Phenology and species determine growing-season albedo increase at the altitudinal limit of shrub growth in the sub-Arctic.

Arctic warming is resulting in reduced snow cover and increased shrub growth, both of which have been associated with altered land surface-atmospheric feedback processes involving sensible heat flux, ground heat flux and biogeochemical cycling. Using field measurements, we show that two common Arctic shrub species (Betula glandulosa and Salix pulchra), which are largely responsible for shrub encroachment in tundra, differed markedly in albedo and that albedo of both species increased as growing season progressed when measured at their altitudinal limit. A moveable apparatus was used to repeatedly measure albedo at six precise spots during the summer of 2012, and resampled in 2013. Contrary to the generally accepted view of shrub-covered areas having low albedo in tundra, full-canopy prostrate B. glandulosa had almost the highest albedo of all surfaces measured during the peak of the growing season. The higher midsummer albedo is also evident in localized MODIS albedo aggregated from 2000 to 2013, which displays a similar increase in growing-season albedo. Using our field measurements, we show the ensemble summer increase in tundra albedo counteracts the generalized effect of earlier spring snow melt on surface energy balance by approximately 40%. This summer increase in albedo, when viewed in absolute values, is as large as the difference between the forest and tundra transition. These results indicate that near future (<50 years) changes in growing-season albedo related to Arctic vegetation change are unlikely to be particularly large and might constitute a negative feedback to climate warming in certain circumstances. Future efforts to calculate energy budgets and a sensible heating feedback in the Arctic will require more detailed information about the relative abundance of different ground cover types, particularly shrub species and their respective growth forms and phenology.

Herbivore diversity effects on Arctic tundra ecosystems: a systematic review.

BACKGROUND: Northern ecosystems are strongly influenced by herbivores that differ in their impacts on the ecosystem. Yet the role of herbivore diversity in shaping the structure and functioning of tundra ecosystems has been overlooked. With climate and land-use changes causing rapid shifts in Arctic species assemblages, a better understanding of the consequences of herbivore diversity changes for tundra ecosystem functioning is urgently needed. This systematic review synthesizes available evidence on the effects of herbivore diversity on different processes, functions, and properties of tundra ecosystems. METHODS: Following a published protocol, our systematic review combined primary field studies retrieved from bibliographic databases, search engines and specialist websites that compared tundra ecosystem responses to different levels of vertebrate and invertebrate herbivore diversity. We used the number of functional groups of herbivores (i.e., functional group richness) as a measure of the diversity of the herbivore assemblage. We screened titles, abstracts, and full texts of studies using pre-defined eligibility criteria. We critically appraised the validity of the studies, tested the influence of different moderators, and conducted sensitivity analyses. Quantitative synthesis (i.e., calculation of effect sizes) was performed for ecosystem responses reported by at least five articles and meta-regressions including the effects of potential modifiers for those reported by at least 10 articles. REVIEW FINDINGS: The literature searches retrieved 5944 articles. After screening titles, abstracts, and full texts, 201 articles including 3713 studies (i.e., individual comparisons) were deemed relevant for the systematic review, with 2844 of these studies included in quantitative syntheses. The available evidence base on the effects of herbivore diversity on tundra ecosystems is concentrated around well-established research locations and focuses mainly on the impacts of vertebrate herbivores on vegetation. Overall, greater herbivore diversity led to increased abundance of feeding marks by herbivores and soil temperature, and to reduced total abundance of plants, graminoids, forbs, and litter, plant leaf size, plant height, and moss depth, but the effects of herbivore diversity were difficult to tease apart from those of excluding vertebrate herbivores. The effects of different functional groups of herbivores on graminoid and lichen abundance compensated each other, leading to no net effects when herbivore effects were combined. In turn, smaller herbivores and large-bodied herbivores only reduced plant height when occurring together but not when occurring separately. Greater herbivore diversity increased plant diversity in graminoid tundra but not in other habitat types. CONCLUSIONS: This systematic review underscores the importance of herbivore diversity in shaping the structure and function of Arctic ecosystems, with different functional groups of herbivores exerting additive or compensatory effects that can be modulated by environmental conditions. Still, many challenges remain to fully understand the complex impacts of herbivore diversity on tundra ecosystems. Future studies should explicitly address the role of herbivore diversity beyond presence-absence, targeting a broader range of ecosystem responses and explicitly including invertebrate herbivores. A better understanding of the role of herbivore diversity will enhance our ability to predict whether and where shifts in herbivore assemblages might mitigate or further amplify the impacts of environmental change on Arctic ecosystems.

After the frass: foraging pikas select patches previously grazed by caterpillars.

Interactions among herbivores can shape the structure of their communities and drive their dynamics. However, detecting herbivore interactions can be challenging when they are deferred in space or time. Moreover, interactions among distantly related groups of herbivores, such as vertebrates and invertebrates, are poorly understood. We investigated the effect of invertebrate herbivory on the subsequent foraging choices of a small alpine-dwelling vertebrate, the collared pika (Ochotona collaris). We carried out a field experiment within pika territories, by presenting them with a choice of foraging sites following manipulation of invertebrate (caterpillar) herbivory. Pikas actively selected areas with increased, recent invertebrate herbivory. While the underlying mechanisms behind this interaction remain unknown, our results demonstrate a positive effect of invertebrate herbivores on subsequent vertebrate foraging preferences for the first time. Even among distantly related taxa, such interactions where one herbivore is cueing on the foraging of another, could drive the creation of herbivory hotspots, with cascading consequences for ecosystem processes.

Global assessment of experimental climate warming on tundra vegetation: heterogeneity over space and time.

Understanding the sensitivity of tundra vegetation to climate warming is critical to forecasting future biodiversity and vegetation feedbacks to climate. In situ warming experiments accelerate climate change on a small scale to forecast responses of local plant communities. Limitations of this approach include the apparent site-specificity of results and uncertainty about the power of short-term studies to anticipate longer term change. We address these issues with a synthesis of 61 experimental warming studies, of up to 20 years duration, in tundra sites worldwide. The response of plant groups to warming often differed with ambient summer temperature, soil moisture and experimental duration. Shrubs increased with warming only where ambient temperature was high, whereas graminoids increased primarily in the coldest study sites. Linear increases in effect size over time were frequently observed. There was little indication of saturating or accelerating effects, as would be predicted if negative or positive vegetation feedbacks were common. These results indicate that tundra vegetation exhibits strong regional variation in response to warming, and that in vulnerable regions, cumulative effects of long-term warming on tundra vegetation - and associated ecosystem consequences - have the potential to be much greater than we have observed to date.

Uniform female-biased sex ratios in alpine willows.

PREMISE OF THE STUDY: The development of biased sex ratios in dioecious plant species has been ascribed to either (1) factors influencing differential adult mortality of male and female plants or (2) factors acting at an early life stage that determine seed sex ratio or seedling survival. METHODS: To discriminate between these two competing hypotheses, we surveyed sex and age of 379 individuals from five species of the genus Salix across 11 alpine valleys in the southwest Yukon. KEY RESULTS: We observed uniformly female-biased sex ratios of approximately 2:1 across all adult age cohorts and patch sizes of the five willow species. No spatial variation in sex ratio occurred that could be associated with site-specific characteristics such as elevation or aspect. CONCLUSIONS: Our results indicate that the female-biased sex ratios in the alpine willow species investigated in this study are not a consequence of ecological processes acting on established adult plants. The sex ratio is instead determined at an early life stage by a mechanism that remains unknown.

What are the effects of herbivore diversity on tundra ecosystems? A systematic review protocol.

BACKGROUND: Changes in the diversity of herbivore communities can strongly influence the functioning of northern ecosystems. Different herbivores have different impacts on ecosystems because of differences in their diets, behaviour and energy requirements. The combined effects of different herbivores can in some cases compensate each other but lead to stronger directional changes elsewhere. However, the diversity of herbivore assemblages has until recently been a largely overlooked dimension of plant-herbivore interactions. Given the ongoing environmental changes in tundra ecosystems, with increased influx of boreal species and changes in the distribution and abundance of arctic herbivores, a better understanding of the consequences of changes in the diversity of herbivore assemblages is needed. This protocol presents the methodology that will be used in a systematic review on the effects of herbivore diversity on different processes, functions and properties of tundra ecosystems. METHODS: This systematic review builds on an earlier systematic map on herbivory studies in the Arctic that identified a relatively large number of studies assessing the effects of multiple herbivores. The systematic review will include primary field studies retrieved from databases, search engines and specialist websites, that compare responses of tundra ecosystems to different levels of herbivore diversity, including both vertebrate and invertebrate herbivores. We will use species richness of herbivores or the richness of functional groups of herbivores as a measure of the diversity of the herbivore assemblages. Studies will be screened in three stages: title, abstract and full text, and inclusion will follow clearly identified eligibility criteria, based on their target population, exposure, comparator and study design. The review will cover terrestrial Arctic ecosystems including the forest-tundra ecotone. Potential outcomes will include multiple processes, functions and properties of tundra ecosystems related to primary productivity, nutrient cycling, accumulation and dynamics of nutrient pools, as well as the impacts of herbivores on other organisms. Studies will be critically appraised for validity, and where studies report similar outcomes, meta-analysis will be performed.

Four decades of plant community change in the Alpine tundra of southwest Yukon, Canada.

Repeat measurements from long-term plots provide precise data for studying plant community change. In 2010, we visited a remote location in Yukon, Canada, where a detailed survey of alpine tundra communities was conducted in 1968. Plant community composition was resurveyed on the same four slopes using the same methods as the original study. Species richness and diversity increased significantly over the 42 years and non-metric multidimensional scaling indicated that community composition had also changed significantly. However, the direction and magnitude of change varied with aspect. Dominant species were not replaced or eliminated but, instead, declined in relative importance. Fine-scale changes in vegetation were evident from repeat photography and dendro-ecological analysis of erect shrubs, supporting the community-level analysis. The period of study corresponds to a mean annual temperature increase of 2 degrees C, suggesting that climate warming has influenced these changes.

Expansion of canopy-forming willows over the twentieth century on Herschel Island, Yukon Territory, Canada.

Canopy-forming shrubs are reported to be increasing at sites around the circumpolar Arctic. Our results indicate expansion in canopy cover and height of willows on Herschel Island located at 70 degrees north on the western Arctic coast of the Yukon Territory. We examined historic photographs, repeated vegetation surveys, and conducted monitoring of long-term plots and found evidence of increases of each of the dominant canopy-forming willow species (Salix richardsonii, Salix glauca and Salix pulchra), during the twentieth century. A simple model of patch initiation indicates that the majority of willow patches for each of these species became established between 1910 and 1960, with stem ages and maximum growth rates indicating that some patches could have established as late as the 1980s. Collectively, these results suggest that willow species are increasing in canopy cover and height on Herschel Island. We did not find evidence that expansion of willow patches is currently limited by herbivory, disease, or growing conditions.

Multi-decadal changes in tundra environments and ecosystems: synthesis of the International Polar Year-Back to the Future project (IPY-BTF).

Understanding the responses of tundra systems to global change has global implications. Most tundra regions lack sustained environmental monitoring and one of the only ways to document multi-decadal change is to resample historic research sites. The International Polar Year (IPY) provided a unique opportunity for such research through the Back to the Future (BTF) project (IPY project #512). This article synthesizes the results from 13 papers within this Ambio Special Issue. Abiotic changes include glacial recession in the Altai Mountains, Russia; increased snow depth and hardness, permafrost warming, and increased growing season length in sub-arctic Sweden; drying of ponds in Greenland; increased nutrient availability in Alaskan tundra ponds, and warming at most locations studied. Biotic changes ranged from relatively minor plant community change at two sites in Greenland to moderate change in the Yukon, and to dramatic increases in shrub and tree density on Herschel Island, and in subarctic Sweden. The population of geese tripled at one site in northeast Greenland where biomass in non-grazed plots doubled. A model parameterized using results from a BTF study forecasts substantial declines in all snowbeds and increases in shrub tundra on Niwot Ridge, Colorado over the next century. In general, results support and provide improved capacities for validating experimental manipulation, remote sensing, and modeling studies.

Plant interactions are unimportant in a subarctic-alpine plant community.

We investigated whether plant interaction intensity in a subarctic-alpine meadow is important for determining community structure and species abundance. Using two common species as phytometers, we measured interaction intensity using a neighbor removal approach. Eight biotic and abiotic variables known to influence species abundance and community structure were measured, with regression trees used to examine how plant interactions and the biotic and abiotic variables were related to species evenness, richness, and phytometer spatial cover. A range of interactions was present, with both strong competition and facilitation present over small-scale abiotic and biotic gradients. Despite the variation in interaction intensity, it was generally unrelated to either community structure or phytometer cover. In other words, plant interactions were intense in many cases but were not important to community structure. This may be due to the prevalence of clonal species in this system and the influence of previous year's interactions on plant survival and patterns of community structure. These results also suggest how conflicting theories of the role of competition in unproductive environments may be resolved. Our findings suggest that plant interactions may be intense in reducing individual growth, while simultaneously not important in the context of community structure. Plant interactions need to be viewed and tested relative to other factors and stresses to accurately evaluate their importance in plant communities, with continued differentiation between the intensity of plant interactions and their relative importance in communities.

Spring warming in Yukon mountains is not amplified by the snow albedo feedback.

Decreasing spring snow cover may amplify Arctic warming through the snow albedo feedback. To examine the impact of snowmelt on increasing temperature we used a 5,000 m elevation gradient in Yukon, Canada, extending from valley-bottom conifer forests, through middle elevation tundra, to high elevation icefields, to compare validated downscaled reanalysis air temperature patterns across elevational bands characterized by different patterns of spring snowmelt. From 2000 to 2014 we observed surface warming of 0.01 °C/a·1,000 m in May (0.14 °C/a at 1,000 m to 0.19 °C/a at 5,000 m), and uniform cooling of 0.09 °C/a in June at all elevations. May temperature trends across elevationally dependent land cover types were highly correlated with each other despite large variations in albedo and snow cover trends. Furthermore, a clear dependency of infrared skin temperature on snow cover mediated albedo decline was observed in tundra, but this was insufficient to influence average diurnal air temperature. We observed negative June temperature trends which we attribute to increasing daytime cloud cover because albedo and snow cover trends were unchanging. We conclude that 8-day and monthly averaged Spring air temperature trends are responding to a synoptic external forcing that is much stronger than the snow albedo feedback in sub-Arctic mountains.

Global negative vegetation feedback to climate warming responses of leaf litter decomposition rates in cold biomes.

Whether climate change will turn cold biomes from large long-term carbon sinks into sources is hotly debated because of the great potential for ecosystem-mediated feedbacks to global climate. Critical are the direction, magnitude and generality of climate responses of plant litter decomposition. Here, we present the first quantitative analysis of the major climate-change-related drivers of litter decomposition rates in cold northern biomes worldwide. Leaf litters collected from the predominant species in 33 global change manipulation experiments in circum-arctic-alpine ecosystems were incubated simultaneously in two contrasting arctic life zones. We demonstrate that longer-term, large-scale changes to leaf litter decomposition will be driven primarily by both direct warming effects and concomitant shifts in plant growth form composition, with a much smaller role for changes in litter quality within species. Specifically, the ongoing warming-induced expansion of shrubs with recalcitrant leaf litter across cold biomes would constitute a negative feedback to global warming. Depending on the strength of other (previously reported) positive feedbacks of shrub expansion on soil carbon turnover, this may partly counteract direct warming enhancement of litter decomposition.

Herbivory mediates grass-endophyte relationships.

Endophytic fungi are plant symbionts living asymptomatically within plant tissues. Neotyphodium spp., which are asexual vertically transmitted systemic fungal endophytes of cool-season grasses, are predicted to be plant mutualists. These endophytes increase host plant resistance to environmental stresses and/or increase the production of alkaloid-based herbivore deterrents. The ubiquity of this defense mutualism is unclear, and a variety of alternative mechanisms may explain the observed variation in infection rates, levels of deterrence, and the maintenance of asexual endophytes in grass populations. We found that grass-endophyte interactions are variable and ordered along an herbivory gradient in an undisturbed subarctic alpine ecosystem. Native grass populations in grazed sites had significantly greater frequency of Neotyphodium infection compared to ungrazed sites. Tillers from grazed sites had significantly higher hyphal densities compared to ungrazed sites. The ability of grass-Neotyphodium constituents to deter vertebrate herbivory in natural systems is thought to be rare. In grazed meadows, we showed that endophyte infection resulted in the deterrence of grazing by native vertebrate herbivores. However, the same herbivores did not distinguish between infected and uninfected grass harvested from ungrazed areas. These results demonstrate that the relationship between vertically transmitted endophytes and grasses in the alpine tundra vary greatly within populations. This may be based in part on defense mutualism and is consistent, under varying levels of herbivory, with the predictions of optimal defense theory.

Higher predation risk for insect prey at low latitudes and elevations.

Biotic interactions underlie ecosystem structure and function, but predicting interaction outcomes is difficult. We tested the hypothesis that biotic interaction strength increases toward the equator, using a global experiment with model caterpillars to measure predation risk. Across an 11,660-kilometer latitudinal gradient spanning six continents, we found increasing predation toward the equator, with a parallel pattern of increasing predation toward lower elevations. Patterns across both latitude and elevation were driven by arthropod predators, with no systematic trend in attack rates by birds or mammals. These matching gradients at global and regional scales suggest consistent drivers of biotic interaction strength, a finding that needs to be integrated into general theories of herbivory, community organization, and life-history evolution.

Moss Mediates the Influence of Shrub Species on Soil Properties and Processes in Alpine Tundra.

In tundra ecosystems, bryophytes influence soil processes directly and indirectly through interactions with overstory shrub species. We experimentally manipulated moss cover and measured seasonal soil properties and processes under two species of deciduous shrubs with contrasting canopy structures, Salix planifolia pulchra and Betula glandulosa-nana complex. Soil properties (seasonal temperature, moisture and C:N ratios) and processes (seasonal litter decomposition and soil respiration) were measured over twelve months. Shrub species identity had the largest influence on summer soil temperatures and soil respiration rates, which were higher under Salix canopies. Mosses were associated with lower soil moisture irrespective of shrub identity, but modulated the effects of shrubs on winter soil temperatures and soil C:N ratios so that moss cover reduced differences in soil winter temperatures between shrub species and reduced C:N ratios under Betula but not under Salix canopies. Our results suggest a central role of mosses in mediating soil properties and processes, with their influence depending on shrub species identity. Such species-dependent effects need to be accounted for when forecasting vegetation dynamics under ongoing environmental changes.

Ecological, evolutionary and social constraints on reproductive effort: are hoary marmots really biennial breeders?

Biennial breeding is a rare life-history trait observed in animal species living in harsh, unproductive environments. This reproductive pattern is thought to occur in 10 of 14 species in the genus Marmota, making marmots useful model organisms for studying its ecological and evolutionary implications. Biennial breeding in marmots has been described as an obligate pattern which evolved as a mechanism to mitigate the energetic costs of reproduction (Evolved Constraint hypothesis). However, recent anecdotal evidence suggests that it is a facultative pattern controlled by annual variation in climate and food availability (Environmental Constraint hypothesis). Finally, in social animals like marmots, biennial breeding could result from reproductive competition between females within social groups (Social Constraint hypothesis). We evaluated these three hypotheses using mark-recapture data from an 8-year study of hoary marmot (Marmota caligata) population dynamics in the Yukon. Annual variation in breeding probability was modeled using multi-state mark-recapture models, while other reproductive life-history traits were modeled with generalized linear mixed models. Hoary marmots were neither obligate nor facultative biennial breeders, and breeding probability was insensitive to evolved, environmental, or social factors. However, newly mature females were significantly less likely to breed than older individuals. Annual breeding did not result in increased mortality. Female survival and, to a lesser extent, average fecundity were correlated with winter climate, as indexed by the Pacific Decadal Oscillation. Hoary marmots are less conservative breeders than previously believed, and the evidence for biennial breeding throughout Marmota, and in other arctic/alpine/antarctic animals, should be re-examined. Prediction of future population dynamics requires an accurate understanding of life history strategies, and of how life history traits allow animals to cope with changes in weather and other demographic influences.

Is dimethylsulfoxide a reliable solvent for extracting chlorophyll under field conditions?

Dimethylsulfoxide (DMSO) appears to be a reliable solvent for extracting chlorophyll (Chl), however, modification of standard methods may be necessary for some species under field conditions. We found that Chl extraction of whole leaf tissue with DMSO incubated at between 25 and 40 degrees C was generally similar to the 80% acetone method, except for one graminoid species that required maceration. There was little effect of incubation temperature or duration of incubation beyond 7 h on extraction efficiency, but DMSO extracts were less stable than acetone extracts during one week of cold storage, especially if they thawed during this period. Since Chl extraction methods may provide variable results, particularly in the field, studies using different solvents should be compared cautiously unless specific methods have been calibrated.

Shrub canopies influence soil temperatures but not nutrient dynamics: An experimental test of tundra snow-shrub interactions.

Shrubs are the largest plant life form in tundra ecosystems; therefore, any changes in the abundance of shrubs will feedback to influence biodiversity, ecosystem function, and climate. The snow-shrub hypothesis asserts that shrub canopies trap snow and insulate soils in winter, increasing the rates of nutrient cycling to create a positive feedback to shrub expansion. However, previous work has not been able to separate the abiotic from the biotic influences of shrub canopies. We conducted a 3-year factorial experiment to determine the influences of canopies on soil temperatures and nutrient cycling parameters by removing ∼0.5 m high willow (Salix spp.) and birch (Betula glandulosa) shrubs, creating artificial shrub canopies and comparing these manipulations to nearby open tundra and shrub patches. Soil temperatures were 4-5°C warmer in January, and 2°C cooler in July under shrub cover. Natural shrub plots had 14-33 cm more snow in January than adjacent open tundra plots. Snow cover and soil temperatures were similar in the manipulated plots when compared with the respective unmanipulated treatments, indicating that shrub canopy cover was a dominant factor influencing the soil thermal regime. Conversely, we found no strong evidence of increased soil decomposition, CO2 fluxes, or nitrate or ammonia adsorbtion under artificial shrub canopy treatments when compared with unmanipulated open tundra. Our results suggest that the abiotic influences of shrub canopy cover alone on nutrient dynamics are weaker than previously asserted.

Influence of food hoarding behavior on the over-winter survival of pikas in strongly seasonal environments.

Food hoarding is a behavioral adaptation of some herbivores to manage food availability through time and space. In strongly seasonal environments, where summer growing seasons are short relative to winter, an earlier start to hoarding should increase the amount of vegetation stored for winter and improve subsequent survival. We examined hoarding behavior ('haying') and its impact on survival for a small alpine lagomorph, the collared pika (Ochotona collaris) in Yukon, Canada. We used a combination of video surveillance, haypile measurements, and survival data from marked individuals of known age and sex. Annual haypile initiation was strongly influenced by age and year. Adult pikas began haying an average of 16 days earlier in 2004 relative to 2005, whereas young of the year (juveniles) did not vary in the timing of haypile initiation. The mean haying rate per hour increased monthly from 3.7 +/- 0.7 trips in June to 6.6 +/- 0.8 trips in August. Simulation analysis estimated the median haypile mass (dry weight) by mid-September to be 5.5 kg. At least 75% of simulated haypiles had a minimum of 90 days (3 months) of food reserves, and 50% of simulated haypiles had a minimum of 177 days (5.9 months) of food reserves by mid-September, supporting the hypothesis that haypiles serve as the primary source of food during winter. Survival was greatest for pikas in 2005 when they began haying prior to 31 July, but the benefits of earlier accumulation of vegetation on survival also varied between years. The implications of earlier spring snowmelt are discussed with respect to pika foraging and overwinter survival.