Glacier-fed streams (GFSs) are harsh environments hosting unique, highly specialized communities. Interestingly, glaciers and their GFSs are also present in Earth's tropical regions, where environmental characteristics contrast with GFS conditions elsewhere. Yet, despite the unique and isolated nature of tropical GFSs, little is known about their inhabitants, even though they may disappear later this century with ongoing climate change. Here, we examined diatom communities from one of the last tropical African GFSs in the Rwenzori Mountains, Uganda, to characterize the composition and diversity of this unique system. Six sediment-associated biofilm samples were collected from two reaches of a stream draining the Mt. Stanley Glacier, and the resident diatom communities were studied morphologically using light and scanning electron microscopy, as well as through the sequencing of amplicons from extracted DNA (18S and rbcL). In general, morphological results agree well with barcoding results, but each individually provides irreplaceable insights. In total, we identify 24 morphotypes utilizing light microscopy, 101 diatom Amplicon Sequence Variants (ASVs) using 18S sequences, and 65 ASVs with rbcL. Across approaches, common genera include Achnanthidium, Psammothidium, Neidium, Cymbopleura, Eunotia, and Pinnularia. However, only about half of the diversity could be assigned to the species level across methodologies, including several of the most common taxa, indicating a high level of uniqueness. Accordingly, one of the most common taxa encountered is described here as a new species, Neidium rwenzoriense sp. nov. Our results emphasize the Rwenzori Mountains as a global hotspot for endemism, and the novelty of disappearing tropical GFSs as diatom habitats.

The rapid melting of mountain glaciers and the vanishing of their streams is emblematic of climate change1,2. Glacier-fed streams (GFSs) are cold, oligotrophic and unstable ecosystems in which life is dominated by microbial biofilms2,3. However, current knowledge on the GFS microbiome is scarce4,5, precluding an understanding of its response to glacier shrinkage. Here, by leveraging metabarcoding and metagenomics, we provide a comprehensive survey of bacteria in the benthic microbiome across 152 GFSs draining the Earth’s major mountain ranges. We find that the GFS bacterial microbiome is taxonomically and functionally distinct from other cryospheric microbiomes. GFS bacteria are diverse, with more than half being specific to a given mountain range, some unique to single GFSs and a few cosmopolitan and abundant. We show how geographic isolation and environmental selection shape their biogeography, which is characterized by distinct compositional patterns between mountain ranges and hemispheres. Phylogenetic analyses furthermore uncovered microdiverse clades resulting from environmental selection, probably promoting functional resilience and contributing to GFS bacterial biodiversity and biogeography. Climate-induced glacier shrinkage puts this unique microbiome at risk. Our study provides a global reference for future climate-change microbiology studies on the vanishing GFS ecosystem. Leveraging metabarcoding and metagenomics, a survey of bacteria in the benthic microbiome across 152 glacier-fed streams (GFSs) provides a global reference for future climate-change microbiology studies on the vanishing GFS ecosystem.

We investigated the root microbiome of an endemo-relict plant species of Mount Olympus, Ramonda (Jankaea) heldreichii (Boiss.) C.B.Clarke, at various altitudes (400m-1200m asl), through amplicon sequencing. Microbial communities (prokaryotes, fungi, protists) revealed the significant impact of roots on the tight rhizosphere (TR) that were less diverse and less altitude-impacted compared with the loose rhizosphere (LR). Prokaryotic α-diversity was highly affected by root, whereas that of fungi was comparatively more sensitive to altitude. The TR-associated taxonomic groups, included well equipped taxa for tolerating biotic and abiotic stresses (drought/metal tolerance, microcystin degradation, psychrotolerance, chitin degradation), while the LR-associated taxa mainly included microorganisms with chemolithoautotrophic potential. Relative abundances of the N-cycling and greenhouse gas associated Nitrosopheraceae, were increased with altitude. We explored the root microbiome of the multi-million year old Jankaea species, and took a step towards understanding mountain microbiology, climate change, and bioprospecting, straight from the steep inclines of Mount Olympus.

Runoff from rapidly melting mountain glaciers is a dominant source of riverine organic carbon in many high-latitude and high-elevation regions. Glacier dissolved organic carbon is highly bioavailable, and its composition likely reflects internal (e.g., autotrophic production) and external (i.e., atmospheric deposition) sources. However, the balance of these sources across Earth's glaciers is poorly understood, despite implications for the mineralization and assimilation of glacier organic carbon within recipient ecosystems. We assessed the molecular-level composition of dissolved organic matter from 136 mountain glacier outflows from 11 regions covering six continents using ultrahigh resolution 21 T mass spectrometry. We found substantial diversity in organic matter composition with coherent and predictable (80% accuracy) regional patterns. Employing stable and radiocarbon isotopic analyses, we demonstrate that these patterns are inherently linked to atmospheric deposition and in situ production. In remote regions like Greenland and New Zealand, the glacier organic matter pool appears to be dominated by in situ production. However, downwind of industrial centers (e.g., Alaska and Nepal), fossil fuel combustion byproducts likely underpin organic matter composition, resulting in older and more aromatic material being exported downstream. These findings highlight that the glacier carbon cycle is spatially distinct, with ramifications for predicting the dynamics and fate of glacier organic carbon concurrent with continued retreat and anthropogenic perturbation.

Most cryospheric ecosystems are energy limited. How their energetics will respond to climate change remains largely unknown. This is particularly true for glacier-fed streams, which interface with the cryosphere and initiate some of Earth’s largest river systems. Here, by studying resource stoichiometry and microbial energetics in 154 glacier-fed streams sampled by the Vanishing Glaciers project across Earth’s major mountain ranges, we show that these ecosystems and their benthic microbiome are overall carbon and phosphorus limited. Threshold elemental ratios and low carbon use efficiencies (median: 0.15) modelled from extracellular enzymatic activities corroborate resource limitation in agreement with maintenance metabolism of benthic microorganisms. Space-for-time substitution analyses suggest that glacier shrinkage will stimulate benthic primary production in glacier-fed streams, thereby relieving microbial metabolism from carbon limitation. Concomitantly, we find that increasing streamwater temperature will probably stimulate microbial growth (temperature sensitivity: 0.62 eV). Consequently, elevated microbial demands for phosphorus, but diminishing inputs from subglacial sources, may intensify phosphorus limitation as glaciers shrink. Our study thus unveils a ‘green transition’ towards autotrophy in the world’s glacier-fed streams, entailing shifts in the energetics of their microorganisms.

The glaciers on Africa's 'Mountains of the Moon' (Rwenzori National Park, Uganda) are predicted to disappear within the next decades owing to climate change. Consequently, the glacier-fed streams (GFSs) that drain them will vanish, along with their resident microbial communities. Despite the relevance of microbial communities for performing ecosystem processes in equatorial GFSs, their ecology remains understudied. Here, we show that the benthic microbiome from the Mt. Stanley GFS is distinct at several levels from other GFSs. Specifically, several novel taxa were present, and usually common groups such as Chrysophytes and Polaromonas exhibited lower relative abundances compared to higher-latitude GFSs, while cyanobacteria and diatoms were more abundant. The rich primary producer community in this GFS likely results from the greater environmental stability of the Afrotropics, and accordingly, heterotrophic processes dominated in the bacterial community. Metagenomics revealed that almost all prokaryotes in the Mt. Stanley GFS are capable of organic carbon oxidation, while greater than 80% have the potential for fermentation and acetate oxidation. Our findings suggest a close coupling between photoautotrophs and other microbes in this GFS, and provide a glimpse into the future for high-latitude GFSs globally where primary production is projected to increase with ongoing glacier shrinkage.

Despite the existence of numerous glaciers on the Mediterranean mountains during the Little Ice Age (LIA), many of these disappeared during the 20th century. However, periglacial conditions are sustained in the formerly glaciated alpine zones, preserving relicts of the late Holocene glacial record. The present climate of the Mediterranean mountains is hostile to glaciation and projected climate trends suggest that the Mediterranean cryosphere will be shrinking with immediate impacts on the water budget of the lowlands. Here we show preliminary results of an extensive fieldwork campaign that focuses on the Holocene reconstruction of the climate and alpine critical zone environmental conditions of Mount Olympus (2918 m) in Greece. A well-preserved sequence of late Holocene glacial moraines dating to ⁓2.5 and ⁓0.6 ka BP, respectively, suggest that the small cirque glaciers were geomorphologically active during the LIA, whereas 30 m deep glacial ice found in a perennial ice cave opens a new window of local and regional continuous climate reconstructions. The extensive snowfields of the mid-20th century have shrunk dramatically but have survived the warmest summers of the 21st century. Below these perennial snowfields a 15 m thick permafrost layer has been discovered during our campaign through 3 Electrical Resistivity Tomography (ERT) profiles, in a location where the mean annual air temperature (MAAT) of the last 10 years is above 0oC, but in agreement with permafrost occurrence in other mountains of the Southern Balkan peninsula. The base horizon of postglacial alpine soils overlying glacial till deposited in a glaciokarstic plateau below the summit, appears cryoturbated whereas the soils are characterized by translocation of clay from the upper to the lower horizon. These observations along with occasional early summer soil freeze and subsequent waterlogging, suggest that the periglacial activity on Mount Olympus continues in a rapidly warming Mediterranean environment. However, regional warming and anomalous early summer convective rainfall that has caused a dramatic reduction in the volume of the perennial ice cave deposits and the near extinction of the perennial snowfields (even after winters with very high snow accumulation) over the past 10 years also threatens this periglacial activity. Altogether these observations show the general decreasing trend of the Mediterranean cryosphere and periglacial activity, and they highlight immediate impacts on karstic aquifer water recharge and water availability in the piedmont and coastal zone of Mount Olympus, especially during the summer season when water demand is very high due to agricultural and touristic activities.

Abstract Nearly 60% of the world’s glaciers are predicted to be lost by the end of the century because of global warming1. The impacts of this environmental change on the glacier-fed stream (GFS) ecosystems are profound, putting their ecological communities at risk2–4. Life in GFSs is dominated by benthic microbial biofilms, which regulate key ecosystem processes and form the basis of the food web5 . However, the biodiversity and biogeography of the biofilm microbiome of the world’s GFSs remain unknown, precluding a mechanistic understanding of its responses to glacier shrinkage. Here, applying metabarcoding to the benthic biofilm bacteria from 148 GFSs draining the world’s major mountain ranges, we unveil their global biodiversity and biogeography. We find that the global GFS benthic microbiome differs from other cryospheric microbiomes and is unexpectedly diverse. A large microbiome fraction (58% of the total amplicon sequence variants) is endemic to single mountain ranges, while 0.4% are disproportionately abundant and form a cosmopolitan core. These microbiome components have overlapping taxonomies, but their compositional turnover is differently shaped by dispersal limitation and environmental selection. Phylogenetic analyses relate prevalent clades to endemicity and microdiversity, the latter potentially buffering the GFS microbiome against climate-change impacts. Our global survey provides the basis for future climate-change microbiology studies6 on a rapidly changing ecosystem.

The biogeochemistry of rapidly retreating Andean glaciers is poorly understood, and Ecuadorian glacier dissolved organic matter (DOM) composition is unknown. This study examined molecular composition and carbon isotopes of DOM from supraglacial and outflow streams ( n = 5 and 14, respectively) across five ice capped volcanoes in Ecuador. Compositional metrics were paired with streamwater isotope analyses (δ 18 O) to assess if outflow DOM composition was associated with regional precipitation gradients and thus an atmospheric origin of glacier DOM. Ecuadorian glacier outflows exported ancient, biolabile dissolved organic carbon (DOC), and DOM contained a high relative abundance (RA) of aliphatic and peptide‐like compounds (≥27%RA). Outflows were consistently more depleted in Δ 14 C‐DOC (i.e., older) compared to supraglacial streams (mean −195.2 and −61.3‰ respectively), perhaps due to integration of spatially heterogenous and variably aged DOM pools across the supraglacial environment, or incorporation of aged subglacial OM as runoff was routed to the outflow. Across Ecuador, Δ 14 C‐DOC enrichment was associated with decreased aromaticity of DOM, due to increased contributions of organic matter (OM) from microbial processes or atmospheric deposition of recently fixed and subsequently degraded OM (e.g., biomass burning byproducts). There was a regional gradient between glacier outflow DOM composition and streamwater δ 18 O, suggesting covariation between regional precipitation gradients and the DOM exported from glacier outflows. Ultimately, this highlights that atmospheric deposition may exert a control on glacier outflow DOM composition, suggesting regional air circulation patterns and precipitation sources in part determine the origins and quality of OM exported from glacier environments.

Soil formation in Mediterranean periglacial landscapes remains poorly understood as the interplay between erosion and aeolian dust accretion in providing parent materials, and mineral weathering and pedogenesis, as dominant post depositional processes, depends on a variety of local and regional factors. Herein, we investigate the balance between erosion and aeolian dust accretion in the formation of an alpine soil profile along an erosional gradient in the periglacial zone of Mount Olympus in Greece. We applied a wide range of analytical methods to 23 samples, from a soil profile developed in a glaciokarstic plateau, from colluvial sediment horizons interbedded in postglacial scree slopes and from modern Sahara dust samples deposited on the snowpack. Colluvial sediment horizons exhibit high concentrations of calcite rich sand and represent the local erosion products. The soil B horizon developed on a glaciokarstic plateau contains high amounts of fine earth and is rich in quartz, mica, plagioclase, clays, and Fe-Ti oxides. Based on its physical and textural characteristics the soil profile is partitioned in a surficial weathered Bw and a lower illuvial Bt horizon that overlies the local regolith composed of fragmented glacial till and slope wash deposits. Radiogenic isotope systematics, textural and mineralogical analysis show that the contribution of Sahara and locally sourced dust to the development of the soil B horizon ranges between 50 and 65%. Cryoturbation results in fine earth translocation from Bw to the Bt horizon, whereas weak pedogenetic modifications of aeolian and bedrock-derived minerals result in magnetic mineral weathering and secondary clay formation. Our findings reveal that, aeolian dust accretion is the dominant process in providing alpine soil parent material and that cryoturbation, weak pedogenesis, and clay mineral alteration occur within the Mediterranean periglacial zone of Mount Olympus.

<p> </p><p>Glacier-fed streams (GFSs) serve as headwaters to many of the world’s largest river networks. Although being characterized by extreme environmental conditions (i.e., low water temperatures, oligotrophy) GFSs host an underappreciated microbial biodiversity, especially within benthic biofilms which play pivotal roles in downstream biogeochemical cycles. Yet, we still lack a global overview of the GFS biofilm microbiome. In addition, little is known on how environmental conditions shape bacterial diversity, and how these relationships drive global distribution patterns. This is particularly important as mountain glaciers are currently vanishing at a rapid pace due to global warming. Here, we used 16S rRNA gene sequencing data from the Vanishing Glaciers project to conduct a first comprehensive analysis of the benthic microbiome from 148 GFSs across 11 mountain ranges. Our analyses revealed marked biogeographic patterns in the GFS microbiome, mainly driven by the replacement of phylogenetically closely related taxa. Strikingly, the GFS microbiome was characterized by pronounced level of endemism, with >58% of the Amplicon Sequence Variants (ASVs) being specific to one mountain range. Consistent with the marked dissimilarities across mountain ranges, we found a very small taxonomic core including only 200 ASVs, yet accounting for >25% of the total relative abundance of the ASVs. Finally, we found that spatial effects such as dispersal limitation, isolation and spatially autocorrelated environmental conditions overwhelmed the effect of the environment by itself on benthic biofilm beta diversity. Our findings shed light on the previously unresolved global diversity and biogeography of the GFS microbiome now at risk across the world’s major mountain ranges because of rapidly shrinking glaciers.</p>

Antimicrobial resistance is an omnipresent phenomenon in the anthropogenically influenced ecosystems. However, its role in shaping microbial community dynamics in pristine environments is relatively unknown.

The present study deals with the determination of Mount Olympus summit (Mytikas), exploiting modern observations such as Global Navigation Satellite Systems (GNSS) and existing geodetic information. The last official Olympus height determination goes back 102 years by the Swiss surveyor M. Kurz. Since then, only unofficial measurement campaigns have taken place. There is a variety of released heights, which range from 2917 to 2919 meters. In September and October of 2022, we conducted a GNSS campaign, focusing on the area of highest Olympus peaks. Employing precise GNSS measurements (accuracy of 1–3 cm), in conjunction with height information from Greece’s National Triangulation Network (5–10 cm accuracy) and an appropriate, recent geoid model (5–6 cm accuracy), we estimate through the so-called GNSS-levelling, the height of Olympus to 2917.727 m with respect to Greece’s officially accepted mean sea level and 2918.390 m with respect to the global vertical datum. Our estimation of Olympus highest peak shows remarkable consistency at the level of 12.8 cm to that of M. Kurz in 1921.

In glacier-fed streams, ecological windows of opportunity allow complex microbial biofilms to develop and transiently form the basis of the food web, thereby controlling key ecosystem processes. Using metagenome-assembled genomes, we unravel strategies that allow biofilms to seize this opportunity in an ecosystem otherwise characterized by harsh environmental conditions. We observe a diverse microbiome spanning the entire tree of life including a rich virome. Various co-existing energy acquisition pathways point to diverse niches and the exploitation of available resources, likely fostering the establishment of complex biofilms during windows of opportunity. The wide occurrence of rhodopsins, besides chlorophyll, highlights the role of solar energy capture in these biofilms while internal carbon and nutrient cycling between photoautotrophs and heterotrophs may help overcome constraints imposed by oligotrophy in these habitats. Mechanisms potentially protecting bacteria against low temperatures and high UV-radiation are also revealed and the selective pressure of this environment is further highlighted by a phylogenomic analysis differentiating important components of the glacier-fed stream microbiome from other ecosystems. Our findings reveal key genomic underpinnings of adaptive traits contributing to the success of complex biofilms to exploit environmental opportunities in glacier-fed streams, which are now rapidly changing owing to global warming.

The shrinking of glaciers is among the most iconic consequences of climate change. Despite this, the downstream consequences for ecosystem processes and related microbiome structure and function remain poorly understood. Here, using a space-for-time substitution approach across 101 glacier-fed streams (GFSs) from six major regions worldwide, we investigated how glacier shrinkage is likely to impact the organic matter (OM) decomposition rates of benthic biofilms. To do this, we measured the activities of five common extracellular enzymes and estimated decomposition rates by using enzyme allocation equations based on stoichiometry. We found decomposition rates to average 0.0129 (% d(-1)), and that decreases in glacier influence (estimated by percent glacier catchment coverage, turbidity, and a glacier index) accelerates decomposition rates. To explore mechanisms behind these relationships, we further compared decomposition rates with biofilm and stream water characteristics. We found that chlorophyll-a, temperature, and stream water N:P together explained 61% of the variability in decomposition. Algal biomass, which is also increasing with glacier shrinkage, showed a particularly strong relationship with decomposition, likely indicating their importance in contributing labile organic compounds to these carbon-poor habitats. We also found high relative abundances of chytrid fungi in GFS sediments, which putatively parasitize these algae, promoting decomposition through a fungal shunt. Exploring the biofilm microbiome, we then sought to identify bacterial phylogenetic clades significantly associated with decomposition, and found numerous positively (e.g., Saprospiraceae) and negatively (e.g., Nitrospira) related clades. Lastly, using metagenomics, we found evidence of different bacterial classes possessing different proportions of EEA-encoding genes, potentially informing some of the microbial associations with decomposition rates. Our results, therefore, present new mechanistic insights into OM decomposition in GFSs by demonstrating that an algal-based "green food web" is likely to increase in importance in the future and will promote important biogeochemical shifts in these streams as glaciers vanish.

Microbial life in glacier-fed streams (GFSs) is dominated by benthic biofilms which fulfill critical ecosystem processes. However, it remains unclear how the bacterial communities of these biofilms assemble in stream ecosystems characterized by rapid turnover of benthic habitats and high suspended sediment loads. Using16S rRNA gene amplicon sequence data collected from 54 GFSs across the Himalayas, European Alps, and Scandinavian Mountains, we found that benthic biofilms harbor bacterial communities that are distinct from the bacterial assemblages suspended in the streamwater. Our data showed a decrease in species richness in the benthic biofilms compared to the bacterial cells putatively free-living in the water. The benthic biofilms also differed from the suspended water fractions in terms of community composition. Differential abundance analyses highlighted bacterial families that were specific to the benthic biofilms and the suspended assemblages. Notably, source-sink models suggested that the benthic biofilm communities are not simply a subset of the suspended assemblages. Rather, we found evidence that deterministic processes (e.g., species sorting) shape the benthic biofilm communities. This is unexpected given the high vertical mixing of water and contained bacterial cells in GFSs and further highlights the benthic biofilm mode of life as one that is determined through niche-related processes. Our findings therefore reveal a "native" benthic biofilm community in an ecosystem that is currently threatened by climate-induced glacier shrinkage. IMPORTANCE Benthic biofilms represent the dominant form of life in glacier-fed streams. However, it remains unclear how bacterial communities within these biofilms assemble. Our findings from glacier-fed streams from three major mountain ranges across the Himalayas, the European Alps and the Scandinavian Mountains reveal a bacterial community associated with benthic biofilms that is distinct from the assemblage in the overlying streamwater. Our analyses suggest that selection is the underlying process to this differentiation. This is unexpected given that bacterial cells that are freely living or attached to the abundant sediment particles suspended in the water continuously mix with the benthic biofilms. The latter colonize loose sediments that are subject to high turnover owing to the forces of the water flow. Our research unravels the existence of a microbiome specific to benthic biofilms in glacier-fed streams, now under major threats due to global warming.

Between the southern margin of the European loess belt and Sahara Desert, thin and irregularly distributed loess deposits occur in Mediterranean mountains. During the most recent deglaciation, along the Pleistocene-Holocene boundary, the deposition of glacial, periglacial and outwash sediments, was the main local source of Mediterranean alpine loess, whereas proximal alluvial planes comprised a secondary source. The mid-Holocene termination of African Humid Period and subsequent aridification of Sahara Desert occurred simultaneously with a change of the regional climate from Atlantic to Mediterranean-dominated, characterized by frequent episodes of southerly winds. This resulted to a change of the loess source, as deflation of quartz rich silts enriched in Zr during intense episodes of Sahara dust transport became more dominant. Here, a 32cm loess profile from the Plateau of Muses (PM), below the summit of Mount Olympus, Greece, is investigated on the basis of grain size, mineralogy, environmental magnetism and geochemistry. Comparisons of loess samples with glacial and periglacial deposits, enables us to differentiate relative contributions of local sources and allochthonous aeolian inputs. Calcite sand rich in feldspars makes up the glacial and periglacial clast free matrix. In contrast, PM loess is composed by clay and fine silt fractions with minor calcite sand contributions. The mineralogical matrix of loess contains quartz, phyllosilicates and mixed layer clays, while its geochemical composition contains high amounts of detrital Fe-Ti oxides and aeolian transported Al and Zr. Based on the multi-proxy approach applied here, the loess profile is partitioned in three layers. Holocene average deposition rates (~2.5 cm/ka) broadly agree with modern Sahara dust deposition (~2.0 cm/ka) and long-term postglacial Mediterranean mountain denudation rates (~0.5 cm/ka). Such low rates provided ample time for post depositional modifications, such as decalcification, deferrification and removal of K, evident from the trends of chemical weathering proxies Ca/Sr, Fe/Ti and K/Rb, respectively.

Glacier-fed streams (GFSs) are extreme and rapidly vanishing ecosystems, and yet they harbor diverse microbial communities. Although our understanding of the GFS microbiome has recently increased, we do not know which microbial clades are ecologically successful in these ecosystems, nor do we understand potentially underlying mechanisms. Ecologically successful clades should be more prevalent across GFSs compared to other clades, which should be reflected as clade-wise distinctly low phylogenetic turnover. However, methods to assess such patterns are currently missing. Here we developed and applied a novel analytical framework, “phyloscore analysis”, to identify clades with lower spatial phylogenetic turnover than other clades in the sediment microbiome across twenty GFSs in New Zealand. These clades constituted up to 44% and 64% of community α-diversity and abundance, respectively. Furthermore, both their α-diversity and abundance increased as sediment chlorophyll a decreased, corroborating their ecological success in GFS habitats largely devoid of primary production. These clades also contained elevated levels of putative microdiversity than others, which could potentially explain their high prevalence in GFSs. This hitherto unknown microdiversity may be threatened as glaciers shrink, urging towards further genomic and functional exploration of the GFS microbiome.