Host tree availability shapes potential distribution of a target epiphytic moss species more than direct climate effects.

Climate change significantly impacts the distribution of woody plants, indirectly influencing the dynamics of entire ecosystems. Understanding species' varied responses to the environment and their reliance on biotic interactions is crucial for predicting the global changes' impact on woodland biodiversity. Our study focusses on Dicranum viride, a moss of conservation priority, and its dependence on specific phorophytes (host trees). Using species distribution modelling (SDM) techniques, we initially modelled its distribution using climate-only variables. As a novel approach, we also modelled the distribution of the main phorophyte species and incorporated them into D. viride SDM alongside climate data. Finally, we analysed the overlap of climatic and geographic niches between the epiphyte and the phorophytes. Inclusion of biotic interactions significantly improved model performance, with phorophyte availability emerging as the primary predictor. This underscores the significance of epiphyte-phorophyte interactions, supported by substantial niche overlap. Predictions indicate a potential decline in the suitability of most of the current areas for D. viride, with noticeable shifts towards the northern regions of Europe. Our study underscores the importance of incorporating biotic interactions into SDMs, especially for dependent organisms. Understanding such connections is essential to implement successful conservation strategies and adapt forest management practices to environmental changes.

Moss species and precipitation mediate experimental warming stimulation of growing season N2 fixation in subarctic tundra.

Climate change in high latitude regions leads to both higher temperatures and more precipitation but their combined effects on terrestrial ecosystem processes are poorly understood. In nitrogen (N) limited and often moss-dominated tundra and boreal ecosystems, moss-associated N2 fixation is an important process that provides new N. We tested whether high mean annual precipitation enhanced experimental warming effects on growing season N2 fixation in three common arctic-boreal moss species adapted to different moisture conditions and evaluated their N contribution to the landscape level. We measured in situ N2 fixation rates in Hylocomium splendens, Pleurozium schreberi and Sphagnum spp. from June to September in subarctic tundra in Sweden. We exposed mosses occurring along a natural precipitation gradient (mean annual precipitation: 571-1155 mm) to 8 years of experimental summer warming using open-top chambers before our measurements. We modelled species-specific seasonal N input to the ecosystem at the colony and landscape level. Higher mean annual precipitation clearly increased N2 fixation, especially during peak growing season and in feather mosses. For Sphagnum-associated N2 fixation, high mean annual precipitation reversed a small negative warming response. By contrast, in the dry-adapted feather moss species higher mean annual precipitation led to negative warming effects. Modelled total growing season N inputs for Sphagnum spp. colonies were two to three times that of feather mosses at an area basis. However, at the landscape level where feather mosses were more abundant, they contributed 50% more N than Sphagnum. The discrepancy between modelled estimates of species-specific N input via N2 fixation at the moss core versus ecosystem scale, exemplify how moss cover is essential for evaluating impact of altered N2 fixation. Importantly, combined effects of warming and higher mean annual precipitation may not lead to similar responses across moss species, which could affect moss fitness and their abilities to buffer environmental changes.

Plasmodesmata dynamics in bryophyte model organisms: secondary formation and developmental modifications of structure and function.

MAIN CONCLUSION: Developing bryophytes differentially modify their plasmodesmata structure and function. Secondary plasmodesmata formation via twinning appears to be an ancestral trait. Plasmodesmata networks in hornwort sporophyte meristems resemble those of angiosperms. All land-plant taxa use plasmodesmata (PD) cell connections for symplasmic communication. In angiosperm development, PD networks undergo an extensive remodeling by structural and functional PD modifications, and by postcytokinetic formation of additional secondary PD (secPD). Since comparable information on PD dynamics is scarce for the embryophyte sister groups, we investigated maturating tissues of Anthoceros agrestis (hornwort), Physcomitrium patens (moss), and Marchantia polymorpha (liverwort). As in angiosperms, quantitative electron microscopy revealed secPD formation via twinning in gametophytes of all model bryophytes, which gives rise to laterally adjacent PD pairs or to complex branched PD. This finding suggests that PD twinning is an ancient evolutionary mechanism to adjust PD numbers during wall expansion. Moreover, all bryophyte gametophytes modify their existing PD via taxon-specific strategies resembling those of angiosperms. Development of type II-like PD morphotypes with enlarged diameters or formation of pit pairs might be required to maintain PD transport rates during wall thickening. Similar to angiosperm leaves, fluorescence redistribution after photobleaching revealed a considerable reduction of the PD permeability in maturating P. patens phyllids. In contrast to previous reports on monoplex meristems of bryophyte gametophytes with single initials, we observed targeted secPD formation in the multi-initial basal meristems of A. agrestis sporophytes. Their PD networks share typical features of multi-initial angiosperm meristems, which may hint at a putative homologous origin. We also discuss that monoplex and multi-initial meristems may require distinct types of PD networks, with or without secPD formation, to control maintenance of initial identity and positional signaling.

Cell death in bryophytes: emerging models to study core regulatory modules and conserved pathways.

This review summarizes recent progress in our current understanding of the mechanisms underlying the cell death pathways in bryophytes, focusing on conserved pathways and particularities in comparison to angiosperms. Regulated cell death (RCD) plays key roles during essential processes along the plant life cycle. It is part of specific developmental programmes and maintains homeostasis of the organism in response to unfavourable environments. Bryophytes could provide valuable models to study developmental RCD processes as well as those triggered by biotic and abiotic stresses. Some pathways analogous to those present in angiosperms occur in the gametophytic haploid generation of bryophytes, allowing direct genetic studies. In this review, we focus on such RCD programmes, identifying core conserved mechanisms and raising new key questions to analyse RCD from an evolutionary perspective.

Dehydration and rehydration differently affect photosynthesis and volatile monoterpenes in bryophytes with contrasting ecological traits.

Bryophytes desiccate rapidly when relative humidity decreases. The capacity to withstand dehydration depends on several ecological and physiological factors. Volatile organic compounds (VOCs) may have a role in enhancing tolerance to desiccating bryophytes. However, the functions of VOCs in bryophytes have received little attention so far. We aimed to investigate the impact of a dehydration-rehydration treatment on primary carbon metabolism and volatile terpenes (VTs) in three bryophytes with contrasting ecological traits: Vessicularia dubyana, Porella platyphylla and Pleurochaete squarrosa. First, we confirmed the desiccation sensitivity gradient of the species. Under fully hydrated conditions, the photosynthetic rate (A) was inversely associated with stress tolerance, with a lower rate in more tolerant species. The partial recovery of A in P. platyphylla and P. squarrosa after rehydration confirmed the desiccation tolerance of these two species. On the other hand, A did not recover after rehydration in V. dubyana. Regarding VT, each species exhibited a distinct VT profile under optimum hydration, with the highest VT pool found in the more desiccation-sensitive species (V. dubyana). However, the observed species-specific VT pattern could be associated with the ecological habitat of each species. P. squarrosa, a moss of dry habitats, may synthesize mainly non-volatile secondary metabolites as stress-defensive compounds. On the other hand, V. dubyana, commonly found submerged, may need to invest photosynthetically assimilated carbon to synthesize a higher amount of VTs to cope with transient water stress occurrence. Further research on the functions of VTs in bryophytes is needed to deepen our understanding of their ecological significance.

Terrestrial mosses as a substrate and potential host for cyanobacteria, green algae and diatoms.

Most studies of terrestrial bryophytes as natural substrates for photosynthetic microorganisms have been performed in the polar regions, where bryophytes are an important part of the ecosystem. As they remain green throughout the year, bryophytes may also be an ideal substrate for epiphytic organisms in temperate regions. The present study investigated the colonization potential and diversity of microalgae on selected plant species in riparian forest and spruce monoculture in a temperate region. It examines whether the presence of algae is related to substrate humidity, the micromorphology of gametophyte or the seasonal availability of substrate. The taxonomic diversity of algae was studied. Cyanobacteria and green algae were cultured on BG-11 agar medium, while diatoms were identified in permanent diatomaceous slides. The alpha- and beta-diversity indices were calculated, and the communities were compared using Bray-Curtis distances and multidimensional correspondence analyses. Our findings indicate that the largest number of alga species were diatoms; however, their presence was only observed in riparian forest and was associated with high humidity. Both aerophilic and freshwater taxa were noted, the latter carried by water from nearby aquatic ecosystem. Green algae were present in both phytocoenoses and humidity appears to have no substantial effect on the degree of colonization; their diversity was low and the group consisted of terrestrial taxa. In two bryophytes growing at the highest humidity, cyanobacteria were only identified in culture. The key factor influencing the degree of microalgae colonization was the humidity of the substrate, which was related to the distance from water.

Impact of changing climate on bryophyte contributions to terrestrial water, carbon, and nitrogen cycles.

Bryophytes, including the lineages of mosses, liverworts, and hornworts, are the second-largest photoautotroph group on Earth. Recent work across terrestrial ecosystems has highlighted how bryophytes retain and control water, fix substantial amounts of carbon (C), and contribute to nitrogen (N) cycles in forests (boreal, temperate, and tropical), tundra, peatlands, grasslands, and deserts. Understanding how changing climate affects bryophyte contributions to global cycles in different ecosystems is of primary importance. However, because of their small physical size, bryophytes have been largely ignored in research on water, C, and N cycles at global scales. Here, we review the literature on how bryophytes influence global biogeochemical cycles, and we highlight that while some aspects of global change represent critical tipping points for survival, bryophytes may also buffer many ecosystems from change due to their capacity for water, C, and N uptake and storage. However, as the thresholds of resistance of bryophytes to temperature and precipitation regime changes are mostly unknown, it is challenging to predict how long this buffering capacity will remain functional. Furthermore, as ecosystems shift their global distribution in response to changing climate, the size of different bryophyte-influenced biomes will change, resulting in shifts in the magnitude of bryophyte impacts on global ecosystem functions.

Morphological Innovation Drives Sperm Release in Bryophytes.

Plant movements for survival are nontrivial. Antheridia in the moss Physcomitrium patens (P. patens) use motion to eject sperm in the presence of water. However, the biological and mechanical mechanisms that actuate the process are unknown. Here, the burst of the antheridium of P. patens, triggered by water, results from elastic instability and is determined by an asymmetric change in cell geometry. The tension generated in jacket cell walls of antheridium arises from turgor pressure, and is further promoted when the inner walls of apex burst in hydration, causing water and cellular contents of apex quickly influx into sperm chamber. The outer walls of the jacket cells are strengthened by NAC transcription factor VNS4 and serve as key morphomechanical innovations to store hydrostatic energy in a confined space in P. patens. However, the antheridium in liverwort Marchantia polymorpha (M. polymorpha) adopts a different strategy for sperm release; like jacket cell outer walls of P. patens, the cells surrounding the antheridium of M. polymorpha appear to play a similar role in the storage of energy. Collectively, the work shows that plants have evolved different ingenious devices for sperm discharge and that morphological innovations can differ.

Changes in bryophyte functional composition during post-fire succession.

Climate and land-use changes are altering fire regimes in many regions around the world. To date, most studies have focused on the effects of altered fire regimes on woody and herbaceous communities, while the mechanisms driving post-fire bryophyte succession remain poorly understood, particularly in Mediterranean-type ecosystems. Here, we examined changes in bryophyte functional composition along a post-fire chronosequence (ranging from 1 to 20+ years) in Pyrenean oak woodlands (northeastern Portugal). To do so, we defined bryophyte functional groups based on seven morphological, reproductive, and life history traits. Then, we fitted linear and structural equation models to disentangle the direct and indirect effects of fire (time since fire and fire intensity), vegetation structure, climate, topography, and edaphic conditions on the abundance of each group. We identified two main functional groups: early colonizers (species with traits associated with strong colonization ability and desiccation tolerance) and perennial stayers (species with high competitive ability, i.e., large perennial mosses). Overall, the abundance of early colonizer species decreased with time since fire and increased with fire intensity, while the opposite was observed for perennial stayers. Thus, successional dynamics reflected a trade-off between species' competitive and colonization abilities, highlighting the role of biotic interactions later in succession. Patterns of functional composition were also consistent with changes in environmental conditions during succession, suggesting that species may experience stressful conditions (i.e., high radiation and low water availability) in early stages of post-fire succession. Our results also indicate that increased fire intensity may alter successional trajectories, leading to long-term changes in bryophyte communities. By understanding the response of bryophyte communities to fire, we were able to identify species with potential use as soil restoration materials.

Biocrusts enhance soil organic carbon stability and regulate the fate of new-input carbon in semiarid desert ecosystems.

Given their global prevalence, dryland (including hyperarid, arid, semiarid, and dry subhumid regions) ecosystems are critical for supporting soil organic carbon (SOC) stocks, with even small changes in such SOC pools affecting the global carbon (C) cycling. Biocrusts play an essential role in supporting C cycling in semiarid ecosystems. However, the influence of biocrusts and their successional stages on SOC and its fraction contents, as well as their role in regulating new input C into SOC fractions remain largely unknown. In this study, we collected continuous samples of bare soil (BS) and three successional stages of biocrust soils (cyanobacterial (CC), low-cover moss (LM), and high-cover moss (HM)) at 0-5 cm depth every month for one year in a semiarid desert ecosystem. We analyzed SOC changes among the samples and their fraction contents including: labile organic C (LOC) (composed of microbial biomass C (MBC), dissolved organic C (DOC), and easily oxidized organic C (EOC)) and recalcitrant organic C (ROC) fractions, soil nutrient content including: ammonium (NH4+-N), nitrate (NO3--N), and available phosphorus (AP), and soil temperature and moisture. We also conducted a 13C pulse-labelling experiment in the field to accurately quantify the effects of biocrust successional stage on exogenous C allocation to SOC fractions. Our results showed that the three successional stages of biocrust (CC-LM-HM) increased SOC and ROC contents by an average of 5.3 ± 3.6 g kg-1 and 4.0 ± 3.0 g kg-1, respectively; and the MBC, DOC, and EOC contents increased by an average of 41.7 ± 24.8 mg kg-1, 28.7 ± 12.6 mg kg-1, and 1.2 ± 0.6 g kg-1, respectively, compared to that of BS. These increases were attributed to an increase in photosynthetic pigment content, higher nutrient levels, and more suitable microclimates (e.g., higher moisture and more moderate temperature) during biocrust succession. More importantly, SOC stability was greatly improved with biocrust succession from cyanobacteria to moss, as evidenced by the reduction in soil EOC:SOC and EOC:ROC ratios by an average of 50 ± 34 % and 99 ± 67 %, respectively, while the ROC:SOC ratio increased by 33 ± 16 % with biocrust succession compared to those of BS. The biocrust SOC, DOC, and MBC 13C contents at different stages were on average 0.096 ± 0.034 mg kg-1, 0.010 ± 0.005 mg kg-1, and 0.014 ± 0.005 mg kg-1 higher than those of BS. Similarly, the allocation of new-input C among the DOC and MBC at different biocrust stages (19 ± 10 %) was significantly higher than that of BS (9 ± 6 %). New-input C into the biocrusts was fixed by microbes (43 ± 18 %) within ∼10 days and converted into other forms of C (85 ± 5 %) after 80 days. Our study provides a new perspective on how biocrusts support C cycling in semiarid desert ecosystems by mediating new C inputs into diverse fractional contents, and highlights the significance of biocrust successional stages in maintaining soil C stocks and stability in the dryland soil system.

Mapping biocrust distribution in China's drylands under changing climate.

Biological soil crusts (biocrusts) are widely distributed in global drylands and have multiple significant roles in regulating dryland soil and ecosystem multifunctionality. However, maps of their distribution over large spatial scales are uncommon and sometimes unreliable, because our current remote sensing technology is unable to efficiently discriminate between biocrusts and vascular plants or even bare soil across different ecosystem and soil types. The lack of biocrust spatial data may limit our ability to detect risks to dryland function or key tipping points. Here, we indirectly mapped biocrust distribution in China's drylands using spatial prediction modeling, based on a set of occurrences of biocrusts (379 in total) and high-resolution soil and environmental data. The results showed that biocrusts currently cover 13.9 % of China's drylands (or 5.7 % of China's total area), with moss-, lichen-, and cyanobacterial-dominated biocrusts each occupying 5.7 % to 10.7 % of the region. Biocrust distribution is mainly determined by soil properties (soil type and contents of gravel and nitrogen), aridity stress, and altitude. Their most favorable habitat is arenosols with low contents of gravel and nitrogen, in climate with a drought index of 0.54 and an altitude of about 500 m. By 2050, climate change will lead to a 5.5 %-9.0 % reduction in biocrust cover. Lichen biocrusts exhibit a high vulnerability to climate change, with potential reductions of up to 19.0 % in coverage. Biocrust cover loss is primarily caused by the combined effects of the elevated temperature and increased precipitation. Our study provides the first high-resolution (250 × 250 m) map of biocrust distribution in China's drylands and offers a reliable approach for mapping regional or global biocrust colonization. We suggest incorporating biocrusts into Earth system models to identify their significant impact on global or regional-scale processes under climate change.

Adaptive evolution of the enigmatic Takakia now facing climate change in Tibet.

The most extreme environments are the most vulnerable to transformation under a rapidly changing climate. These ecosystems harbor some of the most specialized species, which will likely suffer the highest extinction rates. We document the steepest temperature increase (2010-2021) on record at altitudes of above 4,000 m, triggering a decline of the relictual and highly adapted moss Takakia lepidozioides. Its de-novo-sequenced genome with 27,467 protein-coding genes includes distinct adaptations to abiotic stresses and comprises the largest number of fast-evolving genes under positive selection. The uplift of the study site in the last 65 million years has resulted in life-threatening UV-B radiation and drastically reduced temperatures, and we detected several of the molecular adaptations of Takakia to these environmental changes. Surprisingly, specific morphological features likely occurred earlier than 165 mya in much warmer environments. Following nearly 400 million years of evolution and resilience, this species is now facing extinction.

Thermophilisation of communities differs between land plant lineages, land use types and elevation.

Bryophytes provide key ecosystem services at the global scale such as carbon storage and primary production in resource limited habitats, but compared to vascular plants knowledge on how these organisms face recent climate warming is fragmentary. This is particularly critical because bryophytes differ fundamentally from vascular plants in their ecophysiological and biological characteristics, so that community alterations most likely have different dynamics. In a comparative approach, we analysed thermophilisation of bryophyte and vascular plant communities in 1146 permanent plots distributed along an elevational gradient of nearly 3.000 m in Switzerland (Central Europe) that were visited in 5-years intervals between 2001 and 2021. We estimated thermophilisation from changes in unweighted mean temperature indicator values of species, compared it to expected thermophilisation rates given the shift of isotherms and addressed differences between the two lineages, major land use types (managed grasslands, forests, unmanaged open areas), life strategy types (long- and short-lived species) and in elevation. Thermophilisation of bryophyte communities was on average 2.1 times higher than of vascular plant communities and at high elevations it approximated the expected rate given the shift of isotherms. Thermophilisation of both, bryophyte and vascular plant communities was not driven by a loss of cryophilic species but by an increase in thermophilic and mesophilic species, indicating an in-filling process. Furthermore, our data show that thermophilisation is higher in managed grasslands than in forests. We suggest that the higher responsiveness of bryophytes compared to vascular plants depends on their poikilohydry and dispersal capacity and that lower thermophilisation of forests communities is related to the buffering effect of microclimatic conditions in the interior of forests. Our study emphasises the heterogeneity of climate warming effects on plants because response dynamics differ between taxonomic groups as well as between land use types and along elevational gradients.

Nitrogen fixation associated with two cohabiting moss species expresses different patterns under Cu and Zn contamination.

Nitrogen (N2) fixation by moss-associated cyanobacteria is an important N source in pristine ecosystems. Previous studies have shown that moss-associated N2 fixation is sensitive to anthropogenic N pollution. However, we still lack understanding of the effects of other factors derived from anthropogenic sources, such as heavy metal pollution on N2 fixation. To test this, we collected two dominant mosses (Pleurozium schreberi and Spaghnum palustre) from a temperate bog in Denmark and assessed their N2 fixation responses to simulated heavy metal pollution by adding 5 levels (plus a control) of copper (Cu, 0-0.05 mg g dw-1) and zinc (Zn, 0-0.1 mg g dw-1). Metal concentrations in both mosses increased linearly with Cu and Zn addition, but N2 fixation activity associated with S. palustre was to a greater extent negatively affected by both Cu and Zn additions than that associated with P. schreberi. Copper additions even promoted N2 fixation in P. schreberi. Hence, the heavy metal sensitivity of N2-fixing cyanobacteria is dependent on the host moss-species, and the vulnerability of ecosystems towards heavy metal pollution could vary depending on the dominant moss species.

Disentangling climate change from air pollution effects on epiphytic bryophytes.

At the interface between atmosphere and vegetation, epiphytic floras have been largely used as indicators of air quality. The recovery of epiphytes from high levels of SO2 pollution has resulted in major range changes, whose interpretation has, however, been challenged by concomitant variation in other pollutants as well as climate change. Here, we combine historical and contemporary information on epiphytic bryophyte species distributions, climatic conditions, and pollution loads since the 1980s in southern Belgium to disentangle the relative impact of climate change and air pollution on temporal shifts in species composition. The relationship between the temporal variation of species composition, climatic conditions, SO2 , NO2 , O3 , and fine particle concentrations, was analyzed by variation partitioning. The temporal shift in species composition was such, that it was, on average, more than twice larger than the change in species composition observed today among communities scattered across the study area. The main driver, contributing to 38% of this temporal shift in species composition, was the variation of air quality. Climate change alone did not contribute to the substantial compositional shifts in epiphytic bryophyte communities in the course of the last 40 years. As a consequence of the substantial drop of N and S loads over the last decades, present-day variations of epiphytic floras were, however, better explained by the spatial variation of climatic conditions than by extant pollution loads. The lack of any signature of recolonization delays of formerly polluted areas in the composition of modern floras suggests that epiphytic bryophytes efficiently disperse at the landscape scale. We suggest that a monitoring of epiphyte communities at 10-year intervals would be desirable to assess the impact of raising pollution sources, and especially pesticides, whose impact on bryophytes remains poorly documented.

Sub-arctic mosses and lichens show idiosyncratic responses to combinations of winter heatwaves, freezing and nitrogen deposition.

Arctic ecosystems are increasingly exposed to extreme climatic events throughout the year, which can affect species performance. Cryptogams (bryophytes and lichens) provide important ecosystem services in polar ecosystems but may be physiologically affected or killed by extreme events. Through field and laboratory manipulations, we compared physiological responses of seven dominant sub-Arctic cryptogams (three bryophytes, four lichens) to single events and factorial combinations of mid-winter heatwave (6°C for 7 days), re-freezing, snow removal and summer nitrogen addition. We aimed to identify which mosses and lichens are vulnerable to these abiotic extremes and if combinations would exacerbate physiological responses. Combinations of extremes resulted in stronger species responses but included idiosyncratic species-specific responses. Species that remained dormant during winter (March), irrespective of extremes, showed little physiological response during summer (August). However, winter physiological activity, and response to winter extremes, was not consistently associated with summer physiological impacts. Winter extremes affect cryptogam physiology, but summer responses appear mild, and lichens affect the photobiont more than the mycobiont. Accounting for Arctic cryptogam response to multiple climatic extremes in ecosystem functioning and modelling will require a better understanding of their winter eco-physiology and repair capabilities.

A research agenda for nonvascular photoautotrophs under climate change.

Nonvascular photoautotrophs (NVP), including bryophytes, lichens, terrestrial algae, and cyanobacteria, are increasingly recognized as being essential to ecosystem functioning in many regions of the world. Current research suggests that climate change may pose a substantial threat to NVP, but the extent to which this will affect the associated ecosystem functions and services is highly uncertain. Here, we propose a research agenda to address this urgent question, focusing on physiological and ecological processes that link NVP to ecosystem functions while also taking into account the substantial taxonomic diversity across multiple ecosystem types. Accordingly, we developed a new categorization scheme, based on microclimatic gradients, which simplifies the high physiological and morphological diversity of NVP and world-wide distribution with respect to several broad habitat types. We found that habitat-specific ecosystem functions of NVP will likely be substantially affected by climate change, and more quantitative process understanding is required on: (1) potential for acclimation; (2) response to elevated CO2 ; (3) role of the microbiome; and (4) feedback to (micro)climate. We suggest an integrative approach of innovative, multimethod laboratory and field experiments and ecophysiological modelling, for which sustained scientific collaboration on NVP research will be essential.

Chemical Stimulation of Heterocyte Differentiation by the Feather Moss Hylocomium splendens: a Potential New Step in Plant-Cyanobacteria Symbioses.

Cyanobacteria associated with mosses play a key role in the nitrogen (N) cycle in unpolluted ecosystems. Mosses have been found to release molecules that induce morphophysiological changes in epiphytic cyanobionts. Nevertheless, the extent of moss influence on these microorganisms remains unknown. To evaluate how mosses or their metabolites influence N2 fixation rates by cyanobacteria, we assessed the nitrogenase activity, heterocyte frequency and biomass of a cyanobacterial strain isolated from the feather moss Hylocomium splendens and a non-symbiotic strain when they were either growing by themselves, together with H. splendens or exposed to H. splendens water, acetone, ethanol, or isopropanol extracts. The same cyanobacterial strains were added to another moss (Taxiphyllum barbieri) and a liverwort (Monosolenium tenerum) to assess if these bryophytes affect N2 fixation differently. Although no significant increases in nitrogenase activity by the cyanobacteria were observed when in contact with H. splendens shoots, both the symbiotic and non-symbiotic cyanobacteria increased nitrogenase activity as well as heterocyte frequency significantly upon exposure to H. splendens ethanol extracts. Contact with T. barbieri shoots, on the other hand, did lead to increases in nitrogenase activity, indicating low host-specificity to cyanobacterial activity. These findings suggest that H. splendens produces heterocyte-differentiating factors (HDFs) that are capable of stimulating cyanobacterial N2 fixation regardless of symbiotic competency. Based on previous knowledge about the chemical ecology and dynamics of moss-cyanobacteria interactions, we speculate that HDF expression by the host takes place in a hypothetical new step occurring after plant colonization and the repression of hormogonia.

Biocrust impacts on dryland soil water balance: A path toward the whole picture.

As a crucial living feature inhabiting the soil-atmosphere boundary, biocrusts play a vital role in liquid water or vapor transport through surface soil and thus have strong effects on soil water regimes. However, it remains unclear how biocrusts affect annual or multiyear soil water budgets through the regulation of evaporation outputs and non-rainfall water (NRW) or infiltration inputs. Thus, we used automated microlysimeters to continually investigate the differences in evaporation and NRW rates between moss-dominated biocrusts and bare soil at 0-5 cm depth for 2 years. The upper 30 cm of soil moisture (θ) and water storage (W) of bare soil and biocrusts were also monitored. Our results showed that the daily evaporation rate (E) of biocrusts was 17% higher than bare soil. Especially after rainfall events, biocrusts had higher E and larger cumulative evaporation than bare soil. Besides, the daily NRW of biocrusts averaged 15% higher than bare soil over 2 years. Furthermore, biocrusts increased θ by 11%-76% at 0-10 cm depth but decreased θ by 32%-56% at 20-30 cm depth in comparison to bare soil, and they subsequently decreased W by 20% at 0-30 cm depth. Summarized annually, the NRW amount of biocrusts was 19% higher than bare soil, but at the same time, the cumulative evaporation of biocrusts was also 19% higher than bare soil. Finally, biocrusts resulted in more water loss at shallow depth through evaporation and lessened total W throughout 0-30 cm depth of soil. These findings demonstrate that although biocrusts input more NRW into surface soil, these water inputs partially offset their intensified evaporation. Given that all rainfall water infiltrates into the soil in our study system, our findings indicate that biocrusts may have an overall negative effect on soil water balance there, while at the same time increasing water storage and availability of the deeper soil underlying biocrusts.