Effects of plant leachates from four boreal understorey species on soil N mineralization, and white spruce (Picea glauca) germination and seedling growth

BACKGROUND AND AIMS: Natural regeneration of white spruce (Picea glauca) after disturbance has been reported to be very poor. Here a study was made to determine whether C compounds released from understorey species growing together with white spruce could be involved in this regeneration failure, either by (1) changing soil nutrient dynamics, (2) inhibiting germination, and/or (3) delaying seedling growth. METHODS: Foliage leachates were obtained from two shrubs (Ledum palustre and Empetrum hermaphroditum) and one bryophyte (Sphagnum sp.) with high phenolic compound concentrations that have been reported to depress growth of conifers in boreal forests, and, as a comparison, one bryophyte (Hylocomium splendens) with negligible phenolic compounds. Mineral soil from a white spruce forest was amended with plant leachates to examine the effect of each species on net N mineralization. Additionally, white spruce seeds and seedlings were watered with plant leachates to determine their effects on germination and growth. KEY RESULTS: Leachates from the shrubs L. palustre and E. hermaphroditum contained high phenolic compound concentrations and dissolved organic carbon (DOC), while no detectable levels of C compounds were released from the bryophytes Sphagnum sp. or H. splendens. A decrease in net N mineralization was determined in soils amended with L. palustre or E. hermaphroditum leachates, and this effect was inversely proportional to the phenolic concentrations, DOC and leachate C/N ratio. The total percentage of white spruce germination and the growth of white spruce seedlings were similar among treatments. CONCLUSIONS: These results suggest that the shrubs L. palustre and E. hermaphroditum could negatively affect the performance of white spruce due to a decrease in soil N availability, but not by direct effects on plant physiology.     

Bryophyte physiological responses to, and recovery from, long-term nitrogen deposition and phosphorus fertilisation in acidic grassland

Atmospheric nitrogen deposition can cause major declines in bryophyte abundance yet the physiological basis for such declines is not fully understood. Bryophyte physiological responses may also be sensitive bioindicators of both the impacts of, and recovery from, N deposition. Here, responses of tissue nutrients (nitrogen (N), phosphorus (P) and potassium (K): NPK), N and P metabolism enzymes (nitrate reductase and phosphomonoesterase), photosynthetic pigments, chlorophyll fluorescence, sclerophylly and percentage cover of two common bryophytes (Pseudoscleropodium purum and Rhytidiadelphus squarrosus) to long-term (11 yr) enhanced N deposition (+3.5 and +14 g N m(-2) yr(-1)) are reported in factorial combination with P addition. Recovery of responses 22 months after treatment cessation were also assessed. Enhanced N deposition caused up to 90% loss of bryophyte cover but no recovery was observed. Phosphomonoesterase activity and tissue N:P ratios increased up to threefold in response to N loading and showed clear recovery, particularly in P. purum. Smaller responses and recovery were also seen in all chlorophyll fluorescence measurements and altered photosynthetic pigment composition. The P limitation of growth appears to be a key mechanism driving bryophyte loss along with damage to photosystem II. Physiological measurements are more sensitive than measurements of abundance as bioindicators of N deposition impact and of recovery in particular.     

Elevational patterns of carbon,nitrogen and phosphorus in understory bryophytes on the eastern slope of Gongga Mountain,China

Aims The nutrient uptake, requirement and releasing rates of bryophytes are very different from those of tracheophytes. However, it is difficult to make a quantitative evaluation of bryophytes’ roles in nutrient cycling and their specific eco-physiological adaptations due to lack of knowledge of their concentrations and stoichiometric ratios of carbon (C), nitrogen (N) and phosphorus (P). To fill this gap, the present study aims to investigate: (i) what are the elevational trends of C, N and P concentrations and stoichiometric ratios of bryophytes? (ii) whether C, N and P concentrations and stoichiometric ratios of bryophytes differ between different bryophyte types (in terms of the growth form and living substrate)? and (iii) how do the exponent scalings of N and P of bryophytes change along the elevational gradient?     

Differential effects of ammonium and nitrate deposition on fen phanerogams and bryophytes

Question: High atmospheric nitrogen (N) deposition has been shown to affect productivity and species composition of terrestrial ecosystems. This study focused on the differential effects of the two inorganic N forms in atmospheric deposition (i.e. ammonium and nitrate). Methods and location: Nutrient addition experiments were carried out during 4 years in a mesotrophic fen in a low‐deposition area in Ireland. In a factorial design, plots were fertilized with ammonium and/or nitrate, in two doses comparable with 35 and 70 kg N ha^?1 y^?1 and compared with an unfertilized control. Results: Vascular plant biomass as well as bryophyte biomass were not affected by N dose but showed significantly different responses to the N form. In the ammonium‐fertilized plots, vascular plant biomass was higher and moss biomass was lower than the control, while nitrate additions had no effect. Vascular plant species density was high (16 species per 0.49 m²) and was not affected by any of the treatments; bryophyte species density was also high (seven species per 0.04 m²) but showed a significant decrease upon ammonium fertilization. Conclusion: The vulnerability of the mesotrophic vegetation to enhanced atmospheric N deposition depends strongly on the N form. If N would be mainly deposited as NO_x, no detrimental effects on the vegetation will occur. If, however, the deposition is mainly in the form of NH_y, the bryophyte vegetation will be seriously damaged, while the vascular plant vegetation will show an increased biomass production with possible shifts in dominance from Carex and herb species to grasses and shrubs.     

六种人工针叶幼林下地表苔藓植物生物量与碳贮量

调查了岷江上游6种人工针叶幼林(油松林、华山松林、日本落叶松林、云杉林、油松-华山松混交林和云杉-华山松混交林)下地表苔藓植物生物量,测定了C含量并估计了林分地表苔藓植物C贮量,比较分析了它们的差异性.结果表明,6种人工针叶幼林下地表苔藓植物总生物量在3.11～460.36 kg·hm^-2之间,而平均C含量在37.44±0.21%～3.9±0.70%之间,总C贮量在1.12±0.03～168.9±0.92 kg·hm^-2之间,但在样方水平上只有云杉林地表苔藓植物生物量与其它林型间差异明显,落叶松林下C含量与其它差异明显(P＜0.0).6种人工林类型中,云杉林地表苔藓植物总生物量和C贮量最高,华山松林下最低.综合分析表明,样方调查数量与布局对生物量取样精度有重要影响,岷江上游人工林下地表苔藓植物生物量与C贮量较低,林分类型与林分特征有重要影响,而疏伐、修枝等措施是改善人工密林下地表苔藓植物发育,增加生物量与C贮量的有效管理措施.     

Effects of nitrogen additions on above- and belowground carbon dynamics in two tropical forests

Anthropogenic nitrogen (N) deposition is increasing rapidly in tropical regions, adding N to ecosystems that often have high background N availability. Tropical forests play an important role in the global carbon (C) cycle, yet the effects of N deposition on C cycling in these ecosystems are poorly understood. We used a field N-fertilization experiment in lower and upper elevation tropical rain forests in Puerto Rico to explore the responses of above- and belowground C pools to N addition. As expected, tree stem growth and litterfall productivity did not respond to N fertilization in either of these N-rich forests, indicating a lack of N limitation to net primary productivity (NPP). In contrast, soil C concentrations increased significantly with N fertilization in both forests, leading to larger C stocks in fertilized plots. However, different soil C pools responded to N fertilization differently. Labile (low density) soil C fractions and live fine roots declined with fertilization, while mineral-associated soil C increased in both forests. Decreased soil CO₂ fluxes in fertilized plots were correlated with smaller labile soil C pools in the lower elevation forest (R² = 0.65, p < 0.05), and with lower live fine root biomass in the upper elevation forest (R² = 0.90, p < 0.05). Our results indicate that soil C storage is sensitive to N deposition in tropical forests, even where plant productivity is not N-limited. The mineral-associated soil C pool has the potential to respond relatively quickly to N additions, and can drive increases in bulk soil C stocks in tropical forests.     

Response of epiphytic bryophytes to simulated N deposition in a subtropical montane cloud forest in southwestern China

A field manipulation experiment was conducted in a subtropical montane cloud forest in southwestern China to determine the possible responses of epiphytic bryophytes to increasing nitrogen (N) deposition from community to physiology level, and to find sensitive epiphytic bryophytes that may be used as indicators for assessing the degree of N pollution. N addition had significantly negative effects on species richness and cover of the epiphytic bryophyte community. Harmful effects of high N loads were recorded for chlorophyll, growth, and vitality of the species tested. The decline of some epiphytic bryophytes may result from detrimental effects on degradation to photosynthetic pigments. Bazzania himalayana (Mitt.) Schiffn., Bazzania ovistipula (Steph.) Mizut., and Homaliodendron flabellatum (Sm.) Fleisch. are candidates in atmospheric nitrogen monitoring. Epiphytic bryophytes in the montane cloud forest are very sensitive to increasing N deposition and often difficult to recover once they have been destroyed, providing early detection of enhanced N pollution for trees or even the whole forest ecosystem. The inference that increasing N pollution may lead to loss of biodiversity is a concern to the developing economy in western China, and should alert the government to the adverse impacts caused by increased industrial pollution during the process of China’s West Development.     

长白山暗针叶林苔藓植物生物量的研究

在长白山北坡暗针叶林对地面和树附生苔藓植物的生物量进行了测定．地面生苔藓采取样带调查取样法测定,树附生苔藓应用McCune方法对树干和树枝的附生苔藓生物量都做了细致的测定．结果表明,长白山暗针叶林中的苔藓植物分布很不均匀,随海拔变化差异很大,海拔1100m最低,仅为543kg·hm^-2;海拔1250m最高,达5097kg·hm^-2．苔藓植物生物量的变化对生境有很好的指示作用,特别是塔藓和拟垂枝藓的生物量随海拔的变化与森林系统的群落学特点有一定的相关性：在海拔1100～1700m,塔藓的生物量与臭冷杉的重要值变化趋势相近,随海拔升高而减少;拟垂枝藓的生物量与鱼鳞云杉重要值的变化趋势相似,随海拔升高而增加．此外,生物量随海拔的变化表明了不同苔藓植物对环境条件要求的差异,拟垂枝藓比塔藓水分条件要求更高．因此,生物量的研究在植物生理上也有一定的指示作用．     

Multi‐nutrient vs. nitrogen‐only effects on carbon sequestration in grassland soils

Human activities have greatly increased the availability of biologically active forms of nutrients [e.g., nitrogen (N), phosphorous (P), potassium (K), magnesium (Mg)] in many soil ecosystems worldwide. Multi‐nutrient fertilization strongly increases plant productivity but may also alter the storage of carbon (C) in soil, which represents the largest terrestrial pool of organic C. Despite this issue is important from a global change perspective, key questions remain on how the single addition of N or the combination of N with other nutrients might affect C sequestration in human‐managed soils. Here, we use a 19‐year old nutrient addition experiment on a permanent grassland to test for nutrient‐induced effects on soil C sequestration. We show that combined NPKMg additions to permanent grassland have ‘constrained’ soil C sequestration to levels similar to unfertilized plots whereas the single addition of N significantly enhanced soil C stocks (N‐only fertilized soils store, on average, 11 t C ha^?1 more than unfertilized soils). These results were consistent across grazing and liming treatments suggesting that whilst multi‐nutrient additions increase plant productivity, soil C sequestration is increased by N‐only additions. The positive N‐only effect on soil C content was not related to changes in plant species diversity or to the functional composition of the plant community. N‐only fertilized grasslands show, however, increases in total root mass and the accumulation of organic matter detritus in topsoils. Finally, soils receiving any N addition (N only or N in combination with other nutrients) were associated with high N losses. Overall, our results demonstrate that nutrient fertilization remains an important global change driver of ecosystem functioning, which can strongly affect the long‐term sustainability of grassland soil ecosystems (e.g., soils ability to deliver multiple ecosystem services).     

Impact of Land-Use Intensity and Productivity on Bryophyte Diversity in Agricultural Grasslands

While bryophytes greatly contribute to plant diversity of semi-natural grasslands, little is known about the relationships between land-use intensity, productivity, and bryophyte diversity in these habitats. We recorded vascular plant and bryophyte vegetation in 85 agricultural used grasslands in two regions in northern and central Germany and gathered information on land-use intensity. To assess grassland productivity, we harvested aboveground vascular plant biomass and analyzed nutrient concentrations of N, P, K, Ca and Mg. Further we calculated mean Ellenberg indicator values of vascular plant vegetation. We tested for effects of land-use intensity and productivity on total bryophyte species richness and on the species richness of acrocarpous (small & erect) and pleurocarpous (creeping, including liverworts) growth forms separately. Bryophyte species were found in almost all studied grasslands, but species richness differed considerably between study regions in northern Germany (2.8 species per 16 m²) and central Germany (6.4 species per 16 m²) due environmental differences as well as land-use history. Increased fertilizer application, coinciding with high mowing frequency, reduced bryophyte species richness significantly. Accordingly, productivity estimates such as plant biomass and nitrogen concentration were strongly negatively related to bryophyte species richness, although productivity decreased only pleurocarpous species. Ellenberg indicator values for nutrients proved to be useful indicators of species richness and productivity. In conclusion, bryophyte composition was strongly dependent on productivity, with smaller bryophytes that were likely negatively affected by greater competition for light. Intensive land-use, however, can also indirectly decrease bryophyte species richness by promoting grassland productivity. Thus, increasing productivity is likely to cause a loss of bryophyte species and a decrease in species diversity.     

高山森林林窗和生长基质对苔藓植物氮和磷含量的影响

苔藓在森林生态系统养分富集和循环方面有重要作用,苔藓植物在氮磷循环中的作用可能受到森林更新和生长基质的影响.为理解苔藓在森林生态系统养分循环中的作用,研究了高山森林不同位置(林窗中心、林窗边缘、林下)和不同生长基质(活立木、倒木、枯立木、大枯枝、根桩、地表)上的苔藓植物氮磷含量.结果表明:地表苔藓氮含量(3.12 mg·g^-1)显著低于其他生长基质上的苔藓,尽管枯立木附生苔藓氮含量高达17.41 mg·g^-1,但倒木、大枯枝、枯立木和活立木苔藓氮含量差异不显著;最高(1.09 mg·g^-1)和最低(0.61 mg·g^-1)的磷含量分别出现在森林地表和枯立木的苔藓,且森林地表的苔藓磷含量显著高于其他生长基质上的苔藓,但倒木、大枯枝、根桩和活立木上的苔藓磷含量差异不显著.林窗位置显著影响了倒木和大枯枝上的苔藓氮、磷含量,林窗内倒木和大枯枝上的苔藓氮、磷含量显著高于林窗边缘.粗木质残体的类型、腐烂等级对苔藓氮、磷含量的影响存在差异,两者交互作用的影响显著;第Ⅴ腐烂等级倒木上的苔藓氮含量显著高于其他腐烂等级;第Ⅲ腐烂等级大枯枝上的苔藓氮含量显著高于其他腐烂等级;第Ⅱ腐烂等级倒木上的苔藓磷含量显著高于其他腐烂等级;第Ⅳ腐烂等级枯立木上的苔藓磷含量显著高于其他腐烂等级.可见,森林林窗更新和粗木质残体腐解过程会影响苔藓植物的氮、磷含量,同时影响森林生态系统的养分循环过程.     

Bryophytes attenuate anthropogenic nitrogen inputs in boreal forests

Productivity in boreal ecosystems is primarily limited by available soil nitrogen (N), and there is substantial interest in understanding whether deposition of anthropogenically derived reactive nitrogen (N_r) results in greater N availability to woody vegetation, which could result in greater carbon (C) sequestration. One factor that may limit the acquisition of N_r by woody plants is the presence of bryophytes, which are a significant C and N pool, and a location where associative cyanobacterial N‐fixation occurs. Using a replicated stand‐scale N‐addition experiment (five levels: 0, 3, 6, 12, and 50 kg N ha^?1 yr^?1; n=6) in the boreal zone of northern Sweden, we tested the hypothesis that sequestration of N_r into bryophyte tissues, and downregulation of N‐fixation would attenuate N_r inputs, and thereby limit anthropogenic N_r acquisition by woody plants. Our data showed that N‐fixation per unit moss mass and per unit area sharply decreased with increasing N addition. Additionally, the tissue N concentrations of Pleuorzium schreberi increased and its biomass decreased with increasing N addition. This response to increasing N addition caused the P. schreberi N pool to be stable at all but the highest N addition rate, where it significantly decreased. The combined effects of changed N‐fixation and P. schreberi biomass N accounted for 56.7% of cumulative N_r additions at the lowest N_r addition rate, but only a minor fraction for all other treatments. This ‘bryophyte effect’ can in part explain why soil inorganic N availability and acquisition by woody plants (indicated by their δ¹⁵N signatures) remained unchanged up to N addition rates of 12 kg ha^?1 yr^?1 or greater. Finally, we demonstrate that approximately 71.8% of the boreal forest experiences N_r deposition rates at or below 3 kg ha^?1 yr^?1, suggesting that bryophytes likely limit woody plant acquisition of ambient anthropogenic N_r inputs throughout a majority of the boreal forest.     

Bryophyte‐cyanobacteria associations as regulators of the northern latitude carbon balance in response to global change

Ecosystems in the far north, including arctic and boreal biomes, are a globally significant pool of carbon (C). Global change is proposed to influence both C uptake and release in these ecosystems, thereby potentially affecting whether they act as C sources or sinks. Bryophytes (i.e., mosses) serve a variety of key functions in these systems, including their association with nitrogen (N₂)‐fixing cyanobacteria, as thermal insulators of the soil, and producers of recalcitrant litter, which have implications for both net primary productivity (NPP) and heterotrophic respiration. While ground‐cover bryophytes typically make up a small proportion of the total biomass in northern systems, their combined physical structure and N₂‐fixing capabilities facilitate a disproportionally large impact on key processes that control ecosystem C and N cycles. As such, the response of bryophyte‐cyanobacteria associations to global change may influence whether and how ecosystem C balances are influenced by global change. Here, we review what is known about their occurrence and N₂‐fixing activity, and how bryophyte systems will respond to several key global change factors. We explore the implications these responses may have in determining how global change influences C balances in high northern latitudes.     

Growth,Net Production,Litter Decomposition,and Net Nitrogen Accumulation by Epiphytic Bryophytes in a Tropical Montane Forest1

To understand the ecological roles of epiphytic bryophytes in the carbon (C) and nitrogen (N) cycles of a tropical montane forest, we used samples in enclosures to estimate rates of growth, net production, and N accumulation by shoots in the canopy, and litterbags, to estimate rates of decomposition and N dynamics of epiphytic bryophyte litter in the canopy and on the forest floor in Monteverde, Costa Rica. Growth of epiphytic bryophytes was estimated at 30.0–49.9 percent/yr, net production at 122–203 g/m²/yr, and N accumulation at 1.8–3.0 g N/m²/yr. Cumulative mass loss from litterbags after one and two years in the canopy was 17 ± 2 and 19 ± 2 percent (mean ± 1 SE) of initial sample mass, respectively, and mass loss from litter and green shoots in litterbags after one year on the forest floor was 29 ± 2 and 45 ± 3 percent, respectively. Approximately 30 percent of the initial N mass was released rapidly from litter in both locations. Nitrogen loss from green shoots on the forest floor was greater; about 47 percent of the initial N mass was lost within the first three months. There was no evidence for net N immobilization by litter or green shoots, but the remaining N in litter was apparently recalcitrant. Annual net accumulation of C and N by epiphytic bryophytes was estimated at 37–64 g C/m²/yr and 0.8–1.3 g N/m²/yr, respectively. Previous research at this site indicated that epiphytic bryophytes retain inorganic N from atmospheric deposition to the canopy. Therefore, they play a major role in transforming N from mobile to highly recalcitrant forms in this ecosystem.     

Controls on production of bryophytes in an arctic tundra stream

1. Two bryophyte taxa (Hygrohypnum spp. and, to a lesser extent, Fontinalis neomexicana) were abundant in riffles within phosphorus-fertilized reaches of the Kuparuk River (North Slope, Alaska), but were much less common in fertilized pools and virtually absent in unfertilized reaches of the river. We conducted field experiments using stems and clumps of both species and artificial bryophytes to test the hypotheses that bryophyte growth was strongly limited by low phosphorus concentrations in unfertilized reaches, and limited by epiphytes in fertilized pools. 2. Stem tips of Hygrohypnum spp. did not elongate when grown in unfertilized pool and riffle environments. In fertilized reaches, Hygrohypnum elongated significantly, although there was no significant difference in elongation of stem tips placed in pools [2.5 ± 0.9 cm (SD)] as opposed to riffles (2.8 ± 0.9 cm) for 32 days. 3. Stem tips of F. neomexicana elongated significantly in all sites. There was a significant difference in elongation of stem tips in control and fertilized riffles (2.1 ± 1.1 and 4.7 ± 0.1 cm, respectively) but not in tips grown in control and fertilized pools (2.8 ± 0.8 and 2.7 ± 0.9 cm, respectively). 4. Biomass increments in clumps of these same species followed similar patterns except in fertilized pools. Hygrohypnum spp. lost weight in control riffle environments and did not grow in pools, but accumulated 181 ± 44 and 335 ± 200% of initial biomass in fertilized riffles in 1992 (over 32 days) and 1993 (over 44 days), respectively. F. neomexicana accumulated 38 ± 39 and 98 ± 47% of initial biomass in 1992 in unfertilized and fertilized riffles, respectively. Total phosphorus concentrations of both bryophytes in 1992 were significantly greater when grown in fertilized riffles than control riffles. 5. Artificial mosses (untwisted, natural fibre rope) and clumps of Hygrohypnum spp. were used to assess effects of flow regime on derrital and epiphyte accumulation in the fertilized zone. Epiphyte and detrital mass was 4–4.5 times greater on average on artificial mosses in slow-flowing pool environments than in fast-flowing riffle environments. Epiphyte chlorophyll a was 4 times greater on Hygrohypnum clumps in pools than in riffles. This difference was probably brought about by increased detrital deposition and reduced grazing by invertebrates in pools. It is likely that both Hygrohypnum spp. and F. neomexicana could grow throughout the river, but are limited strongly by low phosphorus concentrations in unfertilized reaches and secondarily by detritus accumulation and interference competition with epiphytic algae in fertilized pools.     

The physiological basis of bryophyte production

In the main features of their carbon metabolism and physiological responses, bryophytes behave as normal C_:) plants. However, their small size and frequent poikilohydric habit have important effects on the context in which these characteristics are expressed, and on their environmental physiology. Many are tolerant of drying out to low water contents (c. 5–10%, of dry weight). Photosynthesis declines rapidly with water loss, and resumes with greater or lesser delay on remoistening. The rate and completeness of recovery depend on the intensity and duration of desiccation, and on drought-hardening (perhaps largely related to protection of cell components from oxidative damage) which lakes place as the bryophyte dries. Most bryophytes, including species of well-illuminated habitats, function in effect as shade plants, with low chlorophyll a/b ratios, and become light-saturated at relatively low irradiance. Boundary-layer resistance is critically important in determining water loss from bryophytes in many situations. The time for which a poikilohydric species can photosynthesize after rain is determined by storage capacity and rate of water loss, both strongly influenced by growth-form. In sheltered habitats with extensive bryophyte cover water loss is largely determined by radiation balance, and may be very slow in deeply shaded places. Bryophyte growth-forms must represent an adaptive balance between water economy and needs for light capture and carbon and mineral nutrient acquisition.     

Long-term effects of PO4 fertilization on the distribution of bryophytes in an arctic river

1. Each year since 1983, H₃PO₄ has been added continuously during the ice-free season to a P-limited tundra river (Kuparuk River, North Slope, Alaska). Effects on epilithic metabolism, invertebrate community structure and fish production developed quickly. 2. In 1990, 7 years after fertilization began, we noted extensive coverage by bryophytes within the fertilized reach of the river, where very little had been noted before. Bryophyte biomass from a limited set of quadrats taken in 1990 and 1991 yielded 17 ± 9 (SE) g dry mass m^?2 in control reaches and 322 ± 96 g dry mass m^?2 in fertilized reaches. 3. An initial survey of macroalgal and bryophyte cover in 1991 suggested that the moss Schistidium (Grimmia) agassizii was distributed in both control and fertilized reaches of the river. No clear difference in coverage by this species was found in either reach. 4. In contrast, two species of Hygrohypnum (H. alpestre and H. ochraceum) were found almost exclusively in the fertilized reach. An extensive point transect survey done in 1992, above, within and below the fertilized reach, indicated that increased cover and biomass of Hygrohypnum spp. were confined to the fertilized reach of the river. Detrended correspondence analysis clearly separated the macrophyte and macroalgal communities in the fertilized reach from those in the control and downstream reaches. 5. A fourth bryophyte species (Fontinalis neomexicana) also occurred almost exclusively in the fertilized reach, but was much less abundant than the Hygrohypnum species. 6. Analysis of total N and P in the tissues of the Hygrohypnum spp., and estimates of average coverage (～15%) and biomass (～150g dry weight m^?2) over an 8km fertilized reach, suggest that these species alone may have removed two-thirds of the P added in the fertilizer experiment. The bryophyte community in this river is likely to be the dominant sink for P in the fertilized reach.     

Production and microtopography of bog bryophytes: response to warming and water-table manipulations

Boreal peatlands, which contain a large fraction of the world's soil organic carbon pool, may be significantly affected by changes in climate and land use, with attendant feedback to climate through changes in albedo, fluxes of energy or trace gases, and soil carbon storage. The response of peatlands to changing environmental conditions will probably be dictated in part by scale-dependent topographic heterogeneity, which is known to interact with hydrology, vegetation, nutrients, and emissions of trace gases. Because the bryophyte community can contribute the majority of aboveground production in bogs, we investigated how microscale topography affects the response of bryophyte species production and cover to warming (using overhead infrared lamps) and manipulations of water-table height within experimental mesocosms. We removed 27 intact peat monoliths (2.1-m² surface area, 0.5-0.7 m depth) from a bog in northern Minnesota, USA, and subjected them to three warming and three water-table treatments in a fully crossed factorial design. Between 1994 and 1998, we determined annual production of the four dominant bryophyte taxa within three microtopographic zones (low, medium, and high relative to the water table). We also estimated species cover and calculated changes in topography and roughness of the bryophyte surface through time. Total production of all bryophytes, and production of the individual taxa Polytrichum strictum, Sphagnum magellanicum, and Sphagnum Section Acutifolia, were about 100% greater in low microtopographic zones than in high zones, and about 50% greater in low than in medium zones. Production of bryophytes increased along the gradient of increasing water-table heights, but in most years, total production of bryophytes was negatively correlated with height above the set water table only for the wettest water-table treatment. Although bryophyte production was unaffected by the warming treatments, the bryophyte surface flattened in proportion to the degree of warming. These results indicate that production of bryophytes is driven most strongly by the absolute and relative height of the bryophyte surface above the water table. Predicted changes in water-table height commensurate with changes in surface temperature may thus affect both production and superficial topography of bryophyte communities.     

Microsatellite markers for Hylocomium splendens (Hylocomiaceae)

? Premise of the study: The perennial feather moss Hylocomium splendens is one of the most widely distributed and common bryophytes in the Northern Hemisphere and has, because of its capacity to grow under a wide range of environmental conditions, been used as a biomonitor for atmospheric metal deposition in Europe. ? Methods and Results: We present a multiplex approach for the analysis of 14 microsatellite markers tested on 194 H. splendens gametophytes. Ten of the markers are developed recently, and are presented for the first time in this paper, whereas four were previously developed but have not been used for population genetic investigations. ? Conclusions: The microsatellite markers reported here will provide a powerful tool for further research on population genetic structure in H. splendens.     

Impacts of long-term enhanced UV-B radiation on bryophytes in two sub-Arctic heathland sites of contrasting water availability

Background and Aims

Anthropogenic depletion of stratospheric ozone in Arctic latitudes has resulted in an increase of ultraviolet-B radiation (UV-B) reaching the biosphere. UV-B exposure is known to reduce above-ground biomass and plant height, to increase DNA damage and cause accumulation of UV-absorbing compounds in polar plants. However, many studies on Arctic mosses tended to be inconclusive. The importance of different water availability in influencing UV-B impacts on lower plants in the Arctic has been poorly explored and might partially explain the observed wide variation of responses, given the importance of water in controlling bryophyte physiology. This study aimed to assess the long-term responses of three common sub-Arctic bryophytes to enhanced UV-B radiation (+UV-B) and to elucidate the influence of water supply on those responses.

Methods

Responses of three sub-Arctic bryophytes (the mosses Hylocomium splendens and Polytrichum commune and the liverwort Barbilophozia lycopodioides) to +UV-B for 15 and 13 years were studied in two field experiments using lamps for UV-B enhancement with identical design and located in neighbouring areas with contrasting water availability (naturally mesic and drier sites). Responses evaluated included bryophyte abundance, growth, sporophyte production and sclerophylly; cellular protection by accumulation of UV-absorbing compounds, β-carotene, xanthophylls and development of non-photochemical quenching (NPQ); and impacts on photosynthesis performance by maximum quantum yield (F_v /F_m) and electron transport rate (ETR) through photosystem II (PSII) and chlorophyll concentrations.

Results

Responses were species specific: H. splendens responded most to +UV-B, with reduction in both annual growth (–22 %) and sporophyte production (–44 %), together with increased β-carotene, violaxanthin, total chlorophyll and NPQ, and decreased zeaxanthin and de-epoxidation of the xanthophyll cycle pool (DES). Barbilophozia lycopodioides responded less to +UV-B, showing increased β-carotene and sclerophylly and decreased UV-absorbing compounds. Polytrichum commune only showed small morphogenetic changes. No effect of UV-B on bryophyte cover was observed. Water availability had profound effects on bryophyte ecophysiology, and plants showed, in general, lower growth and ETR, together with a higher photoprotection in the drier site. Water availability also influenced bryophyte responses to +UV-B and, in particular, responses were less detectable in the drier site.

Conclusions

Impacts of UV-B exposure on Arctic bryophytes were significant, in contrast to modest or absent UV-B effects measured in previous studies. The impacts were more easily detectable in species with high plasticity such as H. splendens and less obvious, or more subtle, under drier conditions. Species biology and water supply greatly influences the impact of UV-B on at least some Arctic bryophytes and could contribute to the wide variation of responses observed previously.