Habitat patchiness, ecological connectivity and the uneven recovery of boreal stream ecosystems from an experimental drought

Ongoing climate change is increasing the occurrence and intensity of drought episodes worldwide, including in boreal regions not previously regarded as drought prone, and where the impacts of drought remain poorly understood. Ecological connectivity is one factor that might influence community structure and ecosystem functioning post-drought, by facilitating the recovery of sensitive species via dispersal at both local (e.g. a nearby habitat patch) and regional (from other systems within the same region) scales. In an outdoor mesocosm experiment, we investigated how impacts of drought on boreal stream ecosystems are altered by the spatial arrangement of local habitat patches within stream channels, and variation in ecological connectivity with a regional species pool. We measured basal ecosystem processes underlying carbon and nutrient cycling: (a) algal biomass accrual; (b) microbial respiration; and (c) decomposition of organic matter, and sampled communities of aquatic fungi and benthic invertebrates. An 8-day drought event had strong impacts on both community structure and ecosystem functioning, including algal accrual, leaf decomposition and microbial respiration, with many of these impacts persisting even after water levels had been restored for 3.5 weeks. Enhanced connectivity with the regional species pool and increased aggregation of habitat patches also affected multiple response variables, especially those associated with microbes, and in some cases reduced the effects of drought to a small extent. This indicates that spatial processes might play a role in the resilience of communities and ecosystem functioning, given enough time. These effects were however insufficient to facilitate significant recovery in algal growth before seasonal dieback began in autumn. The limited resilience of ecosystem functioning in our experiment suggests that even short-term droughts can have extended consequences for stream ecosystems in the world's vast boreal region, and especially on the ecosystem processes and services mediated by algal biofilms

Anthropogenic impacts and restoration of boreal spring ecosystems

Abstract Human activities have increasingly altered freshwater ecosystems. Land use is a major driver of habitat loss and land use-related input of nutrients and other pollutants from agriculture, forestry and urbanization have deteriorated water quality. Freshwater research has mainly focused on lakes and streams while the effects of anthropogenic stressors on groundwater-dependent ecosystems (GDEs) are poorly known. Likewise, the effectiveness of ecological restoration in mitigating human disturbance in GDEs remains understudied. In this thesis, I studied the effects of two main anthropogenic stressors – land drainage and groundwater contamination – on boreal spring ecosystems and evaluated the recovery of spring biodiversity and ecosystem functioning after habitat restoration. I applied several structural (macroinvertebrates, bryophytes, leaf-decomposing fungi and groundwater bacteria) and functional (organic matter decomposition and primary productivity) measures to provide a comprehensive insight into these issues. Both stressors modified spring ecosystems. Land drainage reduced the key ecosystem processes. Long-term monitoring of drainage-impacted springs showed a marked biodiversity loss and change of spring-dwelling bryophytes, and no signs of recovery were observed after about 20 years since the intial land drainage. Groundwater contamination, indicated by elevated nitrate and chloride concentrations, altered the structure of spring biota, reduced their taxonomic diversity and suppressed primary productivity in the most severely contaminated springs. Spring restoration improved habitat quality by reducing drainage-induced inflow of surface water, thus re-establishing groundwater-dominated hydrological conditions. Restoration increased abundance of habitat-specialist bryophytes and shifted macroinvertebrate composition towards natural conditions, despite the restoration actions being fairly recent. Anthropogenic activities can thus cause severe structural and functional degradation of spring ecosystems, and their self-recovery potential from these stressors seems low. Habitat restoration bears great promise as a cost-effective approach to mitigate drainage-induced impacts on spring ecosystems, but protection and co-management of groundwater resources are urgently needed to secure the role of springs as biodiversity hotspots in the boreal forest landscape.Tiivistelmä Ihmistoiminta muuttaa yhä enemmän vesiekosysteemejä. Maankäyttö on johtanut elinympäristöjen häviämiseen, ja siihen liittyvä ravinne- ja haitta-ainekuormitus maa- ja metsätaloudesta sekä kaupunkiympäristöistä on merkittävästi huonontanut veden laatua johtaen maailmanlaajuiseen vesiluonnon monimuotoisuuden heikentymiseen. Vesiekosysteemien tutkimus on keskittynyt pääasiassa järvi- ja jokiympäristöihin, kun ihmistoiminnan vaikutukset pohjavesiriippuvaisiin ekosysteemeihin tunnetaan edelleen huonosti. Samoin kunnostusten merkitys pohjavesiriippuvaisten ekosysteemien tilan parantamiseksi on selvittämättä. Väitöskirjassani tarkastelin kahden keskeisen ihmistoiminnan – metsäojituksen ja pohjaveden laadun heikkenemisen – vaikutuksia lähde-ekosysteemeihin sekä arvioin elinympäristökunnostusten vaikutuksia niiden rakenteeseen ja toimintaan. Sovelsin työssäni rakenteellisia (pohjaeläimet, sammalet, lehtikariketta hajottavat sienet ja pohjavesibakteerit) ja toiminnallisia (eloperäisen aineksen hajoaminen ja perustuotanto) mittareita tuottamaan kattavan käsityksen tutkimuskysymyksiini. Sekä metsäojitukset että pohjaveden laadun heikkeneminen aiheuttavat muutoksia lähteiden rakenteessa ja toiminnassa. Metsäojitukset hidastavat keskeisiä ekosysteemitoimintoja ja johtavat lähdesammallajiston muutokseen ja monimuotoisuuden taantumiseen. Pohjaveden pilaantuminen, jota työssä ilmennettiin kohonneilla nitraatti- ja kloridipitoisuuksilla, heikentää lähdelajiston monimuotoisuutta, muuttaa lajikoostumusta ja johtaa perustuotannon laskuun voimakkaimmin kuormitetuissa lähteissä. Kunnostus parantaa lähde-elinympäristön laatua vähentämällä metsäojien aiheuttamaa pintavesivaikutusta palauttaen pohjavesivaltaisen hydrologisen tilan. Lähdekunnostusten myötä lähdesammaleet runsastuvat ja pohjaeläinyhteisön rakenne palautuu luonnontilaisten lähteiden kaltaiseksi, vaikka kunnostuksista on kulunut vasta muutamia vuosia. Väitöskirjan tulokset osoittavat, että ihmisen toiminta voi aiheuttaa muutoksia lähde-ekosysteemien rakenteessa ja toiminnassa ja lähteiden luontainen palautuminen häiriöstä on hidasta. Lähde-elinympäristöjen kunnostus vaikuttaa lupaavalta suojelutoimenpiteeltä metsäojitusten vaikutusten vähentämisessä, mutta lähteiden säilyttäminen monimuotoisena ja suojelullisesti arvokkaana luontotyyppinä edellyttää pohjavesivarojen hallinnan ja tilan suojelun tehostamista

Effect of riparian soil moisture on bacterial, fungal and plant communities and microbial decomposition rates in boreal stream-side forests

Riparian habitats of boreal forests are considered as hotspots for biochemical processes and biodiversity, and varying width riparian buffers have been proposed to protect species diversity of the riparian forests. However, evidence of the role of soil moisture variation in shaping riparian biodiversity and ecosystem functioning remain scarce particularly regarding belowground diversity. We studied how distance from the stream and soil moisture of the riparian zone affected species richness and community composition of plants, bacteria, and fungi as well as microbial decomposition rates. Using a split-plot design with a plant survey and amplicon sequencing for microorganisms we identified taxa associated with different categories of moisture and distance from the stream along six headwater stream-sides in middle boreal forests in Northern Finland. Tea-bag Index was used to assess the decomposition rates. PERMANOVA and linear mixed-effect models were used to analyze the data. Variation in riparian soil moisture influenced species composition and richness of plants and bacteria. Plant communities also changed from herbaceous dominated to shrub dominated with increasing distance from the stream. Fungal communities, however, did not respond to soil moisture or distance from the stream, and there were only slight differences in fungal trophic guilds among moisture and distance categories. Decomposition of organic material by microorganisms was faster adjacent to the stream than further away, and moist riparian areas had higher decomposition rates than drier ones. Decomposition rates were positively related to pH, Ca, Mg and NH4 and soil temperature. Synthesis and applications We show that above- and belowground diversity and microbial decomposition are associated to soil moisture at riparian sites supporting the idea of leaving wider unmanaged buffers in moist habitats to safeguard the overall forest diversity. Our findings further emphasize the need to consider soil moisture when planning the measures for riparian protection as changes in riparian soil moisture could lead to deterioration of organic matter decomposition. Different responses of the examined plant and microbial communities to riparian soil conditions clearly imply that overall riparian diversity cannot be explained based on a single community type, and that different organisms may respond differently to human-induced changes in stream riparian zone. Highlights • Soil moisture influences plant and bacterial diversity, but not fungal. • Microbial driven decomposition is faster in moist riparian areas than on dry ones. • Wider buffers in moist habitats could help safeguard the overall forest diversity. • Overall riparian diversity cannot be explained based on a single community type. • Ecosystem functioning should be considered in riparian protection planning

Effect of riparian soil moisture on bacterial, fungal and plant communities and microbial decomposition rates in boreal stream-side forests

Highlights •Soil moisture influences plant and bacterial diversity, but not fungal. •Microbial driven decomposition is faster in moist riparian areas than on dry ones. •Wider buffers in moist habitats could help safeguard the overall forest diversity. •Overall riparian diversity cannot be explained based on a single community type. •Ecosystem functioning should be considered in riparian protection planning. Abstract Riparian habitats of boreal forests are considered as hotspots for biochemical processes and biodiversity, and varying width riparian buffers have been proposed to protect species diversity of the riparian forests. However, evidence of the role of soil moisture variation in shaping riparian biodiversity and ecosystem functioning remain scarce particularly regarding belowground diversity. We studied how distance from the stream and soil moisture of the riparian zone affected species richness and community composition of plants, bacteria, and fungi as well as microbial decomposition rates. Using a split-plot design with a plant survey and amplicon sequencing for microorganisms we identified taxa associated with different categories of moisture and distance from the stream along six headwater stream-sides in middle boreal forests in Northern Finland. Tea-bag Index was used to assess the decomposition rates. PERMANOVA and linear mixed-effect models were used to analyze the data. Variation in riparian soil moisture influenced species composition and richness of plants and bacteria. Plant communities also changed from herbaceous dominated to shrub dominated with increasing distance from the stream. Fungal communities, however, did not respond to soil moisture or distance from the stream, and there were only slight differences in fungal trophic guilds among moisture and distance categories. Decomposition of organic material by microorganisms was faster adjacent to the stream than further away, and moist riparian areas had higher decomposition rates than drier ones. Decomposition rates were positively related to pH, Ca, Mg and NH4 and soil temperature. Synthesis and applications We show that above- and belowground diversity and microbial decomposition are associated to soil moisture at riparian sites supporting the idea of leaving wider unmanaged buffers in moist habitats to safeguard the overall forest diversity. Our findings further emphasize the need to consider soil moisture when planning the measures for riparian protection as changes in riparian soil moisture could lead to deterioration of organic matter decomposition. Different responses of the examined plant and microbial communities to riparian soil conditions clearly imply that overall riparian diversity cannot be explained based on a single community type, and that different organisms may respond differently to human-induced changes in stream riparian zone

Bacterial communities in a subarctic stream network:spatial and seasonal patterns of benthic biofilm and bacterioplankton

Abstract Water-column bacterial communities are assembled by different mechanisms at different stream network positions, with headwater communities being controlled by mass effects (advection of bacteria from terrestrial soils) while downstream communities are mainly driven by environmental sorting. Conversely, benthic biofilms are colonized largely by the same set of taxa across the entire network. However, direct comparisons of biofilm and bacterioplankton communities along whole stream networks are rare. We used 16S rRNA gene amplicon sequencing to explore the spatiotemporal variability of benthic biofilm (2 weeks old vs. mature biofilm) and water-column communities at different network positions of a subarctic stream from early summer to late autumn. Amplicon sequence variant (ASV) richness of mature biofilm was about 2.5 times higher than that of early biofilm, yet the pattern of seasonality was the same, with the highest richness in midsummer. Biofilm bacterial richness was unrelated to network position whereas bacterioplankton diversity was negatively related to water residence time and distance from the source. This pattern of decreasing diversity along the network was strongest around midsummer and diminished greatly as water level increased towards autumn. Biofilm communities were phylogenetically clustered at all network positions while bacterioplankton assemblages were phylogenetically clustered only at the most downstream site. Both early and mature biofilm communities already differed significantly between upstream (1st order) and midstream (2nd order) sections. Network position was also related to variation in bacterioplankton communities, with upstream sites harbouring substantially more unique taxa (44% of all upstream taxa) than midstream (20%) or downstream (8%) sites. Some of the taxa that were dominant in downstream sections were already present in the upmost headwaters, and even in riparian soils, where they were very rare (relative abundance <0.01%). These patterns in species diversity and taxonomic and phylogenetic community composition of the riverine bacterial metacommunity were particularly strong for water-column communities, whereas both early and mature biofilm exhibited weaker spatial patterns. Our study demonstrated the benefits of studying bacterioplankton and biofilm communities simultaneously to allow testing of ecological hypotheses about biodiversity patterns in freshwater bacteria

Streams and riparian forests depend on each other:a review with a special focus on microbes

Abstract In this review, we draw together the research on the two-way connection of streams and their riparian forests of the boreal zone from ecological points of view. Although the knowledge about stream-riparian interactions has increased considerably recently, in practice, riparian zones are still mainly seen as buffers for nutrient and sediment loading. However, recent research has shown that riparian forests disproportionately foster regional biodiversity and maintain stream ecosystem functions and diversity. On the other hand, streams contribute to riparian diversity and ecosystem functions. Microbes are key drivers of global biochemical cycles, and they also interact with plants and animals. The knowledge on microbial communities and understanding of processes they drive has considerably increased due to recent development in microbial profiling methods. However, microbes have been largely neglected in former reviews. Thus, this overview has a special focus on the role of microorganisms in controlling stream-riparian interaction. We also review the land-use pressures that are threatening biodiversity and ecosystem processes of riparian zones in forested landscapes. In addition, we review the possible effects of climate change on stream-riparian interactions. Finally, we outline the research gaps that call for future research

Bacterial communities at a groundwater-surface water ecotone:gradual change or abrupt transition points along a contamination gradient?

Summary Microbial communities contribute greatly to groundwater quality, but the impacts of land-use practices on bacteria in groundwaters and groundwater-dependent ecosystems remain poorly known. With 16S rRNA gene amplicon sequencing, we assessed bacterial community composition at the groundwater-surface water ecotone of boreal springs impacted by urbanization and agriculture, using spring water nitrate-N as a surrogate of contamination. We also measured the rate of a key ecosystem process, organic matter decomposition. We documented a recurrent pattern across all major bacterial phyla where diversity started to decrease at unexpectedly low nitrate-N concentrations (100–300 μg L−1). At 400 NO3−-N μg L−1, 25 bacterial exact sequence variants showed a negative response, resulting in a distinct threshold in bacterial community composition. Chthonomonas, Acetobacterales and Hyphomicrobium were the most sensitive taxa, while only three taxa (Duganella, Undibacterium and Thermoanaerobaculaceae) were enriched due to increased contamination. Decomposition rate responded unimodally to increasing nitrate-N concentration, with a peak rate at ~400 NO3−-N μg L−1, parallelly with a major shift in bacterial community composition. Our results emphasize the utility of bacterial communities in the assessment of groundwater-dependent ecosystems. They also call for a careful reconsideration of threshold nitrate values for defining groundwater ecosystem health and protecting their microbial biodiversity

Host species shape the community structure of culturable endophytes in fruits of wild berry species (Vaccinium myrtillus L., Empetrum nigrum L. and Vaccinium vitis-idaea L.)

Abstract Wild berries are interesting research subjects due to their rich sources of health-beneficial phenolic compounds. However, the internal microbial communities, endophytes, associated with the wild berry fruits are currently unknown. Endophytes are bacteria or fungi inhabiting inside plant tissues, and their functions vary depending on the host species and environmental parameters. The present study aimed to examine community composition of fungal and bacterial endophytes in fruits of three wild berry species (bilberry Vaccinium myrtillus L., lingonberry Vaccinium vitis-idaea L. and crowberry Empetrum nigrum L.) and the effects of host plant species and their growth sites on shaping the endophytic communities. We found that the endophytic community structures differed between the berry species, and fungi were predominant over bacteria in the total endophytic taxa. We identified previously unknown endophytic fungal taxa including Angustimassarina, Dothidea, Fellozyma, Pseudohyphozyma, Hannaella coprosmae and Oberwinklerozyma straminea. A role of soluble phenolic compounds, the intracellular components in wild berry fruits, in shaping the endophytic communities is proposed. Overall, our study demonstrates that each berry species harbors a unique endophytic community of microbes

Habitat patchiness, ecological connectivity and the uneven recovery of boreal stream ecosystems from an experimental drought

Abstract Ongoing climate change is increasing the occurrence and intensity of drought episodes worldwide, including in boreal regions not previously regarded as drought prone, and where the impacts of drought remain poorly understood. Ecological connectivity is one factor that might influence community structure and ecosystem functioning post‐drought, by facilitating the recovery of sensitive species via dispersal at both local (e.g. a nearby habitat patch) and regional (from other systems within the same region) scales. In an outdoor mesocosm experiment, we investigated how impacts of drought on boreal stream ecosystems are altered by the spatial arrangement of local habitat patches within stream channels, and variation in ecological connectivity with a regional species pool. We measured basal ecosystem processes underlying carbon and nutrient cycling: (a) algal biomass accrual; (b) microbial respiration; and (c) decomposition of organic matter, and sampled communities of aquatic fungi and benthic invertebrates. An 8‐day drought event had strong impacts on both community structure and ecosystem functioning, including algal accrual, leaf decomposition and microbial respiration, with many of these impacts persisting even after water levels had been restored for 3.5 weeks. Enhanced connectivity with the regional species pool and increased aggregation of habitat patches also affected multiple response variables, especially those associated with microbes, and in some cases reduced the effects of drought to a small extent. This indicates that spatial processes might play a role in the resilience of communities and ecosystem functioning, given enough time. These effects were however insufficient to facilitate significant recovery in algal growth before seasonal dieback began in autumn. The limited resilience of ecosystem functioning in our experiment suggests that even short‐term droughts can have extended consequences for stream ecosystems in the world‘s vast boreal region, and especially on the ecosystem processes and services mediated by algal biofilms