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Rising atmospheric CO 2 may stimulate future forest productivity, possibly increasing carbon storage in terrestrial ecosystems, but how tropospheric ozone will modify this response is unknown. Because of the importance of fine roots to the belowground C cycle, we monitored fine-root biomass and associated C fluxes in regenerating stands of trembling aspen, and mixed stands of trembling aspen and paper birch at FACTS-II, the Aspen FACE project in Rhinelander, Wisconsin. Free-air CO 2 enrichment (FACE) was used to elevate concentrations of CO 2 (average enrichment concentration 535 µl l -1 ) and O 3 (53 nl l -1 ) in developing forest stands in 1998 and 1999. Soil respiration, soil pCO 2 , and dissolved organic carbon in soil solution (DOC) were monitored biweekly. Soil respiration was measured with a portable infrared gas analyzer. Soil pCO 2 and DOC samples were collected from soil gas wells and tension lysimeters, respectively, at depths of 15, 30, and 125 cm. Fine-root biomass averaged 263 g m -2 in control plots and increased 96% under elevated CO 2 . The increased root biomass was accompanied by a 39% increase in soil respiration and a 27% increase in soil pCO 2 . Both soil respiration and pCO 2 exhibited a strong seasonal signal, which was positively correlated with soil temperature. DOC concentrations in soil solution averaged ~12 mg l -1 in surface horizons, declined with depth, and were little affected by the treatments. A simplified belowground C budget for the site indicated that native soil organic matter still dominated the system, and that soil respiration was by far the largest flux. Ozone decreased the above responses to elevated CO 2 , but effects were rarely statistically significant. We conclude that regenerating stands of northern hardwoods have the potential for substantially greater C input to soil due to greater fine-root production under elevated CO 2 . Greater fine-root biomass will be accompanied by greater soil C efflux as soil respiration, but leaching losses of C will probably be unaffected.

Effects of CO 2 on plants are often confounded with effects of nutrition. Coleman et al. (1993) showed that nitrogen concentrations in plants grown under different CO 2 levels did not differ if plants of equal sizes were compared but varied at equal times and equal sizes for different levels of nitrogen additions. I will suggest why this must occur under the conditions of the experiment. I will also suggest why nitrogen productivity should be used to interpret experimental results rather than the dubious nitrogen use efficiency.

This study examined the independent and interactive effects of elevated carbon dioxide (CO) and ozone (O) on the foliar quality of two deciduous trees species and the performance of two outbreak herbivore species. Trembling aspen ( Populus tremuloides) and paper birch ( Betula papyrifera) were grown at the Aspen FACE research site in northern Wisconsin, USA, under four combinations of ambient and elevated CO and O. We measured the effects of elevated CO and O on aspen and birch phytochemistry and on gypsy moth ( Lymantria dispar) and forest tent caterpillar ( Malacosoma disstria) performance. Elevated CO nominally affected foliar quality for both tree species. Elevated O negatively affected aspen foliar quality, but only marginally influenced birch foliar quality. Elevated CO slightly improved herbivore performance, while elevated O decreased herbivore performance, and both responses were stronger on aspen than birch. Interestingly, elevated CO largely offset decreased herbivore performance under elevated O. Nitrogen, lignin, and C:N were identified as having strong influences on herbivore performance when larvae were fed aspen, but no significant relationships were observed for insects fed birch. Our results support the notion that herbivore performance can be affected by atmospheric change through altered foliar quality, but how herbivores will respond will depend on interactions among CO, O, and tree species. An emergent finding from this study is that tree age and longevity of exposure to pollutants may influence the effects of elevated CO and O on plant-herbivore interactions, highlighting the need to continue long-term atmospheric change research. [ABSTRACT FROM AUTHOR]

Light with decreased red:far-red (R:FR) ratios may signal neighbor presence and trigger plant developmental responses. There is some evidence that plant canopies forage towards increased R:FR ratios, but it is unclear to what extent R:FR versus the total amount of photosynthetically active radiation (PAR) influences canopy foraging responses among forest trees. The objective of this study was to examine the relative importance of PAR and R:FR as photosensory cues leading to tree canopy foraging responses. Seedlings of Betula papyrifera Marshall (paper birch) were grown in an experimental garden. Each seedling was germinated and grown in its own shading structure and exposed to two spatially separated light environments, in a factorial design of PAR and R:FR. Plant canopy foraging was evaluated at the end of one growing season in terms of canopy displacement, canopy area, leaf number, direction of stem lean, petiole aspect, and lamina aspect with respect to experimental light treatments. Leaf number and canopy area were greater on the high PAR sides of plants, irrespective of the R:FR treatment. Seedling canopies were displaced towards the direction of high PAR, but this relationship was not significant across all treatments. Petiole aspect was random and showed no significant directedness towards any of the light treatments. Lamina aspect and the direction of stem lean were distributed towards the direction of high PAR, irrespective of the R:FR treatment. Overall, first-year B. papyrifera seedlings used PAR, rather than R:FR ratio, as a photosensory cue for canopy light foraging. [ABSTRACT FROM AUTHOR]

Rising atmospheric CO2 may stimulate future forest productivity, possibly increasing carbon storage in terrestrial ecosystems, but how tropospheric ozone will modify this response is unknown. Because of the importance of fine roots to the belowground C cycle, we monitored fine-root biomass and associated C fluxes in regenerating stands of trembling aspen, and mixed stands of trembling aspen and paper birch at FACTS-II, the Aspen FACE project in Rhinelander, Wisconsin. Free-air CO2 enrichment (FACE) was used to elevate concentrations of CO2 (average enrichment concentration 535 µl l–1) and O3 (53 nl l–1) in developing forest stands in 1998 and 1999. Soil respiration, soil pCO2, and dissolved organic carbon in soil solution (DOC) were monitored biweekly. Soil respiration was measured with a portable infrared gas analyzer. Soil pCO2 and DOC samples were collected from soil gas wells and tension lysimeters, respectively, at depths of 15, 30, and 125 cm. Fine-root biomass averaged 263 g m–2 in control plots and increased 96% under elevated CO2. The increased root biomass was accompanied by a 39% increase in soil respiration and a 27% increase in soil pCO2. Both soil respiration and pCO2 exhibited a strong seasonal signal, which was positively correlated with soil temperature. DOC concentrations in soil solution averaged ~12 mg l–1 in surface horizons, declined with depth, and were little affected by the treatments. A simplified belowground C budget for the site indicated that native soil organic matter still dominated the system, and that soil respiration was by far the largest flux. Ozone decreased the above responses to elevated CO2, but effects were rarely statistically significant. We conclude that regenerating stands of northern hardwoods have the potential for substantially greater C input to soil due to greater fine-root production under elevated CO2. Greater fine-root biomass will be accompanied by greater soil C efflux as soil respiration, but leaching losses of C will probably be unaffected. [ABSTRACT FROM AUTHOR]

Allometry studies the change in scale between two dimensions of an organism. The metabolic theory of ecology predicts invariant allometric scaling exponents, while empirical studies evidenced inter- and intra-specific variations. This work aimed at identifying the sources of variations of the allometric exponents at both inter- and intra-specific levels using stem analysis from 9,363 trees for ten Eastern Canada species with a large shade-tolerance gradient. Specifically, the yearly allometric exponents, α [volume ( v) and diameter at breast height (DBH)], β [ v and height ( h)], and γ ( h and DBH) were modelled as a function of tree age for each species. α, and γ increased with tree age and then reached a plateau ranging from 2.45 to 3.12 for α, and 0.874-1.48 for γ. Pine species presented a local maximum. No effect of tree age on β was found for conifers, while it increased until a plateau ranging from 3.71 to 5.16 for broadleaves. The influence of shade tolerance on the growth trajectories was then explored. In the juvenile stage, α, and γ increased with shade tolerance while β was shade-tolerance independent. In the mature stage, β increased with shade tolerance, whereas γ decreased and α was shade-tolerance independent. The interaction between development stage and shade tolerance for allometric exponents demonstrates the importance of the changing functional requirements of trees for resource allocation at both the inter- and intra-specific level. These results indicate the need to also integrate specific functional traits, growth strategies and allocation, in allometric theoretical frameworks. [ABSTRACT FROM AUTHOR]

Evidence from tropical forests suggests understory habitats are associated with a high risk of disease, which may prevent the establishment of vulnerable tree species; in contrast, canopy gaps can act as refuges from these pathogens. However, few studies have investigated the impacts of pathogens on regeneration in temperate forests. To determine whether losses to fungi of seeds of Betula papyrifera, a light-loving species, varied between habitats that differed in their degree of openness, we applied fungicide to seeds buried in old fields, treefall gaps, and forest understory sites. We found that the application of fungicide significantly reduced losses in all habitats, relative to control values. This effect was habitat-dependent: the benefit of fungicide was greater in forest understory than in openings. This suggests that B. papyrifera is prevented from establishing in understory environments in part by its susceptibility to pathogen attack, and not solely because of a high light requirement. [ABSTRACT FROM AUTHOR]

Herbivory can influence ecosystem productivity, but recent evidence suggests that damage by herbivores modulates potential productivity specific to damage type. Because productivity is linked to photosynthesis at the leaf level, which in turn is influenced by atmospheric CO concentrations, we investigated how different herbivore damage types alter component processes of photosynthesis under ambient and elevated atmospheric CO. We examined spatial patterns in chlorophyll fluorescence and the temperature of leaves damaged by leaf-chewing, gall-forming, and leaf-folding insects in aspen trees as well as by leaf-chewing insects in birch trees under ambient and elevated CO at the aspen free-air CO enrichment (FACE) site in Wisconsin. Both defoliation and gall damage suppressed the operating efficiency of photosystem II (ΦPSII) in remaining leaf tissue, and the distance that damage propagated into visibly undamaged tissue was marginally attenuated under elevated CO. Elevated CO increased leaf temperatures, which reduced the cooling effect of gall formation and freshly chewed leaf tissue. These results provide mechanistic insight into how different damage types influence the remaining, visibly undamaged leaf tissue, and suggest that elevated CO may reduce the effects of herbivory on the primary photochemistry controlling photosynthesis. [ABSTRACT FROM AUTHOR]

An introduction is presented in which the editor discusses various articles in the issue on topics including climate cycles in desert perennial shrubs, California Coast Redwood tree-ring isotope chronology, and droughts in Amazonian trees.

This paper combines published and original data on sexual size dimorphism, reproductive behavior, and habitat types in turtles. Our major finding is that observed patterns of sexual size dimorphism correlate with habitat type and male mating strategy. (1) In most terrestrial species, males engage in combat with each other. Males typically grow larger than females. (2) In semiaquatic and 'bottom-walking' aquatic species, male combat is less common, but males often forcibly inseminate females. As in terrestrial species, males are usually larger than females. (3) In truly aquatic species, male combat and forcible insemination are rare. Instead, males utilize elaborate precoital displays, and female choice is highly important. Males are usually smaller than females. We interpret these correlations between sexual behavior and size dimorphism in terms of sexual selection theory: males are larger than females when large male size evolves as an adaptation to increase success in male combat, or to enable forcible insemination of females. In contrast, males are usually smaller than females where small size in males evolves to increase mobility (and hence, ability to locate females), or because selection for increased fecundity may result in increased female size. In turtle species with male combat or forcible insemination, the degree of male size superiority increases with mean species body size. [ABSTRACT FROM AUTHOR]

Early in his career, Russ Monson produced a series of influential eco-physiological papers that helped lay the foundation for the study of C4 plant evolution. Among the most important was a 1984 paper with Maurice Ku and Gerry Edwards that outlined the pathway for the evolutionary bridge from C3 to C4 photosynthesis. This model proposed C4 photosynthesis arose out of a shuttle that imported photorespiratory metabolites into bundle sheath (BS) cells, where glycine decarboxylase cleaved off CO2, allowing it to accumulate and be efficiently refixed by BS Rubisco. By the mid-1990's, Monson's research focus had shifted away from C4 plants, save for one 2003 paper on C3 versus C4 stomatal control with Travis Huxman, and a series of critical reviews on C4 evolution. These reviews heavily influenced the modern synthesis of C4 evolutionary studies, which incorporates phylogenomic understanding with physiological, molecular, and structural characterizations of trait shifts in multiple evolutionary lineages. Subsequent research supported the Monson et al. model from 1984, by showing a glycine shuttle occurs in nearly all C3–C4 intermediate species identified. Monson also examined the physiological controls over the ecological distribution of C3, C3–C4 intermediate, and C4 photosynthesis, building our understanding of the fitness value of the intermediate and C4 pathway in relevant microenvironments. By establishing the foundation for discoveries that followed, Russ Monson can rightly be considered a leading pioneer contributing to the evolutionary biology of C4 photosynthesis. [ABSTRACT FROM AUTHOR]

Geographic patterns of intraspecific variations in traits related to photosynthesis and biomass were examined in two separate common garden experiments using seed collected from 26 Sitka alder (Alnus sinuata Rydb.) and 18 paper birch (Betula papyrifera Marsh.) populations from climatically diverse locations in British Columbia, Canada. Exchange rates of carbon dioxide and water vapour were measured on 2-year-old seedlings to determine the maximum net instantaneous photosynthetic rate, mesophyll conductance, stomatal conductance, and photosynthetic water use efficiency. Height, stem diameter, root and shoot dry mass and fall frost hardiness data were also obtained. Mean population maximum photosynthetic rate ranged from 10.35 to 14.57 µmol CO2 m–2 s–1 in Sitka alder and from 14.76 to 17.55 µmol CO2 m–2 s–1 in paper birch. Based on canonical correlation analyses, populations from locations with colder winters and shorter (but not necessarily cooler) summers had higher maximum photosynthetic rates implying the existence of an inverse relationship between leaf longevity and photosynthetic capacity. Significant canonical variates based on climatic variables derived for the seed collection sites explained 58% and 41% of variation in the rate of photosynthesis in Sitka alder and paper birch, respectively. Since growing season length is reflected in date of frost hardiness development, an intrinsic relationship was found between photosynthetic capacity and the level of fall frost hardiness. The correlation was particularly strong for paper birch (r=–0.77) and less strong for Sitka alder (r=–0.60). Mean population biomass accumulation decreased with increased climate coldness. These patterns may be consequential for evaluation of the impact of climate change and extension of the growing season on plant communities. [ABSTRACT FROM AUTHOR]

The exploration of factors and processes affecting biodiversity loss is central to nature management and wildlife conservation, but only recently has knowledge about the absence of species been recognized as a valuable asset to understand the current biodiversity crisis. In this paper, we explore the dark diversity (species that belong to a site-specific species pool but that are not locally present) of breeding birds in Denmark assessed through species co-occurrence patterns. We apply a nation-wide atlas survey of breeding birds (with a 5 × 5 km resolution), to investigate how landscape characteristics may influence avian diversity, and whether threatened and near threatened species are more likely to occur in dark diversity than least concern (LC) species. On average, the dark diversity constituted 41% of all species belonging to the site-specific species pools and threatened and near-threatened species had a higher probability of belonging to the dark diversity than least concern species. Habitat heterogeneity was negatively related to dark diversity and the proportional cover of intensive agriculture positively related, implying that homogeneous landscapes dominated by agricultural interests led to more absent avian species. Finally, we found significant effects of human disturbance and distance to the coast, indicating that more breeding bird species were missing when human disturbance was high and in near-coastal areas. Our study provides the first attempt to investigate dark diversity among birds and highlights how important landscape characteristics may shape breeding bird diversity and reveal areas of considerable species impoverishment. [ABSTRACT FROM AUTHOR]

Lichens are well-suited organisms for experimental herbivory studies because their secondary compounds, assumed to deter grazing, can be non-destructively extracted. Thalli of 17 lichen species from various habitats were cut in two equal parts; compounds were extracted from one part by acetone, the other served as a control. These two pieces were offered as a paired choice to the generalist herbivore snailCepaea hortensis. Control thalli of all lichens were consumed at a low rate regardless of their investments in acetone-extractable lichen compounds; naturally compound-deficient lichen species were not preferred compared to those with high contents. However, for extracted thalli, there was a highly significant positive correlation between rate of consumption and the extracted compound contents. These data imply that herbivore defence has evolved in different directions in different lichens. Studied members of Parmeliaceae, common in oligotrophic habitats, have high contents of carbon-rich acetone-soluble compounds; these lichens became highly palatable to snails subsequent to acetone rinsing. Extracted lichen compounds were applied to pieces of filter paper and fed to snails. Extracts from members of the Parmeliaceae significantly deterred feeding on paper. Such data suggest that generalist herbivores may have shaped evolution in the widespread and highly diverse Parmeliaceae towards high investments in lichen compounds. On the other hand, lichens belonging to the Physciaceae and Teloschistales, common in nutrient-enriched habitats, are deficient in, or have low concentrations of, lichen compounds. Such lichens did not become more palatable after acetone rinsing. The orange anthraquinone compound parietin, restricted to the Teloschistales, and which has previously been found to protect against excess light, did not deter grazing. [ABSTRACT FROM AUTHOR]