Bjorn J. M. Robroek

The effect of temperature on growth and competition between Sphagnum species

Peat bogs play a large role in the global sequestration of C, and are often dominated by different Sphagnum species. Therefore, it is crucial to understand how Sphagnum vegetation in peat bogs will respond to global warming. We performed a greenhouse experiment to study the effect of four temperature treatments (11.2, 14.7, 18.0 and 21.4°C) on the growth of four Sphagnum species: S. fuscum and S. balticum from a site in northern Sweden and S. magellanicum and S. cuspidatum from a site in southern Sweden. In addition, three combinations of these species were made to study the effect of temperature on competition. We found that all species increased their height increment and biomass production with an increase in temperature, while bulk densities were lower at higher temperatures. The hollow species S. cuspidatum was the least responsive species, whereas the hummock species S. fuscum increased biomass production 13-fold from the lowest to the highest temperature treatment in monocultures. Nutrient concentrations were higher at higher temperatures, especially N concentrations of S. fuscum and S. balticum increased compared to field values. Competition between S. cuspidatum and S. magellanicum was not influenced by temperature. The mixtures of S. balticum with S. fuscum and S. balticum with S. magellanicum showed that S. balticum was the stronger competitor, but it lost competitive advantage in the highest temperature treatment. These findings suggest that species abundances will shift in response to global warming, particularly at northern sites where hollow species will lose competitive strength relative to hummock species and southern species.

Plant functional types define magnitude of drought response in peatland CO2 exchange.

Peatlands are important sinks for atmospheric carbon (C), yet the role of plant functional types (PFTs) for C sequestration under climatic perturbations is still unclear. A plant-removal experiment was used to study the importance of vascular PFTs for the net ecosystem CO2 exchange (NEE) during (i.e., resistance) and after (i.e., recovery) an experimental drought. The removal of PFTs caused a decrease of NEE, but the rate differed between microhabitats (i.e., hummocks and lawns) and the type of PFTs. Ericoid removal had a large effect on NEE in hummocks, while the graminoids played a major role in the lawns. The removal of PFTs did not affect the resistance or the recovery after the experimental drought. We argue that the response of Sphagnum mosses (the only PFT present in all treatments) to drought is dominant over that of coexisting PFTs. However, we observed that the moment in time when the system switched from C sink to C source during the drought was controlled by the vascular PFTs. In the light of climate change, the shifts in species composition or even the loss of certain PFTs are expected to strongly affect the future C dynamics in response to environmental stress.

Field Simulation of Global Change: Transplanting Northern Bog Mesocosms Southward

A large proportion of northern peatlands consists of Sphagnum-dominated ombrotrophic bogs. In these bogs, peat mosses (Sphagnum) and vascular plants occur in an apparent stable equilibrium, thereby sustaining the carbon sink function of the bog ecosystem. How global warming and increased nitrogen (N) deposition will affect the species composition in bog vegetation is still unclear. We performed a transplantation experiment in which mesocosms with intact vegetation were transplanted southward from north Sweden to north-east Germany along a transect of four bog sites, in which both temperature and N deposition increased. In addition, we monitored undisturbed vegetation in control plots at the four sites of the latitudinal gradient. Four growing seasons after transplantation, ericaceous dwarf shrubs had become much more abundant when transplanted to the warmest site which also had highest N deposition. As a result ericoid aboveground biomass in the transplanted mesocosms increased most at the southernmost site, this site also had highest ericoid biomass in the undisturbed vegetation. The two dominant Sphagnum species showed opposing responses when transplanted southward; Sphagnum balticum height increment decreased, whereas S. fuscum height increment increased when transplanted southward. Sphagnum production did not differ significantly among the transplanted mesocosms, but was lowest in the southernmost control plots. The dwarf shrub expansion and increased N concentrations in plant tissues we observed, point in the direction of a positive feedback toward vascular plant-dominance suppressing peat-forming Sphagnum in the long term. However, our data also indicate that precipitation and phosphorus availability influence the competitive balance between Sphagnum, dwarf shrubs and graminoids.

Peatland vascular plant functional types affect methane dynamics by altering microbial community structure

Peatlands are natural sources of atmospheric methane (CH4), an important greenhouse gas. It is established that peatland methane dynamics are controlled by both biotic and abiotic conditions, yet the interactive effect of these drivers is less studied and consequently poorly understood. Climate change affects the distribution of vascular plant functional types (PFTs) in peatlands. By removing specific PFTs, we assessed their effects on peat organic matter chemistry, microbial community composition and on potential methane production (PMP) and oxidation (PMO) in two microhabitats (lawns and hummocks). Whilst PFT removal only marginally altered the peat organic matter chemistry, we observed considerable changes in microbial community structure. This resulted in altered PMP and PMO. PMP was slightly lower when graminoids were removed, whilst PMO was highest in the absence of both vascular PFTs (graminoids and ericoids), but only in the hummocks. Path analyses demonstrate that different plant-soil interactions drive PMP and PMO in peatlands and that changes in biotic and abiotic factors can have auto-amplifying effects on current CH4 dynamics.Synthesis. Changing environmental conditions will, both directly and indirectly, affect peatland processes, causing unforeseen changes in CH4 dynamics. The resilience of peatland CH4 dynamics to environmental change therefore depends on the interaction between plant community composition and microbial communities.

An unexpected role for mixotrophs in the response of peatland carbon cycling to climate warming

Mixotrophic protists are increasingly recognized for their significant contribution to carbon (C) cycling. As phototrophs they contribute to photosynthetic C fixation, whilst as predators of decomposers, they indirectly influence organic matter decomposition. Despite these direct and indirect effects on the C cycle, little is known about the responses of peatland mixotrophs to climate change and the potential consequences for the peatland C cycle. With a combination of field and microcosm experiments, we show that mixotrophs in the Sphagnum bryosphere play an important role in modulating peatland C cycle responses to experimental warming. We found that five years of consecutive summer warming with peaks of +2 to +8°C led to a 50% reduction in the biomass of the dominant mixotrophs, the mixotrophic testate amoebae (MTA). The biomass of other microbial groups (including decomposers) did not change, suggesting MTA to be particularly sensitive to temperature. In a microcosm experiment under controlled conditions, we then manipulated the abundance of MTA, and showed that the reported 50% reduction of MTA biomass in the field was linked to a significant reduction of net C uptake (-13%) of the entire Sphagnum bryosphere. Our findings suggest that reduced abundance of MTA with climate warming could lead to reduced peatland C fixation.

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.

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.

Snow cover manipulation effects on microbial community structure and soil chemistry in a mountain bog

Background and AimsAlterations in snow cover driven by climate change may impact ecosystem functioning, including biogeochemistry and soil (microbial) processes. We elucidated the effects of snow cover manipulation (SCM) on above-and belowground processes in a temperate peatland.MethodsIn a Swiss mountain-peatland we manipulated snow cover (addition, removal and control), and assessed the effects on Andromeda polifolia root enzyme activity, soil microbial community structure, and leaf tissue and soil biogeochemistry.ResultsReduced snow cover produced warmer soils in our experiment while increased snow cover kept soil temperatures close-to-freezing. SCM had a major influence on the microbial community, and prolonged ‘close-to-freezing’ temperatures caused a shift in microbial communities toward fungal dominance. Soil temperature largely explained soil microbial structure, while other descriptors such as root enzyme activity and pore-water chemistry interacted less with the soil microbial communities.ConclusionsWe envisage that SCM-driven changes in the microbial community composition could lead to substantial changes in trophic fluxes and associated ecosystem processes. Hence, we need to improve our understanding on the impact of frost and freeze-thaw cycles on the microbial food web and its implications for peatland ecosystem processes in a changing climate; in particular for the fate of the sequestered carbon.

Sensitivity of blanket peat vegetation and hydrochemistry to local disturbances

At the ecosystem scale, peatlands can be extremely resilient to perturbations. Yet, they are very sensitive to local disturbances, especially mechanical perturbations (e.g. trampling). The effects of these disturbances on vegetation, and potential effects on hydrochemical conditions along the peat surface, however, are largely unknown. We used three research tracks (paths researchers use to access their study sites) differing in time of abandonment to investigate the impact of local disturbance (trampling) on the vegetation and its short-term (< or = 2 year) recovery in a flagship research blanket peatland. Additionally, we examined the effects of local disturbance on fluvial runoff events and the concentrations of dissolved organic carbon (DOC) and particulate organic carbon (POC) in runoff water. Local disturbance heavily impacted peat vegetation, resulting in large areas of scarred and churned peat. Recovery of vascular plants along abandoned tracks was slow, but a functional Sphagnum layer re-established after just one year. The absence of vegetation elicited an increase in the number of runoff events along the tracks, by which POC runoff from the tracks increased. POC concentrations were highest in the surface water from the recently abandoned track, while they were low in the runoff water from the track abandoned longest and the undisturbed control track. We attribute this to the relatively fast recovery of the Sphagnum vegetation. DOC concentrations did not differ significantly either spatially or temporally in surface runoff or soil solution waters. While at an ecosystem scale local disturbances may be negligible in terms of carbon loss, our data points to the need for further research on the potential long-term effects of local disturbance on the vegetation, and significant effects on local scale carbon fluxes. Moreover, the effects of disturbances could be long-lasting and their role on ecosystem processes should not be underestimated.

Subordinate plants mitigate drought effects on soil ecosystem processes by stimulating fungi

1. The subordinate insurance hypothesis suggests that highly diverse communities contain greater numbers of subordinate species than less diverse communities. It has previously been reported that subordinate species can improve grassland productivity during drought, but the underlying mechanisms remain undetermined. 2. Using a combination of subordinate species removal and summer drought, we show that soil processes play a critical role in community resistance to drought. Interestingly, subordinate species drive soil microbial community structure and largely mitigate the effect of drought on grassland soil functioning. Our results highlight subordinate species in shifting the balance within the phospholipid fatty acid (PLFA) microbial community towards more fungal dominance. 3. Fungal communities promoted by subordinate species were more resistant to drought and maintained higher rates of litter decomposition and soil respiration. These results emphasize the important role of subordinate species in mitigating drought effects on soil ecosystem functions. Reciprocal effects between fungi and subordinate species explain also how subordinate species better resisted to drought conditions. 4. Our results point to a delayed plant-soil feedback following environmental perturbation. Additionally, they extend the diversity insurance hypothesis by showing that more diverse communities not only contain species well adapted to perturbations, but also species with higher impacts on soil microbial communities and related ecosystem functions.