Diurnal temperature range as a key predictor of plants' elevation ranges globally.

A prominent hypothesis in ecology is that larger species ranges are found in more variable climates because species develop broader environmental tolerances, predicting a positive range size-temperature variability relationship. However, this overlooks the extreme temperatures that variable climates impose on species, with upper or lower thermal limits more likely to be exceeded. Accordingly, we propose the 'temperature range squeeze' hypothesis, predicting a negative range size-temperature variability relationship. We test these contrasting predictions by relating 88,000 elevation range sizes of vascular plants in 44 mountains to short- and long-term temperature variation. Consistent with our hypothesis, we find that species' range size is negatively correlated with diurnal temperature range. Accurate predictions of short-term temperature variation will become increasingly important for extinction risk assessment in the future.

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Life-history traits predict responses of wild bees to climate variation.

Life-history traits, which are physical traits or behaviours that affect growth, survivorship and reproduction, could play an important role in how well organisms respond to environmental change. By looking for trait-based responses within groups, we can gain a mechanistic understanding of why environmental change might favour or penalize certain species over others. We monitored the abundance of at least 154 bee species for 8 consecutive years in a subalpine region of the Rocky Mountains to ask whether bees respond differently to changes in abiotic conditions based on their life-history traits. We found that comb-building cavity nesters and larger bodied bees declined in relative abundance with increasing temperatures, while smaller, soil-nesting bees increased. Further, bees with narrower diet breadths increased in relative abundance with decreased rainfall. Finally, reduced snowpack was associated with reduced relative abundance of bees that overwintered as prepupae whereas bees that overwintered as adults increased in relative abundance, suggesting that overwintering conditions might affect body size, lipid content and overwintering survival. Taken together, our results show how climate change may reshape bee pollinator communities, with bees with certain traits increasing in abundance and others declining, potentially leading to novel plant-pollinator interactions and changes in plant reproduction.

Grass species identity shapes communities of root and leaf fungi more than elevation.

Fungal symbionts can buffer plants from environmental extremes and may affect host capacities to acclimate, adapt, or redistribute under environmental change; however, the distributions of fungal symbionts along abiotic gradients are poorly described. Fungal mutualists should be the most beneficial in abiotically stressful environments, and the structure of networks of plant-fungal interactions likely shift along gradients, even when fungal community composition does not track environmental stress. We sampled 634 unique combinations of fungal endophytes and mycorrhizal fungi, grass species identities, and sampling locations from 66 sites across six replicate altitudinal gradients in the western Colorado Rocky Mountains. The diversity and composition of leaf endophytic, root endophytic, and arbuscular mycorrhizal (AM) fungal guilds and the overall abundance of fungal functional groups (pathogens, saprotrophs, mutualists) tracked grass host identity more closely than elevation. Network structures of root endophytes become more nested and less specialized at higher elevations, but network structures of other fungal guilds did not vary with elevation. Overall, grass species identity had overriding influence on the diversity and composition of above- and belowground fungal endophytes and AM fungi, despite large environmental variation. Therefore, in our system climate change may rarely directly affect fungal symbionts. Instead, fungal symbiont distributions will most likely track the range dynamics of host grasses.

Mammalian herbivores restrict the altitudinal range limits of alpine plants.

Although rarely experimentally tested, biotic interactions have long been hypothesised to limit low-elevation range boundaries of species. We tested the effects of herbivory on three alpine-restricted plant species by transplanting plants below (novel), at the edge (limit), or in the centre (core) of their current elevational range and factorially fencing-out above- and belowground mammals. Herbivore damage was greater in range limit and novel habitats than in range cores. Exclosures increased plant biomass and reproduction more in novel habitats than in range cores, suggesting demographic costs of novel interactions with herbivores. We then used demographic models to project population growth rates, which increased 5-20% more under herbivore exclosure at range limit and novel sites than in core habitats. Our results identify mammalian herbivores as key drivers of the low-elevation range limits of alpine plants and indicate that upward encroachment of herbivores could trigger local extinctions by depressing plant population growth.

Macroecological context predicts species' responses to climate warming.

Context-dependencies in species' responses to the same climate change frustrate attempts to generalize and make predictions based on experimental and observational approaches in biodiversity science. Here, we propose predictability may be enhanced by explicitly incorporating macroecological context into analyses of species' responses to climate manipulations. We combined vascular plant species' responses to an 8-year, 12-site turf transplant climate change experiment set in southwestern Norway with climate niche data from the observed 151 species. We used the difference between a species' mean climate across their range and climate conditions at the transplant site ("climate differences") to predict colonization probability, extinction probability, and change in abundance of a species at a site. In analyses across species that ignore species-specific patterns, colonization success increased as species' distribution optima were increasingly warmer than the experimental target site. Extinction probability increased as species' distribution optima were increasingly colder than the target site. These patterns were reflected in change in abundance analyses. We found weak responses to increased precipitation in these oceanic climates. Climate differences were better predictors of species' responses to climate manipulations than range size. Interestingly, similar patterns were found when analyses focused on variation in species-specific responses across sites. These results provide an experimental underpinning to observational studies that report thermophilization of communities and suggest that space-for-time substitutions may be valid for predicting species' responses to climate warming, given other conditions are accounted for (e.g., soil nutrients). Finally, we suggest that this method of putting climate change experiments into macroecological context has the potential to generalize and predict species' responses to climate manipulations globally.

Weak latitudinal gradients in insect herbivory for dominant rangeland grasses of North America.

Patterns of insect herbivory may follow predictable geographical gradients, with greater herbivory at low latitudes. However, biogeographic studies of insect herbivory often do not account for multiple abiotic factors (e.g., precipitation and soil nutrients) that could underlie gradients. We tested for latitudinal clines in insect herbivory as well as climatic, edaphic, and trait-based drivers of herbivory. We quantified herbivory on five dominant grass species over 23 sites across the Great Plains, USA. We examined the importance of climate, edaphic factors, and traits as correlates of herbivory. Herbivory increased at low latitudes when all grass species were analyzed together and for two grass species individually, while two other grasses trended in this direction. Higher precipitation was related to more herbivory for two species but less herbivory for a different species, while higher specific root length was related to more herbivory for one species and less herbivory for a different species. Taken together, results highlight that climate and trait-based correlates of herbivory can be highly contextual and species-specific. Patterns of insect herbivory on dominant grasses support the hypothesis that herbivory increases toward lower latitudes, though weakly, and indicates that climate change may have species-specific effects on plant-herbivore interactions.

Correction to: Soil microbes that may accompany climate warming increase alpine plant production.

The DOI link to the data in the Acknowledgments section of the article was incorrect. The proper link to the data is.

Soil microbes that may accompany climate warming increase alpine plant production.

Climate change is causing species with non-overlapping ranges to come in contact, and a key challenge is to predict the consequences of such species re-shuffling. Experiments on plants have focused largely on novel competitive interactions; other species interactions, such as plant-microbe symbioses, while less studied, may also influence plant responses to climate change. In this greenhouse study, we evaluated interactions between soil microbes and alpine-restricted plant species, simulating a warming scenario in which low-elevation microbes migrate upslope into the distribution of alpine plants. We examined three alpine grasses from the Rocky Mountains, CO, USA (Poa alpina, Festuca brachyphylla, and Elymus scribneri). We used soil inocula from within (resident) or below (novel) the plants' current elevation range and examined responses in plant biomass, plant traits, and fungal colonization of roots. Resident soil inocula from the species' home range decreased biomass to a greater extent than novel soil inocula. The depressed growth in resident soils suggested that these soils harbor more carbon-demanding microbes, as plant biomass generally declined with greater fungal colonization of roots, especially in resident soil inocula. Although plant traits did not respond to the provenance of soil inocula, specific leaf area declined and root:shoot ratio increased when soil inocula were sterilized, indicating microbial mediation of plant trait expression. Contrary to current predictions, our findings suggest that if upwardly migrating microbes were to displace current soil microbes, alpine plants may benefit from this warming-induced microbial re-shuffling.

Altitudinal gradients fail to predict fungal symbiont responses to warming.

Climate change is shifting altitudinal species ranges, with potential to disrupt species interactions. Altitudinal gradient studies and warming experiments can both increase understanding of climate effects on species interactions, but few studies have used both together to improve predictions. We examined whether plant-fungal symbioses responded similarly to altitude and 23 yr of experimental warming. Root- and leaf-associated fungi, which can mediate plants' climate sensitivity, responded divergently to elevation vs. warming. Fungal colonization, diversity, and composition varied with altitude, but climate variables were generally weak predictors; other factors such as host plant identity, plant community composition, or edaphic variables likely drive fungal altitudinal distributions. Manipulated warming altered fungal colonization, but not composition or diversity. Leaf symbionts were more sensitive to climate and experimental warming than root symbionts. Altitudinal patterns and responses to warming differed among host plant species and fungal groups, indicating that predicting climate effects on symbioses will require tracking both host and symbiont identities. Combining experimental and observational methods can yield valuable insight into how climate change may alter plant-symbiont interactions, but our results indicate that altitude does not always serve as an adequate proxy for warming effects on fungal symbionts of plants.

Plant Identity Influences Foliar Fungal Symbionts More Than Elevation in the Colorado Rocky Mountains.

Despite colonizing nearly every plant on Earth, foliar fungal symbionts have received little attention in studies on the biogeography of host-associated microbes. Evidence from regional scale studies suggests that foliar fungal symbiont distributions are influenced both by plant hosts and environmental variation in climate and soil resources. However, previous surveys have focused on either one plant host across an environmental gradient or one gradient and multiple plant hosts, making it difficult to disentangle the influence of host identity from the influence of the environment on foliar endophyte communities. We used a culture-based approach to survey fungal symbiont composition in the leaves of nine C3 grass species along replicated elevation gradients in grasslands of the Colorado Rocky Mountains. In these ecosystems, the taxonomic richness and composition of foliar fungal symbionts were mostly structured by the taxonomic identity of the plant host rather than by variation in climate. Plant traits related to size (height and leaf length) were the best predictors of foliar fungal symbiont composition and diversity, and composition did not vary predictably with plant evolutionary history. The largest plants had the most diverse and distinctive fungal communities. These results suggest that across the ~ 300 m elevation range that we sampled, foliar fungal symbionts may indirectly experience climate change by tracking the shifting distributions of plant hosts rather than tracking climate directly.