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.

Moss functional trait ecology: Trends, gaps, and biases in the current literature.

Functional traits are critical tools in plant ecology for capturing organism-environment interactions based on trade-offs and making links between organismal and ecosystem processes. While broad frameworks for functional traits have been developed for vascular plants, we lack the same for bryophytes, despite an escalation in the number of studies on bryophyte functional trait in the last 45 years and an increased recognition of the ecological roles bryophytes play across ecosystems. In this review, we compiled data from 282 published articles (10,005 records) that focused on functional traits measured in mosses and sought to examine trends in types of traits measured, capture taxonomic and geographic breadth of trait coverage, reveal biases in coverage in the current literature, and develop a bryophyte-function index (BFI) to describe the completeness of current trait coverage and identify global gaps to focus research efforts. The most commonly measured response traits (those related to growth/reproduction in individual organisms) and effect traits (those that directly affect community/ecosystem scale processes) fell into the categories of morphology (e.g., leaf area, shoot height) and nutrient storage/cycling, and our BFI revealed that these data were most commonly collected from temperate and boreal regions of Europe, North America, and East Asia. However, fewer than 10% of known moss species have available functional trait information. Our synthesis revealed a need for research on traits related to ontogeny, sex, and intraspecific plasticity and on co-measurement of traits related to water relations and bryophyte-mediated soil processes.

Responses of Biocrust and Associated Soil Bacteria to Novel Climates Are Not Tightly Coupled.

Climate change is expanding drylands even as land use practices degrade them. Representing ∼40% of Earth's terrestrial surface, drylands rely on biological soil crusts (biocrusts) for key ecosystem functions including soil stability, biogeochemical cycling, and water capture. Understanding how biocrusts adapt to climate change is critical to understanding how dryland ecosystems will function with altered climate. We investigated the sensitivity of biocrusts to experimentally imposed novel climates to track changes in productivity and stability under both warming and cooling scenarios. We established three common gardens along an elevational-climate gradient on the Colorado Plateau. Mature biocrusts were collected from each site and reciprocally transplanted intact. Over 20 months we monitored visible species composition and cover, chlorophyll a, and the composition of soil bacterial communities using high throughput sequencing. We hypothesized that biocrusts replanted at their home site would show local preference, and biocrusts transplanted to novel environments would maintain higher cover and stability at elevations higher than their origin, compared to at elevations lower than their origin. We expected responses of the visible biocrust cover and soil bacterial components of the biocrust community to be coupled, with later successional taxa showing higher sensitivity to novel environments. Only high elevation sourced biocrusts maintained higher biocrust cover and community stability at their site of origin. Biocrusts from all sources had higher cover and stability in the high elevation garden. Later successional taxa decreased cover in low elevation gardens, suggesting successional reversal with warming. Visible community composition was influenced by both source and transplant environment. In contrast, soil bacterial community composition was not influenced by transplant environments but retained fidelity to the source. Thus, responses of the visible and soil bacterial components of the biocrust community were not coupled. Synthesis: Our results suggest biocrust communities are sensitive to climate change, and loss of species and function can be expected, while associated soil bacteria may be buffered against rapid change.

Strategies of desiccation tolerance vary across life phases in the moss Syntrichia caninervis.

PREMISE: Desiccation tolerance (DT) is a widespread phenomenon among land plants, and variable ecological strategies for DT are likely to exist. Using Syntrichia caninervis, a dryland moss and model system used in DT studies, we hypothesized that DT is lowest in juvenile (protonemal) tissues, highest in asexual reproductive propagules (gemmae), and intermediate in adults (shoots). We tested the long-standing hypothesis of an inherent constitutive strategy of DT in this species. METHODS: Plants were rapidly dried to levels of equilibrating relative humidity (RHeq) ranging from 0 to 93%. Postrehydration recovery was assessed using chlorophyll fluorescence, regeneration rates, and visual tissue damage. For each life phase, we estimated the minimum rate of drying (RoDmin ) at RHeq = 42% that did not elicit damage 24 h postrehydration. RESULTS: DT strategy varied with life phase, with adult shoots having the lowest RoDmin (10-25 min), followed by gemmae (3-10 h) and protonema (14-20 h). Adult shoots exhibited no detectable damage 24 h postrehydration following a rapid-dry only at the highest RHeq used (93%), but when dried to lower RHs the response declined to <50% of control fluorescence values. Notably, immediately following rehydration (0 h postrehydration), shoots were damaged below control levels of fluorescence regardless of the RHeq, thus implicating damage. CONCLUSIONS: Life phases of the moss S. caninervis had a range of strategies from near constitutive (adult shoots) to demonstrably inducible (protonema). A new response variable for assessing degree of DT is introduced as the minimum rate of drying from which full recovery occurs.

Morphological and physiological traits in relation to carbon balance in a diverse clade of dryland mosses.

Plant functional trait analyses have focused almost exclusively on vascular plants, but bryophytes comprise ancient and diverse plant lineages that have widespread global distributions and important ecological functions in terrestrial ecosystems. We examined a diverse clade of dryland mosses, Syntrichia, and studied carbon balance during a precipitation event (C-balance), a functional trait related to physiological functioning, desiccation tolerance, survival, and ecosystem carbon and nitrogen cycling. We examined variability in C-balance among 14 genotypes of Syntrichia and measured an additional 10 physiological and 13 morphological traits at the cell, leaf, shoot, and clump level. C-balance varied 20-fold among genotypes, and highest C-balances were associated with long, narrow leaves with awns, and small cells with thick cell walls, traits that may influence water uptake and retention during a precipitation event. Ordination analyses revealed that the axis most strongly correlated with C-balance included the maximum chlorophyll fluorescence, Fm , indicating the importance of photosystem II health for C exchange. C-balance represents a key functional trait in bryophytes, but its measurement is time intensive and not feasible to measure on large scales. We propose two models (using physiological and morphological traits) to predict C-balance, whereby identifying simpler to measure traits for trait databases.

Sex-specific morphological and physiological differences in the moss Ceratodon purpureus (Dicranales).

Background and Aims: Dioecy and sexual dimorphism occur in many terrestrial plant species but are especially widespread among the bryophytes. Despite the prevalence of dioecy in non-vascular plants, surprisingly little is known about how fine-scale sex-specific cell and leaf morphological traits are correlated with sex-specific physiology and population sex ratios. Such data are critical to understanding the inter-relationship between sex-specific morphological and physiological characters and how their relationship influences population structure. In this study, these data types were assessed to determine how they vary across three populations within one moss species and whether fine-scale morphological traits scale up to physiological and sex ratio characteristics. Methods: Twenty cell-, leaf- and canopy-level traits and two photochemical measurements were compared between sexes and populations of the dioecious moss Ceratodon purpureus . Field population-expressed sex ratios were obtained for the same populations. Key Results: Male and female plants differed in cell, leaf and photochemical measures. These sexual dimorphisms were female biased, with females having larger and thicker leaves and greater values for chlorophyll fluorescence-based, leaf photochemistry measurements than males. Female traits were also more variable than male traits. Interestingly, field population sex ratios were significantly male biased in two study populations and female biased in the third study population. Conclusions: The results demonstrate that the larger morphology and the greater physiological output of female C. purpureus gametophytes compared with males occurs across populations and is likely to have significant effects on resource allocation and biotic interactions. However, this high level of dimorphism does not explain population sex ratio variation in the three study populations tested. This research lays the groundwork for future studies on how differential sex-specific variation in cell and leaf traits influences bryophyte plant fitness.