Scale-dependent effects of herbivory on moss communities in Arctic wetlands: A 25-year experiment.

Arctic ecosystems are undergoing rapid changes, including increasing disturbance by herbivore populations, which can affect plant species coexistence and community assemblages. Although the significance of mosses in Arctic wetlands is well recognized, the long-term influence of medium-sized herbivores on the composition of moss communities has received limited attention. We used data from a long-term (25 years) Greater Snow Goose (Anser caerulescens atlanticus) exclusion experiment in Arctic tundra wetlands to assess changes in the composition of moss communities at multiple spatial scales (cell, 4 cm2; quadrat, 100 cm2; exclosure, 16 m2). We investigated how snow goose grazing and grubbing can alter the composition of the moss community by measuring changes in alpha and beta diversity, as well as in the strength of plant interspecific interactions between moss species. Our results indicate that goose foraging significantly increased species diversity (richness, evenness, and inverse Simpson index) of moss communities at the cell and quadrat scales but not the exclosure scale. Goose foraging reduced the dissimilarity (beta diversity) of moss communities at all three scales, mainly due to decreased species turnover. Furthermore, goose foraging increased positive interaction between moss species pairs. These findings emphasize the critical role of geese in promoting moss species coexistence and increasing homogeneity in Arctic wetlands. This study illustrates how top-down regulation by herbivores can alter plant communities in Arctic wetlands and highlights the importance of considering herbivores when examining the response of Arctic plant biodiversity to future climate change.

Rewetting increases vegetation cover and net growing season carbon uptake under fen conditions after peat-extraction in Manitoba, Canada.

The moss layer transfer technique has been developed to restore the carbon sequestration function and typical vegetation of Sphagnum-dominated peatlands after peat extraction in North America. However, the technique does not lead to successful bryophyte establishment when applied to peatlands with a richer residual fen peat. Therefore, we evaluated an alternative method of active rewetting and passive vegetation establishment using vegetation surveys and carbon dioxide and methane (CH4) flux measurements at a post-extracted fen in southern Manitoba, Canada. After one growing season post-rewetting, wetland vegetation established and the site was a net carbon sink over the growing season. However, high abundance of Carex lasiocarpa 10 years post-treatment led to higher CH4 emissions than the reference ecosystem. Successful establishment of wetland vegetation is attributed to the area being surrounded by undisturbed fens that can provide a local source of plant propagules. Bryophyte expansion was less successful than vascular plants, likely due to episodic flooding and shading from the sedge communities. Therefore, careful management of water levels to just below the peat surface is needed if reference vegetation community recovery is the goal of restoration. Water level management will also play a key role in controlling CH4 emissions to maximize carbon sequestration potential.

Multi-year net ecosystem carbon balance of a restored peatland reveals a return to carbon sink.

Peatlands after drainage and extraction are large sources of carbon (C) to the atmosphere. Restoration, through re-wetting and revegetation, aims to return the C sink function by re-establishing conditions similar to that of an undrained peatland. However, the time needed to re-establish C sequestration is not well constrained due to the lack of multi-year measurements. We measured over 3 years the net ecosystem exchange of CO2 (NEE), methane ( F CH 4 ), and dissolved organic carbon (DOC) at a restored post-extraction peatland (RES) in southeast Canada (restored 14 years prior to the start of the study) and compared our observations to the C balance of an intact reference peatland (REF) that has a long-term continuous flux record and is in the same climate zone. Small but significant differences in winter respiration driven by temperature were mainly responsible for differences in cumulative NEE between years. Low growing season inter-annual variability was linked to constancy of the initial spring water table position, controlled by the blocked drainage ditches and the presence of water storage structures (bunds and pools). Half-hour F CH 4 at RES was small except when Typha latifolia-invaded drainage ditches were in the tower footprint; this effect at the ecosystem level was small as ditches represent a minor fraction of RES. The restored peatland was an annual sink for CO2 (-90 ± 18 g C m-2  year-1 ), a source of CH4 (4.4 ± 0.2 g C m-2  year-1 ), and a source of DOC (6.9 ± 2.2 g C m-2  year-1 ), resulting in mean net ecosystem uptake of 78 ± 17 g C m-2  year-1 . Annual NEE at RES was most similar to wetter, more productive years at REF. Integrating structures to increase water retention, alongside re-establishing key species, have been effective at re-establishing the net C sink rate to that of an intact peatland.

The importance of pH and sand substrate in the revegetation of saline non-waterlogged peat fields.

A partially peat-extracted coastal bog contaminated by seawater was barren and required revegetation as a wetland. Peat fields were rectangular in shape, cambered in cross-section profile, and separated by drainage ditches. Common to all peat fields were symmetrical patterns in micro-topography with slopes between differences in elevation. Saline non-waterlogged slopes of ∼5% occurred as a symmetrical pair on each side of the crest of the cambered profile, at one end of each peat field. Three rows were laid across this slope (Top, Middle, and Bottom rows) and transplanted with naturally-growing plant species with their sand substrate, in three experiments, and grown for a year. In the Spartina pectinata experiment, bare root stem sections were also planted. Another experiment was conducted to determine changes in the characteristics of a volume of sand when incubated in saline peat fields. We found the salinity of peat increased with moisture downslope, and pH decreased with increase in salinity. S. pectinata grew best when planted with its sand substrate compared with bare root stem section, and when planted in Bottom rows. Juncus balticus had excellent growth in all rows. Unexpectedly, Festuca rubra that was inconspicuous beneath the J. balticus canopy in the natural donor site grew densely within the J. balticus sods. Agrostis stolonifera grew well but seemed to show intolerance to the surrounding acidic peat by curling up its stolons. The pH of the incubated sand volume was much higher than the surrounding peat. These studies suggest that recognition of plant niches and pH manipulation are important in the revegetation of disturbed Sphagnum peatlands that are found abundantly in the northern hemisphere. Results are also relevant to the reclamation of other disturbed lands.

Manure derived biochar can successfully replace phosphate rock amendment in peatland restoration.

Phosphate rock fertilization is commonly used in peatland restoration to promote the growth of Polytrichum strictum, a nurse plant which aids the establishment of Sphagnum mosses. The present study tested whether 1) phosphorus fertilization facilitates the germination of P. strictum spores and 2) biochar derived from local pig manure can replace imported phosphate rock currently used in peatland restoration. Various doses of biochar were compared to phosphate rock to test its effect directly on P. strictum stem regeneration (in Petri dishes in a growth chamber) and in a simulation of peatland restoration with the moss layer transfer technique (in mesocoms in a greenhouse). Phosphorus fertilization promoted the germination of P. strictum spores as well as vegetative stem development. Biochar can effectively replace phosphate rock in peatland restoration giving a new waste management option for rural regions with phosphorus surpluses. As more available phosphorus was present in biochar, an addition of only 3-9 g m(-2) of pig manure biochar is recommended during the peatland restoration process, which is less than the standard dose of phosphate rock (15 g m(-2)).

Evolution of niche preference in Sphagnum peat mosses.

Peat mosses (Sphagnum) are ecosystem engineers-species in boreal peatlands simultaneously create and inhabit narrow habitat preferences along two microhabitat gradients: an ionic gradient and a hydrological hummock-hollow gradient. In this article, we demonstrate the connections between microhabitat preference and phylogeny in Sphagnum. Using a dataset of 39 species of Sphagnum, with an 18-locus DNA alignment and an ecological dataset encompassing three large published studies, we tested for phylogenetic signal and within-genus changes in evolutionary rate of eight niche descriptors and two multivariate niche gradients. We find little to no evidence for phylogenetic signal in most component descriptors of the ionic gradient, but interspecific variation along the hummock-hollow gradient shows considerable phylogenetic signal. We find support for a change in the rate of niche evolution within the genus-the hummock-forming subgenus Acutifolia has evolved along the multivariate hummock-hollow gradient faster than the hollow-inhabiting subgenus Cuspidata. Because peat mosses themselves create some of the ecological gradients constituting their own habitats, the classic microtopography of Sphagnum-dominated peatlands is maintained by evolutionary constraints and the biological properties of related Sphagnum species. The patterns of phylogenetic signal observed here will instruct future study on the role of functional traits in peatland growth and reconstruction.

Ecological restoration of rich fens in Europe and North America: from trial and error to an evidence-based approach.

Fens represent a large array of ecosystem services, including the highest biodiversity found among wetlands, hydrological services, water purification and carbon sequestration. Land-use change and drainage has severely damaged or annihilated these services in many parts of North America and Europe; restoration plans are urgently needed at the landscape level. We review the major constraints on the restoration of rich fens and fen water bodies in agricultural areas in Europe and disturbed landscapes in North America: (i) habitat quality problems: drought, eutrophication, acidification, and toxicity, and (ii) recolonization problems: species pools, ecosystem fragmentation and connectivity, genetic variability, and invasive species; and here provide possible solutions. We discuss both positive and negative consequences of restoration measures, and their causes. The restoration of wetland ecosystem functioning and services has, for a long time, been based on a trial-and-error approach. By presenting research and practice on the restoration of rich fen ecosystems within agricultural areas, we demonstrate the importance of biogeochemical and ecological knowledge at different spatial scales for the management and restoration of biodiversity, water quality, carbon sequestration and other ecosystem services, especially in a changing climate. We define target processes that enable scientists, nature managers, water managers and policy makers to choose between different measures and to predict restoration prospects for different types of deteriorated fens and their starting conditions.

Landscape analysis of nutrient-enriched margins (lagg) in ombrotrophic peatlands.

Scientific knowledge of the wet zone - the lagg - that tends to form at the edge of ombrotrophic peatlands is surprisingly limited. In this study, we aim to improve the understanding of the ecohydrological functions of this transition by describing the form and abiotic controls of the laggs and margins of bog peatlands. Data collected in wells and piezometers along 10 transects (within 6 bogs), of the New Brunswick Eastern Lowlands are used to analyse the hydraulic and hydrochemical gradients, while airborne LiDAR data provides new insight on the geomorphology and the vegetation patterns of the bog-lagg-mineral transition zone. Based on their geomorphic character, the study transects are placed into 2 categories: confined and unconfined. Laggs of confined transition are found in a topographic depression, between the bog and a mineral slope >1%, while laggs of unconfined transitions are adjacent to a flat (≤1%) or receding mineral slope (sloping away from the lagg). Water level (4 ± 9 cm vs. -3 ± 9 cm), pH (4.8 ± 0.9 vs. 4.2 ± 0.4), electrical conductivity (ECcorr) (105 ± 52 µS cm(-1) vs. 52 ± 28 µS cm(-1)) and peat depth (55 ± 9 cm vs. 30 ± 9 cm) are found to be higher, respectively, for the confined laggs than for the unconfined. Saturated hydraulic conductivity (Ksat) of the lagg's upper peat layer resembles that of bog environments, but quickly reduces with depth, impeding vertical water flow. The greatest abiotic control of the lagg appears to be topography, which affects water flow rates and direction, thus water chemistry, nutrient transport and availability, hence vegetation characteristics. Our results suggest that the features of the transition zone that include the lagg, influence the quantity and variability of water within the adjacent peatland, and should be considered as integral part of the peatland complex.

Impacts of oil sands process water on fen plants: implications for plant selection in required reclamation projects.

Fen plant growth in peat contaminated with groundwater discharges of oil sands process water (OSPW) was assessed in a greenhouse over two growing seasons. Three treatments (non-diluted OSPW, diluted OSPW and rainwater) were tested on five vascular plants and four mosses. All vascular plants tested can grow in salinity and naphthenic acids levels currently produced by oil sands activity in northwestern Canada. No stress sign was observed after both seasons. Because of plant characteristics, Carex species (C. atherodes and C. utriculata) and Triglochin maritima would be more useful for rapidly restoring vegetation and creating a new peat-accumulating system. Groundwater discharge of OSPW proved detrimental to mosses under dry conditions and ensuring adequate water levels would be crucial in fen creation following oil sands exploitation. Campylium stellatum would be the best choice to grow in contaminated areas and Bryum pseudotriquetrum might be interesting as it has spontaneously regenerated in all treatments.

Trophic interactions in a high arctic snow goose colony.

We examined the role of trophic interactions in structuring a high arctic tundra community characterized by a large breeding colony of greater snow geese (Chen caerulescens atlantica). According to the exploitation ecosystem hypothesis of Oksanen et al. (1981), food chains are controlled by top-down interactions. However, because the arctic primary productivity is low, herbivore populations are too small to support functional predator populations and these communities should thus be dominated by the plant/ herbivore trophic-level interaction. Since 1990, we have been monitoring annual abundance and productivity of geese, the impact of goose grazing, predator abundance (mostly arctic foxes, Alopex lagopus) and the abundance of lemmings, the other significant herbivore in this community, on Bylot Island, Nunavut, Canada. Goose grazing consistently removed a significant proportion of the standing crop (∼40%) in tundra wetlands every year. Grazing changed plant community composition and reduced the production of grasses and sedges to a low-level equilibrium compared to the situation where the presence of geese had been removed. Lemming cyclic fluctuations were strong and affected fox reproduction. Fox predation on goose eggs was severe and generated marked annual variation in goose productivity. Predation intensity on geese was closely related to the lemming cycle, a consequence of an indirect interaction between lemming and geese via shared predators. We conclude that, contrary to the exploitation ecosystem hypothesis, both the plant/herbivore and predator/prey interactions are significant in this arctic community.