Han Y. H. Chen

Understory Vegetation Dynamics of North American Boreal Forests

Understory vegetation is the most diverse and least understood component of North American boreal forests. Understory communities are important as they act as drivers of overstory succession and nutrient cycling. The objective of this review was to examine how understory vegetation abundance, composition, and diversity change with stand development after a major stand replacing disturbance. Understory vegetation abundance and diversity increase rapidly after fire, in response to abundant resources and an influx of disturbance adapted species. The highest diversity occurs within the first 40 years following fire, and declines indefinitely thereafter as a result of decreasing productivity and increased dominance of a small number of late successional feather mosses and woody plant species. Vascular plant and bryophyte/lichen communities undergo very different successional changes. Vascular plant communities are dynamic and change more dramatically with time after fire, whereas bryophyte and lichen communities are much slower to establish and change over time. Considerable variations in these processes exist depending on canopy composition, site condition, regional climate, and frequently occurring non-stand-replacing disturbances. Forest management practices represent a unique disturbance process and can result in different understory vegetation communities from those observed for natural processes, with potential implications for overstory succession and long-term productivity. Because of the importance of understory vegetation on nutrient cycling and overstory composition, post-harvest treatments emulating stand-replacing fire are required to maintain understory diversity, composition, and promote stand productivity in boreal forests.

Fine Root Biomass, Production, Turnover Rates, and Nutrient Contents in Boreal Forest Ecosystems in Relation to Species, Climate, Fertility, and Stand Age: Literature Review and Meta-Analyses

Fine roots <2 mm in diameter play a key role in regulating the biogeochemical cycles of ecosystems and are important to our understanding of ecosystem responses to global climate changes. Given the sensitivity of fine roots, especially in boreal region, to climate changes, it is important to assess whether and to what extent fine roots in this region change with climates. Here, in this synthesis, a data set of 218 root studies were complied to examine fine root patterns in the boreal forest in relation to site and climatic factors. The mean fine root biomass in the boreal forest was 5.28 Mg ha−1, and the production of fine roots was 2.82 Mg ha−1 yr−1, accounting for 32% of annual net primary production of the boreal forest. Fine roots in the boreal forest on average turned over 1.07 times per year. Fine roots contained 50.9 kg ha−1 of nitrogen (N) and 3.63 kg ha−1 of phosphorous (P). In total, fine roots in the boreal forest ecosystems contain 6.1 × 107 Mg N and 4.4×106Mg P pools, respectively, about 10% of the global nutrients of fine roots. Fine root biomass, production, and turnover rate generally increased with increasing mean annual temperature and precipitation. Fine root biomass in the boreal forest decreased significantly with soil N and P availability. With increasing stand age, fine root biomass increased until about 100 years old for forest stands and then leveled off or decreased thereafter. These results of meta analysis suggest that environmental factors strongly influence fine root biomass, production, and turnover in boreal forest, and future studies should place a particular emphasis on the root-environment relationships.

Positive biodiversity-productivity relationship predominant in global forests.

Global biodiversity and productivity The relationship between biodiversity and ecosystem productivity has been explored in detail in herbaceous vegetation, but patterns in forests are far less well understood. Liang et al. have amassed a global forest data set from >770,000 sample plots in 44 countries. A positive and consistent relationship can be discerned between tree diversity and ecosystem productivity at landscape, country, and ecoregion scales. On average, a 10% loss in biodiversity leads to a 3% loss in productivity. This means that the economic value of maintaining biodiversity for the sake of global forest productivity is more than fivefold greater than global conservation costs. Science, this issue p. 196 Global forest inventory records suggest that biodiversity loss would result in a decline in forest productivity worldwide. INTRODUCTION The biodiversity-productivity relationship (BPR; the effect of biodiversity on ecosystem productivity) is foundational to our understanding of the global extinction crisis and its impacts on the functioning of natural ecosystems. The BPR has been a prominent research topic within ecology in recent decades, but it is only recently that we have begun to develop a global perspective. RATIONALE Forests are the most important global repositories of terrestrial biodiversity, but deforestation, forest degradation, climate change, and other factors are threatening approximately one half of tree species worldwide. Although there have been substantial efforts to strengthen the preservation and sustainable use of forest biodiversity throughout the globe, the consequences of this diversity loss pose a major uncertainty for ongoing international forest management and conservation efforts. The forest BPR represents a critical missing link for accurate valuation of global biodiversity and successful integration of biological conservation and socioeconomic development. Until now, there have been limited tree-based diversity experiments, and the forest BPR has only been explored within regional-scale observational studies. Thus, the strength and spatial variability of this relationship remains unexplored at a global scale. RESULTS We explored the effect of tree species richness on tree volume productivity at the global scale using repeated forest inventories from 777,126 permanent sample plots in 44 countries containing more than 30 million trees from 8737 species spanning most of the global terrestrial biomes. Our findings reveal a consistent positive concave-down effect of biodiversity on forest productivity across the world, showing that a continued biodiversity loss would result in an accelerating decline in forest productivity worldwide. The BPR shows considerable geospatial variation across the world. The same percentage of biodiversity loss would lead to a greater relative (that is, percentage) productivity decline in the boreal forests of North America, Northeastern Europe, Central Siberia, East Asia, and scattered regions of South-central Africa and South-central Asia. In the Amazon, West and Southeastern Africa, Southern China, Myanmar, Nepal, and the Malay Archipelago, however, the same percentage of biodiversity loss would lead to greater absolute productivity decline. CONCLUSION Our findings highlight the negative effect of biodiversity loss on forest productivity and the potential benefits from the transition of monocultures to mixed-species stands in forestry practices. The BPR we discover across forest ecosystems worldwide corresponds well with recent theoretical advances, as well as with experimental and observational studies on forest and nonforest ecosystems. On the basis of this relationship, the ongoing species loss in forest ecosystems worldwide could substantially reduce forest productivity and thereby forest carbon absorption rate to compromise the global forest carbon sink. We further estimate that the economic value of biodiversity in maintaining commercial forest productivity alone is

FIRE, LOGGING, AND OVERSTORY AFFECT UNDERSTORY ABUNDANCE, DIVERSITY, AND COMPOSITION IN BOREAL FOREST

166 billion to

Stand Structural Dynamics of North American Boreal Forests

490 billion per year. Although representing only a small percentage of the total value of biodiversity, this value is two to six times as much as it would cost to effectively implement conservation globally. These results highlight the necessity to reassess biodiversity valuation and the potential benefits of integrating and promoting biological conservation in forest resource management and forestry practices worldwide. Global effect of tree species diversity on forest productivity. Ground-sourced data from 777,126 global forest biodiversity permanent sample plots (dark blue dots, left), which cover a substantial portion of the global forest extent (white), reveal a consistent positive and concave-down biodiversity-productivity relationship across forests worldwide (red line with pink bands representing 95% confidence interval, right). The biodiversity-productivity relationship (BPR) is foundational to our understanding of the global extinction crisis and its impacts on ecosystem functioning. Understanding BPR is critical for the accurate valuation and effective conservation of biodiversity. Using ground-sourced data from 777,126 permanent plots, spanning 44 countries and most terrestrial biomes, we reveal a globally consistent positive concave-down BPR, showing that continued biodiversity loss would result in an accelerating decline in forest productivity worldwide. The value of biodiversity in maintaining commercial forest productivity alone—US

Influence of Environmental Variability on Root Dynamics in Northern Forests

166 billion to 490 billion per year according to our estimation—is more than twice what it would cost to implement effective global conservation. This highlights the need for a worldwide reassessment of biodiversity values, forest management strategies, and conservation priorities.

Is understory plant species diversity driven by resource quantity or resource heterogeneity

Understory vegetation plays a critical role in boreal ecosystems. Despite this, quantitative evaluation of the factors controlling understory vegetation abundance, diversity, and composition in the most diverse boreal forest region in North America is lacking. This study examined the dynamics of understory vegetation of stands of fire origin and tested effects of overstory composition and logging vs. fire on the understory vegetation dynamics in Ontario, Canada. Understory vegetation communities were sampled in 68 stands of conifer, mixed-wood, and deciduous overstory type ranging from 7 to 201 years postfire for stands of fire origin, and from 7 to 31 years for stands of logging origin. For stands of fire origin, total cover and species richness followed similar trends for the three overstory types and were highest in the intermediate-aged stands (72-90 years). Trends in cover and richness, however, differed significantly for vascular and nonvascular plant groups. Vascular cover and species richness were generally higher under deciduous stands, and lower on older stands, while nonvascular species richness was highest under conifer stands and increased with time since fire. Neither species richness nor compositional turnover was higher under mixed-wood stands; mixed-wood stands were compositionally intermediate to conifer and deciduous stands. Multivariate analysis using multiple-response permutation procedures indicated that understory communities were compositionally distinct for all overstory types and showed no convergence with increasing time since fire. Compared with postfire stands of similar ages, post-logged stands had similar total understory cover and richness. Vascular cover and richness, however, were higher on post-logged stands, and nonvascular cover and richness were lower. Stands of logging and fire origin were compositionally distinct for all overstory types and ages. Compositional differences appeared to be driven by higher preestablished rhizomatous species and few pyrophilic species on post-logged sites. Understory vegetation communities in the central boreal shield appear to support the intermediate disturbance hypothesis. Understory richness, however, was not negatively associated with high cover values as predicted by the intermediate disturbance hypothesis. Moreover, richness appears to be highest on sites with high light availability, suggesting that boreal understory communities are influenced more by plant tolerances for low resources, rather than by competition.

Tree species diversity increases fine root productivity through increased soil volume filling

Stand structure, the arrangement and interrelationships of live and dead trees, has been linked to forest regeneration, nutrient cycling, wildlife habitat, and climate regulation. The objective of this review was to synthesize literature on stand structural dynamics of North American boreal forests, addressing both live tree and coarse woody debris (CWD) characteristics under different disturbance mechanisms (fire, clearcut, wind, and spruce budworm), while identifying regional differences based on climate and surficial deposit variability. In fire origin stands, both live tree and CWD attributes are influenced initially largely by the characteristics of the stand replacing fire and later increasingly by autogenic processes. Differences in stand structure have also been observed between various stand cover types. Blowdown and insect outbreaks are two significant non-stand replacing disturbances that can alter forest stand structure through removing canopy trees, freeing up available growing space, and creating microsites for new trees to establish. Climate and surficial deposits are highly variable in the boreal forest due to its extensive geographic range, influencing stand and landscape structure by affecting tree colonization, stand composition, successional trajectories, CWD dynamics, and disturbance regimes including regional fire cycles. Further, predicted climate change scenarios are likely to cause regional-specific alterations in stand and landscape structure, with the implications on ecosystem components including wildlife, biodiversity, and carbon balance still unclear. Some stand structural attributes are found to be similar between clearcut and fire origin stands, but others appear to be quite different. Future research shall focus on examining structural variability under both disturbance regimes and management alternatives emulating both stand replacing and non-stand replacing natural disturbances. Referee: Professor Andrew Gordon, Department of Environmental Biology, Ontario Agricultural College, University of Guelph, Guelph, ON N1G 2W1 Canada

Exposure to ambient black carbon derived from a unique inventory and high-resolution model

Plant root systems are highly dynamic over various temporal and spatial scales, and are responsive to changes in environmental conditions. The objective of this review is to describe the dynamics of root structure and function in boreal and northern temperate forests and examine how edaphic and climatic characteristics and intra- and interspecific root competition impact root dynamics. Fine roots exhibit distinct seasonal trends of production and mortality. Over the extent of stand development, coarse root biomass increases until maturity, while the response of fine roots remains unclear. Roots are predominantly restricted to the upper soil layers, and spatial variability of roots in the horizontal direction decreases with decreasing root size. Root/shoot ratio decreases gradually through stand development. On nutrient-rich sites, roots are more concentrated around respective stems and root systems are more branched than on nutrient-poor sites. Plants generally root deeper under low soil moisture conditions, while roots tend to grow horizontally into rich rather than poor patches of soil. Plants adapt their biomass allocation strategies to edaphic and climatic variation according to the functional equilibrium hypothesis. Although root production is projected to increase, providing nutrients are not limiting, following elevated carbon dioxide concentrations and temperatures, how root turnover and above- and below-ground carbon allocation may change remains uncertain. Stands composed of species with different rooting characteristics may attain greater root production compared to single-species stands or mixtures of similar species from reduced exploitative competition. Alternatively, plants can produce greater root biomass with a competing plant than growing alone as a result of self root discrimination. Future research is needed to address how elevated carbon dioxide concentrations and temperatures will feedback upon soil resource availability to influence plant responses from the organism- to the ecosystem-level.

Biomass offsets little or none of permafrost carbon release from soils, streams, and wildfire: an expert assessment

What maintains plant species diversity has been the subject of much debate with no general consensus. In forest ecosystems in which understory plants account for the majority of floristic diversity, a crucial question is whether understory plant diversity is driven by resource quantity or resource heterogeneity. This study sought to reconcile the two hypotheses in relation to their effects on understory plant diversity in forest ecosystems. A database of studies that investigated the effects of resources on understory plant diversity was compiled and analyzed using log-linear models. Whether resource quantity or resource heterogeneity is the determinant of understory plant diversity in individual studies was dependent on stand successional stage(s), presence or absence of intermediate disturbance, and forest biome within which the studies were conducted. Resource quantity was found to govern species diversity in both young and mature stands, whereas resource heterogeneity dominated in old-growth stands. Resource quantity remained the important driver in both disturbed and undisturbed forests, but resource heterogeneity played an important role in disturbed forests. We argue that neither resource quantity nor heterogeneity alone structures species diversity in forest ecosystems, but rather their influences on understory plant diversity vary with stand development and disturbances in forest ecosystems.