Juul Limpens

Atmospheric nitrogen deposition promotes carbon loss from peat bogs

Peat bogs have historically represented exceptional carbon (C) sinks because of their extremely low decomposition rates and consequent accumulation of plant remnants as peat. Among the factors favoring that peat accumulation, a major role is played by the chemical quality of plant litter itself, which is poor in nutrients and characterized by polyphenols with a strong inhibitory effect on microbial breakdown. Because bogs receive their nutrient supply solely from atmospheric deposition, the global increase of atmospheric nitrogen (N) inputs as a consequence of human activities could potentially alter the litter chemistry with important, but still unknown, effects on their C balance. Here we present data showing the decomposition rates of recently formed litter peat samples collected in nine European countries under a natural gradient of atmospheric N deposition from ≈0.2 to 2 g·m−2·yr−1. We found that enhanced decomposition rates for material accumulated under higher atmospheric N supplies resulted in higher carbon dioxide (CO2) emissions and dissolved organic carbon release. The increased N availability favored microbial decomposition (i) by removing N constraints on microbial metabolism and (ii) through a chemical amelioration of litter peat quality with a positive feedback on microbial enzymatic activity. Although some uncertainty remains about whether decay-resistant Sphagnum will continue to dominate litter peat, our data indicate that, even without such changes, increased N deposition poses a serious risk to our valuable peatland C sinks.

Ecosystem responses to reduced and oxidised nitrogen inputs in European terrestrial habitats

While it is well established that ecosystems display strong responses to elevated nitrogen deposition, the importance of the ratio between the dominant forms of deposited nitrogen (NH(x) and NO(y)) in determining ecosystem response is poorly understood. As large changes in the ratio of oxidised and reduced nitrogen inputs are occurring, this oversight requires attention. One reason for this knowledge gap is that plants experience a different NH(x):NO(y) ratio in soil to that seen in atmospheric deposits because atmospheric inputs are modified by soil transformations, mediated by soil pH. Consequently species of neutral and alkaline habitats are less likely to encounter high NH(4)(+) concentrations than species from acid soils. We suggest that the response of vascular plant species to changing ratios of NH(x):NO(y) deposits will be driven primarily by a combination of soil pH and nitrification rates. Testing this hypothesis requires a combination of experimental and survey work in a range of systems.

Climatic modifiers of the response to nitrogen deposition in peat-forming Sphagnum mosses: a meta-analysis

Peatlands in the northern hemisphere have accumulated more atmospheric carbon (C) during the Holocene than any other terrestrial ecosystem, making peatlands long-term C sinks of global importance. Projected increases in nitrogen (N) deposition and temperature make future accumulation rates uncertain. Here, we assessed the impact of N deposition on peatland C sequestration potential by investigating the effects of experimental N addition on Sphagnum moss. We employed meta-regressions to the results of 107 field experiments, accounting for sampling dependence in the data. We found that high N loading (comprising N application rate, experiment duration, background N deposition) depressed Sphagnum production relative to untreated controls. The interactive effects of presence of competitive vascular plants and high tissue N concentrations indicated intensified biotic interactions and altered nutrient stochiometry as mechanisms underlying the detrimental N effects. Importantly, a higher summer temperature (mean for July) and increased annual precipitation intensified the negative effects of N. The temperature effect was comparable to an experimental application of almost 4 g N m(-2)  yr(-1) for each 1°C increase. Our results indicate that current rates of N deposition in a warmer environment will strongly inhibit C sequestration by Sphagnum-dominated vegetation.

Expansion of Sphagnum fallax in bogs: striking the balance between N and P availability

Abstract Nitrogen deposition may cause shifts in the Sphagnum species composition of bogs, ultimately affecting the conservation value of these systems. We studied the effects of N and P on the expansion of S. fallax and S. flexuosum in bogs. We related historical census data of S. fallax, S. flexuosum, and four of their accompanying species to changes in N deposition. In addition, we conducted two fertilization experiments with N and P; one at a low deposition site with S. flexuosum and one at two high deposition sites with S. fallax. Finally, we related existing data on capitulum N and P concentrations of S. fallax to its abundance in the field. A relative increase in observed frequency of S. fallax coincided with an historical increase in N deposition in the Netherlands. There was no indication that S. fallax consistently outcompeted one of the other five Sphagnum species; the observed frequency of the Sphagnum species analysed was rather stable with time. The census data on S. flexuosum did not indicate a response to N deposition, but the species expanded at the low N deposition site when extra N was applied. In contrast, the expansion of S. fallax at the high deposition sites was limited by P. Organic nutrient concentrations suggested that when S. fallax can maintain a capitulum N concentration of 7 mg g-1 or higher and a P concentration of 0.7 mg g-1 or higher the species can grow to dominate. We conclude that S. fallax will gradually colonize an increasing number of new habitats in areas with a low, albeit increasing, N deposition, but will only grow to dominate when P supply is adequate. Then, the expansion of S. fallax may lead to ousting of the other Sphagnum species present.

Can frequent precipitation moderate the impact of drought on peatmoss carbon uptake in northern peatlands

Northern peatlands represent a large global carbon store that can potentially be destabilized by summer water table drawdown. Precipitation can moderate the negative impacts of water table drawdown by rewetting peatmoss (Sphagnum spp.), the ecosystems key species. Yet, the frequency of such rewetting required for it to be effective remains unknown. We experimentally assessed the importance of precipitation frequency for Sphagnum water supply and carbon uptake during a stepwise decrease in water tables in a growth chamber. CO2 exchange and the water balance were measured for intact cores of three peatmoss species (Sphagnum majus, Sphagnum balticum and Sphagnum fuscum) representative of three hydrologically distinct peatland microhabitats (hollow, lawn and hummock) and expected to differ in their water table-precipitation relationships. Precipitation contributed significantly to peatmoss water supply when the water table was deep, demonstrating the importance of precipitation during drought. The ability to exploit transient resources was species-specific; S. fuscum carbon uptake increased linearly with precipitation frequency for deep water tables, whereas carbon uptake by S. balticum and S. majus was depressed at intermediate precipitation frequencies. Our results highlight an important role for precipitation in carbon uptake by peatmosses. Yet, the potential to moderate the impact of drought is species-specific and dependent on the temporal distribution of precipitation.

Positive shrub-tree interactions facilitate woody encroachment in boreal peatlands

1. Boreal ecosystems are warming roughly twice as fast as the global average, resulting in woody expansion that could further speed up the climate warming. Boreal peatbogs are waterlogged systems that store more than 30% of the global soil carbon. Facilitative effects of shrubs and trees on the establishment of new individuals could increase tree cover with profound consequences for the structure and functioning of boreal peatbogs, carbon sequestration and climate. 2. We conducted two field experiments in boreal peatbogs to assess the mechanisms that explain tree seedling recruitment and to estimate the strength of positive feedbacks between shrubs and trees. We planted seeds and seedlings of Pinus sylvestris in microsites with contrasting water-tables and woody cover and manipulated both shrub canopy and root competition. We monitored seedling emergence, growth and survival for up to four growing seasons and assessed how seedling responses related to abiotic and biotic conditions. 3. We found that tree recruitment is more successful in drier topographical microsites with deeper water-tables. On these hummocks, shrubs have both positive and negative effects on tree seedling establishment. Shrub cover improved tree seedling condition, growth and survival during the warmest growing season. In turn, higher tree basal area correlates positively with soil nutrient availability, shrub biomass and abundance of tree juveniles. 4. Synthesis. Our results suggest that shrubs facilitate tree colonization of peatbogs which further increases shrub growth. These facilitative effects seem to be stronger under warmer conditions suggesting that a higher frequency of warmer and dry summers may lead to stronger positive interactions between shrubs and trees that could eventually facilitate a shift from moss to tree-dominated systems.

Glasshouse vs field experiments: do they yield ecologically similar results for assessing N impacts on peat mosses?

• Peat bogs have accumulated more atmospheric carbon (C) than any other terrestrial ecosystem today. Most of this C is associated with peat moss (Sphagnum) litter. Atmospheric nitrogen (N) deposition can decrease Sphagnum production, compromising the C sequestration capacity of peat bogs. The mechanisms underlying the reduced production are uncertain, necessitating multifactorial experiments. • We investigated whether glasshouse experiments are reliable proxies for field experiments for assessing interactions between N deposition and environment as controls on Sphagnum N concentration and production. We performed a meta-analysis over 115 glasshouse experiments and 107 field experiments. • We found that glasshouse and field experiments gave similar qualitative and quantitative estimates of changes in Sphagnum N concentration in response to N application. However, glasshouse-based estimates of changes in production--even qualitative assessments-- diverged from field experiments owing to a stronger N effect on production response in absence of vascular plants in the glasshouse, and a weaker N effect on production response in presence of vascular plants compared to field experiments. • Thus, although we need glasshouse experiments to study how interacting environmental factors affect the response of Sphagnum to increased N deposition, we need field experiments to properly quantify these effects.

PRECIPITATION DETERMINES THE PERSISTENCE OF HOLLOW SPHAGNUM SPECIES ON HUMMOCKS

On raised bogs, the distribution of Sphagnum species is determined by their distance to the water table, but occasionally species are able to survive outside their niche. Hollow species that persist in hummock vegetation are assumed to profit from the higher water content of the surrounding hummock species, although the mechanism responsible is unclear. In this study, we elucidated the role of lateral hummock water transport (LHWT) and precipitation on the water content of hollow species occurring in hummocks. This was tested using a full factorial field transplantation experiment with cores of Sphagnum cuspidatum in a high and a low hummock. Treatments included direct precipitation (present or absent) and LHWT (present or absent). Fresh weights of the cores were measured at regular time intervals. Our results show a relatively large effect of precipitation on the water content in both the high and low hummock, whereas LHWT only seemed to be an important source of water in the high hummock, which was relatively dry. Furthermore, LHWT played an important role only after large precipitation events, suggesting that lateral water transport is indirectly affected by rain. This study shows that precipitation alone can explain the persistence of hollow species in high hummocks, whereas it was less important for hollow species in low hummocks. Our data suggest that the survival and potential expansion of hollow species in higher hummocks strongly depends on the intensity and frequency of rain events. Changes in precipitation patterns may result in a loss of Sphagnum diversity in hummocks.

Spatio-temporal trends of nitrogen deposition and climate effects on Sphagnum productivity in European peatlands

To quantify potential nitrogen (N) deposition impacts on peatland carbon (C) uptake, we explored temporal and spatial trends in N deposition and climate impacts on the production of the key peat forming functional group (Sphagnum mosses) across European peatlands for the period 1900-2050. Using a modelling approach we estimated that between 1900 and 1950 N deposition impacts remained limited irrespective of geographical position. Between 1950 and 2000 N deposition depressed production between 0 and 25% relative to 1900, particularly in temperate regions. Future scenarios indicate this trend will continue and become more pronounced with climate warming. At the European scale, the consequences for Sphagnum net C-uptake remained small relative to 1900 due to the low peatland cover in high-N areas. The predicted impacts of likely changes in N deposition on Sphagnum productivity appeared to be less than those of climate. Nevertheless, current critical loads for peatlands are likely to hold under a future climate.

Embryo dune development drivers: beach morphology, growing season precipitation, and storms

For development of embryo dunes on the highly dynamic land–sea boundary, summer growth and the absence of winter erosion are essential. Other than that, however, we know little about the specific conditions that favour embryo dune development. This study explores the boundary conditions for early dune development to enable better predictions of natural dune expansion. Using a 30 year time series of aerial photographs of 33 sites along the Dutch coast, we assessed the influence of beach morphology (beach width and tidal range), meteorological conditions (storm characteristics, wind speed, growing season precipitation, and temperature), and sand nourishment on early dune development. We examined the presence and area of embryo dunes in relation to beach width and tidal range, and compared changes in embryo dune area to meteorological conditions and whether sand nourishment had been applied. We found that the presence and area of embryo dunes increased with increasing beach width. Over time, embryo dune area was negatively correlated with storm intensity and frequency. Embryo dune area was positively correlated with precipitation in the growing season and sand nourishment. Embryo dune area increased in periods of low storm frequency and in wet summers, and decreased in periods of high storm frequency or intensity. We conclude that beach morphology is highly influential in determining the potential for new dune development, and wide beaches enable development of larger embryo dune fields. Sand nourishment stimulates dune development by increasing beach width. Finally, weather conditions and non-interrupted sequences of years without high-intensity storms determine whether progressive dune development will take place. Copyright