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1. The circumboreal/circumpolar N2-fixing lichen Stereocaulon vesuvianum is among the most widespread and abundant fruticose species in montane Britain but has lost the capacity to fix N2 over large areas of the country. 2. To investigate whether loss of N2-fixation in S. vesuvianum is linked to increased N deposition, we examined thallus morphology, physiology and chemistry at twelve locations representing an N deposition gradient of 3–40 kg ha−1 year−1. Measurements were made in parallel on a non-N2-fixing reference species (Parmelia saxatilis). The presence or absence of cephalodia (N2-fixing nodules containing the cyanobacterium Stigonema sp) was recorded in over 500 herbarium specimens of S. vesuvianum dating back to 1820. 3. Cephalodium abundance in S. vesuvianum, and 15N concentration in S. vesuvianum and P. saxatilis, were strongly negatively correlated with N deposition and particularly with dry deposited N; cephalodia do not form at total N deposition rates ≥8–9 kg ha−1 year−1. Other morphological oddities in S. vesuvianum at N-polluted sites include increased apothecium (fungal reproductive structure) production and green algal biofilm development. Biofilm covered thalli without cephalodia lacked nitrogenase activity and cephalodia at sites where they rarely develop had nitrogenase activities typical for this species. The presence or absence of cephalodia in herbarium specimens of S. vesuvianum suggest that the present-day N-deposition linked gradient in N2-fixing capacity did not exist in the 19th century and largely developed between 1900–1940. 4. Synthesis. We provide clear evidence that N2-fixing capacity in S. vesuvianum has been lost in regions subjected to many decades of enhanced atmospheric N deposition. This loss is consistent with established models of diazotrophy, which identify supply of combined N as an inhibitor of N2-fixation. Progressive depletion of thallus 15N with increasing N deposition is in line with available data indicating that m

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.

Himalayan forests are biodiverse and support the cultural and economic livelihoods of their human communities. They are bounded to the south by the Indo-Gangetic Plain, which has among the highest concentrations of atmospheric ammonia globally. This source of excess nitrogen pushes northwards into the Himalaya, generating concern that Himalayan forests will be impacted. To estimate the extent to which atmospheric nitrogen is impacting Himalayan forests we focussed on lichen epiphytes, which are a well-established bioindicator for atmospheric nitrogen pollution. First, we reviewed published literature describing nitrogen thresholds (critical levels and loads) at which lichen epiphytes are affected, identifying a mean and confidence intervals based on previous research conducted across a diverse set of biogeographic and ecological settings. Second, we used estimates from previously published atmospheric chemistry models (EMEP-WRF and UKCA-CLASSIC) projected to the Himalaya with contrasting spatial resolution and timescales to characterise model variability. Comparing the lichen epiphyte critical levels and loads with the atmospheric chemistry model projections, we created preliminary estimates of the extent to which Himalayan forests are impacted by excess nitrogen; this equated to c. 80–85% and c. 95–98% with respect to ammonia and total nitrogen deposition, respectively. Recognising that lichens are one of the most sensitive bioindicators for atmospheric nitrogen pollution, our new synthesis of previous studies on this topic generated concern that most Himalayan forests are at risk from excess nitrogen. This is a desk-based study that now requires verification through biological surveillance, for which we provide key recommendations.