values of the best model. When fit to the plot-level data, the dominant feature of the best model was a strong intensity–survival correlation which varied from fire to fire, and depended on plot-level community structure. In some fires, community structure and survival became more tightly coupled at larger scales, whereas fire intensity became less important. These results support the view that fire disturbance patterns are influenced by cross-scale interactions between community structure and fire intensity.     

中国北方林生产力变化趋势及其影响因子分析

森林生产力是反映森林固碳能力的重要指标,是进行碳循环研究的重要环节。用模拟生态系统生物地球化学循环的CENTURY模型,模拟中国北方林(兴安落叶松林)近35a来的生产力动态,用3种趋势分析方法,检验了其变化趋势,并用多元线性回归模型分析了中国北方林生产力的年际波动与气温降水年际波动的关系,以及气温和降水对我国北方林生产力的影响程度。结果表明：中国北方林生产力呈增加的趋势,平均年增长率为0．34％;气温与森林生产力呈显著负相关,对森林生产力的贡献因子为4．0977;降水与森林生产力呈弱的正相关,其对森林生产力的贡献因子为0．3902。从而说明近35a来森林生产力的增加除了受气温降水等非生物因素的影响外,还受其它因素的影响;另外说明以气候变暖为标志的全球变化会对森林生产力产生重要的影响。     

Effects of Soil Abiotic and Biotic Factors on Tree Seedling Regeneration Following a Boreal Forest Wildfire

Ecosystems - Wildfire disturbance is important for tree regeneration in boreal ecosystems. A considerable amount of literature has been published on how wildfires affect boreal forest regeneration....     

Seasonal Variation in Seed Dispersal by Tamarins Alters Seed Rain in a Secondary Rain Forest

Reduced dispersal of large seeds into degraded areas is one of the major factors limiting rain forest regeneration, as many seed dispersers capable of transporting large seeds avoid these sites with a limited forest cover. However, the small size of tamarins allows them to use small trees, and hence to disperse seeds into young secondary forests. Seasonal variations in diet and home range use might modify their contribution to forest regeneration through an impact on the seed rain. For a 2-yr period, we followed a mixed-species group of tamarins in Peru to determine how their role as seed dispersers in a 9-yr-old secondary-growth forest varied across seasons. These tamarins dispersed small to large seeds of 166 tree species, 63 of which were into a degraded area. Tamarins’ efficiency in dispersing seeds from primary to secondary forest varied across seasons. During the late wet season, high dietary diversity and long forays in secondary forest allowed them to disperse large seeds involved in later stages of regeneration. This occurred precisely when tamarins spent a more equal amount of time eating a high diversity of fruit species in primary forest and pioneer species in secondary forest. We hypothesized that well-balanced fruit availability induced the movement of seed dispersers between these 2 habitats. The noteworthy number of large-seeded plant species dispersed by such small primates suggests that tamarins play an important, but previously neglected, role in the regeneration and maintenance of forest structure.     

Patterns of Seed Size,Germination and Seed Viability of Tropical Tree Species in Southern India1

Seed weight, days to germination and seed viability were observed for 99 species growing in the Western Ghats of Karnataka, India. Seed size was strongly correlated with days to germination; smaller seeds germinated faster than larger seeds. Species which flowered during the rainy season had lighter seeds than species which flowered during the dry season. It was also found that seed size and viability of seeds were related to the season of fruiting. Species which fruit during the rainy season had heavier seeds and shorter viability than species which fruit during the dry season. These flowering and fruiting patterns and varying seed sizes are argued to be adaptations to the time of dispersal, time of moisture availability in the habitat and seedling survival.     

A novel stochastic method for reconstructing daily precipitation times-series using tree-ring data from the western Canadian Boreal Forest

Tree ring data provide proxy records of historical hydroclimatic conditions that are widely used for reconstructing precipitation time series. Most previous applications are limited to annual time scales, though information about daily precipitation would enable a range of additional analyses of environmental processes to be investigated and modelled. We used statistical downscaling to simulate stochastic daily precipitation ensembles using dendrochronological data from the western Canadian boreal forest. The simulated precipitation series were generally consistent with observed precipitation data, though reconstructions were poorly constrained during short periods of forest pest outbreaks. The proposed multiple temporal scale precipitation reconstruction can generate annual daily maxima and persistent monthly wet and dry episodes, so that the observed and simulated ensembles have similar precipitation characteristics (i.e. magnitude, peak, and duration)—an improvement on previous modelling studies. We discuss how ecological disturbances may limit reconstructions by inducing non-linear responses in tree growth, and conclude with suggestions of possible applications and further development of downscaling methods for dendrochronological data.     

Tree Diversity Explains Variation in Ecosystem Function in a Neotropical Forest in Panama

Many experimental studies show that a decline in species number has a negative effect on ecosystem function, however less is known about this pattern in natural communities. We examined the relative importance of environment, space, and diversity on ecosystem function, specifically tree carbon storage in four plant types (understory/canopy; trees/palms), in a tropical forest in central Panama. The objectives of this study were to detect the relationship between tree diversity and carbon storage given the environmental and spatial variation that occur in natural forests and to determine which species diversity measure is more important to tree carbon storage: richness or dominance. We used redundancy analyses to partition the effect of these sources of variation on tree carbon storage. We showed that together, environment, space, and diversity accounted for 43 percent of tree carbon storage, where diversity (19%) alone is the most important source of variation and explained more variation than space (13%) and environment (1%) together. Therefore, even in natural forests where substantial environment and spatial variation can be found, it is still possible to detect the effect of diversity on ecosystem function at scales relevant to conservation. Moreover, both richness and dominance are important to explain the variation on tree carbon storage in natural forests suggesting that these two diversity measures are complementary. Thus, tree diversity is important to predict tree carbon storage in hyperdiverse forests.     

Nutrient and Carbon Additions to the Microbial Soil Community and its Impact on Tree Seedlings in a Boreal Spruce Forest

We have added glucose and nutrients to manipulate soil microbial activity and nutrient availability in a boreal spruce forest to study the performance of birch and spruce seedlings in relation to the soil microbial community. The proportion of aboveground biomass in the seedlings was largest in plots amended with extra nutrients, while ectomycorrhizal (ECM) colonisation was low in these plots. ECM appeared beneficial for growth of both species, but only at low levels of colonisation (<25% ECM colonised root-tips). The soil microbial biomass, as determined by total PLFA, was largest in plots treated with glucose and there was a significant negative relationship between birch seedling size and levels of total PLFA in soil. This could be taken to suggest that poor seedling growth was due to nutrient limitation caused by microbial assimilation. However, the treatment response of the birch seedlings was generally weak, and spruce often showed no response at all to the addition of nutrients and glucose. The most consistent parameter for the variation in plant performance, as well as for the microbial soil community, was the block-effect. This suggests a strong spatial structure in the soil microbial community, and that this structure was robust with respect to our treatments even though they continued over a 3-year period.     

香港次生林树种种子生理生态学特性的研究

１９８９～１９９１年在温室和野外测定了香港４０种次生林树木种类的种子寿命和发芽率．结果显示，在所测试的４０种天然林树种中，６０％的树种的种子寿命短于１年，仅有８个树种的种子寿命在１年以上．种子的寿命与种子在土壤中的发芽能力呈负相关的关系．种子发芽测定结果表明，大多数树种都可以在强光照条件（９５％的日光）下发芽，但３５％的测试树种不能在天然林下（０７％～２％的日光）发芽．最后从植物生理生态的角度，探讨种子特征与香港次生林区系发展及香港现有次生林的演替现状     

Spatiotemporal Variations of Fire Frequency in Central Boreal Forest

Determination of the direct causal factors controlling wildfires is key to understanding wildfire–vegetation–climate dynamics in a changing climate and for developing sustainable management strategies for biodiversity conservation and maintenance of long-term forest productivity. In this study, we sought to understand how the fire frequency of a large mixedwood forest in the central boreal shield varies as a result of temporal and spatial factors. We reconstructed the fire history of an 11,600-km² area located in the northwestern boreal forest of Ontario, using archival data of large fires occurring since 1921 and dendrochronological dating for fires prior to 1921. The fire cycle decreased from 295 years for the period of 1820–1920 to approximately 100 years for the period of 1921–2008. Spatially, fire frequency increased with latitude, attributable to higher human activities that have increased fragmentation and fire suppression in the southern portion of the study area. Fire frequency also increased with distance to waterbodies, and was higher on Podzols that were strongly correlated with moderate drainage and coniferous vegetation. The temporal increase of fire frequency in the central region, unlike western and eastern boreal forests where fire frequency has decreased, may be a result of increased warm and dry conditions associated with climate change in central North America, suggesting that the response of wildfire to global climate change may be regionally individualistic. The significant spatial factors we found in this study are in agreement with other wildfire studies, indicating the commonality of the influences by physiographic features and human activities on regional fire regimes across the boreal forest. Overall, wildfire in the central boreal shield is more frequent than that in the wetter eastern boreal region and less frequent than that in the drier western boreal region, confirming a climatic top-down control on the fire activities of the entire North American boreal forest.