Jennifer L. Baltzer

EDAPHIC SPECIALIZATION IN TROPICAL TREES: PHYSIOLOGICAL CORRELATES AND RESPONSES TO RECIPROCAL TRANSPLANTATION

Recent research has documented the importance of edaphic factors in de- termining the habitat associations of tree species in many tropical rain forests, but the underlying mechanisms for edaphic associations are unclear. At Sepilok Forest Reserve, Sabah, Malaysian Borneo, two main soil types derived from sandstone (ridges) and alluvium (valleys) differ in nutrient and water availability and are characterized by forests differing markedly in species composition, structure, and understory light availability. We use both survey and reciprocal transplants to examine physiological adaptations to differences in light, nutrient, and water availability between these soil types, and test for the importance of resource-use efficiency in determining edaphic specialization. Photosynthetic surveys for congeneric and confamilial pairs (one species per soil type) of edaphic specialists and for generalists common to both soil types show that species specializing on sandstone- derived soil had lower stomatal conductance at a given assimilation rate than those occurring on alluvial soil and also had greater instantaneous and integrated water-use efficiencies. Foliar dark respiration rates per unit photosynthesis were higher for sandstone ridge than alluvial lowland specialists. We suggest that these higher respiration rates are likely due to increases in photosynthetic enzyme concentrations to compensate for lower internal CO 2 concentrations resulting from increased stomatal closure. This is supported by lower pho- tosynthetic nitrogen-use efficiencies in the sandstone ridge specialists. Generalist species had lower water-use efficiencies than sandstone ridge specialists when growing on the drier, sandy ridgetops, but their nitrogen-use efficiencies did not differ from the species specialized to the more resource-rich alluvial valleys. We varied light environment and soil nutrient availability in a reciprocal transplant experiment involving two specialist species from each soil type. Edaphic specialist species, when grown on the soil type for which they were not specialized, were not capable of acclimatory shifts to achieve similar resource-use effi- ciencies as species specialized to that soil type. We conclude that divergent water-use strategies are an important mechanism underlying differences in edaphic associations and thus contributing to maintenance of high local tree species diversity in Bornean rain forests.

BAAD: a biomass and allometry database for woody plants

Understanding how plants are constructed—i.e., how key size dimensions and the amount of mass invested in different tissues varies among individuals—is essential for modeling plant growth, carbon stocks, and energy fluxes in the terrestrial biosphere. Allocation patterns can differ through ontogeny, but also among coexisting species and among species adapted to different environments. While a variety of models dealing with biomass allocation exist, we lack a synthetic understanding of the underlying processes. This is partly due to the lack of suitable data sets for validating and parameterizing models. To that end, we present the Biomass And Allometry Database (BAAD) for woody plants. The BAAD contains 259 634 measurements collected in 176 different studies, from 21 084 individuals across 678 species. Most of these data come from existing publications. However, raw data were rarely made public at the time of publication. Thus, the BAAD contains data from different studies, transformed into standard units and variable names. The transformations were achieved using a common workflow for all raw data files. Other features that distinguish the BAAD are: (i) measurements were for individual plants rather than stand averages; (ii) individuals spanning a range of sizes were measured; (iii) plants from 0.01–100 m in height were included; and (iv) biomass was estimated directly, i.e., not indirectly via allometric equations (except in very large trees where biomass was estimated from detailed sub-sampling). We included both wild and artificially grown plants. The data set contains the following size metrics: total leaf area; area of stem cross-section including sapwood, heartwood, and bark; height of plant and crown base, crown area, and surface area; and the dry mass of leaf, stem, branches, sapwood, heartwood, bark, coarse roots, and fine root tissues. We also report other properties of individuals (age, leaf size, leaf mass per area, wood density, nitrogen content of leaves and wood), as well as information about the growing environment (location, light, experimental treatment, vegetation type) where available. It is our hope that making these data available will improve our ability to understand plant growth, ecosystem dynamics, and carbon cycling in the worlds vegetation.

Linear disturbances on discontinuous permafrost: implications for thaw-induced changes to land cover and drainage patterns

Within the zone of discontinuous permafrost, linear disturbances such as winter roads and seismic lines severely alter the hydrology, ecology, and ground thermal regime. Continued resource exploration in this environment has created a need to better understand the processes causing permafrost thaw and concomitant changes to the terrain and ground cover, in order to efficiently reduce the environmental impact of future exploration through the development of best management practices. In a peatland 50 km south of Fort Simpson, NWT, permafrost thaw and the resulting ground surface subsidence have produced water-logged linear disturbances that appear not to be regenerating permafrost, and in many cases have altered the land cover type to resemble that of a wetland bog or fen. Subsidence alters the hydrology of plateaus, developing a fill and spill drainage pattern that allows some disturbances to be hydrologically connected with adjacent wetlands via surface flow paths during periods of high water availability. The degree of initial disturbance is an important control on the extent of permafrost thaw and thus the overall potential recovery of the linear disturbance. Low impact techniques that minimize ground surface disturbance and maintain original surface topography by eliminating windrows are needed to minimize the impact of these linear disturbances.

Coordination of foliar and wood anatomical traits contributes to tropical tree distributions and productivity along the Malay-Thai Peninsula

Drought is a critical factor in plant species distributions. Much research points to its relevance even in moist tropical regions. Recent studies have begun to elucidate mechanisms underlying the distributions of tropical tree species with respect to drought; however, how such desiccation tolerance mechanisms correspond with the coordination of hydraulic and photosynthetic traits in determining species distributions with respect to rainfall seasonality deserves attention. In the present study, we used a common garden approach to quantify inherent differences in wood anatomical and foliar physiological traits in 21 tropical tree species with either widespread (occupying both seasonal and aseasonal climates) or southern (restricted to aseasonal forests) distributions with respect to rainfall seasonality. Use of congeneric species pairs and phylogenetically independent contrast analyses allowed examination of this question in a phylogenetic framework. Widespread species opted for wood traits that provide biomechanical support and prevent xylem cavitation and showed associated reductions in canopy productivity and consequently growth rates compared with southern species. These data support the hypothesis that species having broader distributions with respect to climatic variability will be characterized by traits conducive to abiotic stress tolerance. This study highlights the importance of the well-established performance vs. stress tolerance trade-off as a contributor to species distributions at larger scales.

Rainfall seasonality and pest pressure as determinants of tropical tree species' distributions.

Drought and pests are primary abiotic and biotic factors proposed as selective filters acting on species distributions along rainfall gradients in tropical forests and may contribute importantly to species distributional limits, performance, and diversity gradients. Recent research demonstrates linkages between species distributions along rainfall gradients and physiological drought tolerance; corresponding experimental examinations of the contribution of pest pressure to distributional limits and potential interactions between drought and herbivory are limited. This study aims to quantitate differential performance and herbivory as a function of species range limits across a climatic and floristic transition in Southeast Asia. Khao Chong Botanical Garden, Thailand and Pasoh Forest Reserve, Malaysia straddle the Kangar-Pattani Line. A reciprocal transplantation across a seasonality gradient was established using two groups of species (“widespread” taxa whose distributions include seasonally dry forests and “aseasonal” taxa whose distributions are limited to aseasonal forests). Growth, biomass allocation, survival, and herbivory were monitored for 19 months. Systematic differences in performance were a function of species distribution in relation to rainfall seasonality. In aseasonal Pasoh, aseasonal species had both greater growth and survivorship than widespread species. These differences were not a function of differential herbivory as widespread and aseasonal species experienced similar damage in the aseasonal forest. In seasonally dry Khao Chong, widespread species showed higher survivorship than aseasonal species, but these differences were only apparent during drought. We link this differential performance to physiological mechanisms as well as differential tolerance of biotic pressure during drought stress. Systematic decreases in seedling survival in aseasonal taxa during drought corresponded with previously documented physiological differences and may be exacerbated by herbivore damage. These results have important implications for tropical diversity and community composition in light of predicted increases in the frequency and severity of drought in hyperdiverse tropical forests.

Losing Legacies, Ecological Release, and Transient Responses: Key Challenges for the Future of Northern Ecosystem Science

Northern ecosystem processes play out across scales that are rare elsewhere on contemporary earth: large ranging predator–prey systems are still operational, invasive species are rare, and large-scale natural disturbances occur extensively. Disturbances in the far north affect huge areas of land and are difficult to control or manage. Historically, disturbance patterns and processes ranging across a number of spatio-temporal scales have played an important role in the resilience of northern ecosystems. However, due to interactions with a warming climate, these disturbances are now erasing key legacies of the last millennia of ecosystem processes. Building on the concepts of legacies and cross-scale interactions, we highlight several general conceptual issues that represent key challenges for the future of northern ecosystem science, but that also have relevance to other biomes.

Decreased competitive interactions drive a reverse species richness latitudinal gradient in subarctic forests

The tendency for species richness to decrease toward the poles is one of the best-characterized patterns in biogeography. The mechanisms behind this pattern have received much attention, yet very few studies have investigated very high-latitude communities. Here, using data from 134 permanent sample plots from 60 degrees to 68 degrees N, we show that boreal forest plant communities in northwestern Canada increase in richness toward the poles, despite a strong increase in climatic harshness. We hypothesized three possible explanations for this pattern: (1) historical biogeography, (2) reduced competition for light at high latitudes (biotic interactions), and (3) changes in soil characteristics with latitude. We used multidimensional scaling to investigate the community composition at each site and found no clustering of communities by latitude, suggesting that historical biogeography was not constraining site diversity. We then investigated the mechanisms behind this gradient using both abiotic (climate and soil) and biotic (tree stand characteristics) variables in a multiple factor analysis. We found that the best predictor of species richness is an environmental gradient that describes an inverse relationship between temperature and tree-stand density, suggesting that reduced competition for light due to reduced tree growth at low temperatures at higher latitudes allows greater species richness. This study shows that low energy availability and climatic harshness may not be limiting species richness toward the poles, rather, abiotic effects act instead on the strength of biotic interactions.

Soil organic layer combustion in boreal black spruce and jack pine stands of the Northwest Territories, Canada

Increased fire frequency, extent and severity are expected to strongly affect the structure and function of boreal forest ecosystems. In this study, we examined 213 plots in boreal forests dominated by black spruce (Picea mariana) or jack pine (Pinus banksiana) of the Northwest Territories, Canada, after an unprecedentedly large area burned in 2014. Large fire size is associated with high fire intensity and severity, which would manifest as areas with deep burning of the soil organic layer (SOL). Our primary objectives were to estimate burn depth in these fires and then to characterise landscapes vulnerable to deep burning throughout this region. Here we quantify burn depth in black spruce stands using the position of adventitious roots within the soil column, and in jack pine stands using measurements of burned and unburned SOL depths. Using these estimates, we then evaluate how burn depth and the proportion of SOL combusted varies among forest type, ecozone, plot-level moisture and stand density. Our results suggest that most of the SOL was combusted in jack pine stands regardless of plot moisture class, but that black spruce forests experience complete combustion of the SOL only in dry and moderately well-drained landscape positions. The models and calibrations we present in this study should allow future research to more accurately estimate burn depth in Canadian boreal forests.

The world's biomes and primary production as a triple tragedy of the commons foraging game played among plants

Plants appear to produce an excess of leaves, stems and roots beyond what would provide the most efficient harvest of available resources. One way to understand this overproduction of tissues is that excess tissue production provides a competitive advantage. Game theoretic models predict overproduction of all tissues compared with non-game theoretic models because they explicitly account for this indirect competitive benefit. Here, we present a simple game theoretic model of plants simultaneously competing to harvest carbon and nitrogen. In the model, a plants fitness is influenced by its own leaf, stem and root production, and the tissue production of others, which produces a triple tragedy of the commons. Our model predicts (i) absolute net primary production when compared with two independent global datasets; (ii) the allocation relationships to leaf, stem and root tissues in one dataset; (iii) the global distribution of biome types and the plant functional types found within each biome; and (iv) ecosystem responses to nitrogen or carbon fertilization. Our game theoretic approach removes the need to define allocation or vegetation type a priori but instead lets these emerge from the model as evolutionarily stable strategies. We believe this to be the simplest possible model that can describe plant production.

Contributions of C-Band SAR Data and Polarimetric Decompositions to Subarctic Boreal Peatland Mapping

The objective of this paper is to assess the accuracy of C-band synthetic aperture radar (SAR) datasets in mapping peatland types over a region of Canadas subarctic boreal zone. This paper assessed contributions of quad-polarization linear backscatter intensities (σ°HH, σ°HV, σ°VV), image textures, and two polarimetric scattering decompositions: 1) Cloude–Pottier, and 2) Freeman–Durden. Four quad-polarimetric RADADSAT-2 images were studied at incidence angles of 19.4°, 23.1°, 45.8°, and 48.1°. The influence of combining dual-angular information acquired within a short temporal span was also assessed. These C-band SAR data were used to classify peatlands according to isolated flat bogs (bogs), channel fens (fens), raised peat plateaus (plateaus), and forested uplands (uplands) using a supervised support vector machine (SVM) classifier. Numerous classifications were examined to compare the unique contributions of these variables to classification accuracy. Results suggest linear backscatter variables in isolation produce comparable classification results with those of the Freeman–Durden and Cloude–Pottier decompositions. Combining polarimetric decomposition and texture data into classifications with linear backscatter data resulted in only minor (∼1–3%) improvement. Combining classifications from small and large incidence angles (dual-angular) significantly improved classification results over those of a single image. Classification accuracy was the highest for isolated bogs and open water surfaces, whereas fens, uplands, and plateaus had lower accuracies. The highest accuracy classification (84% and kappa coefficient of 0.80) used a dual-angular approach, with additional decomposition and texture information. However, it is noted that texture information rarely improved classification results across all tests. This approach identified isolated flat bogs, channel fens, and raised peat plateaus with >76% producers accuracies.