Forage quality in tundra grasslands under herbivory: Silicon-based defences, nutrients and their ratios in grasses

1. Herbivore-induced changes in both leaf silicon-based defence and nutrient levels are potential mechanisms through which grazers alter the quality of their own grass supply. In tundra grasslands, herbivores have been shown to increase nutrient contents of grasses; yet, it is an open question whether they also increase grass silicon-based defence levels. Here, we asked if, and to what extent, herbivores affect silicon content and silicon:nutrient ratios of grasses found in tundra grasslands. 2. We performed an herbivore-interaction field-experiment spanning four tundragrassland sites. At each site, we established reindeer-open and reindeer-exclusion plots in tundra-patches that had been disturbed or not by small rodents during the previous winter, for a total of 96 plots. We randomly collected over 1,150 leaf samples of inherently silicon-rich and silicon-poor grass species throughout a growing season and analysed silicon, nitrogen and phosphorus contents of each leaf. 3. Small-rodent winter disturbance did not affect grass silicon content, but increased grass quality (i.e. lowered silicon:nutrient ratios) by enhancing nutrient levels of both silicon-rich (+20%–22%) and silicon-poor (+26%–34%) grasses. Reindeer summer herbivory increased the quality of silicon-rich grasses by decreasing their silicon content (−7%). However, the two herbivores together offset both these quality increments in silicon-rich grasses, thus reducing their quality towards the level of those found in the absence of herbivores and further enhancing their silicon:nutrient ratios (+13%–22%) relative to silicon-poor grasses. 4. Synthesis. We provide the first community-level, field-based assessment of how herbivory-driven changes in both leaf silicon-based defence and nutrient levels alter grass-forage quality in tundra grasslands. Herbivores did not promote a net silicon accumulation in grasses, but rather enhanced their overall quality. Yet, the magnitude of these quality increments varied depending on the herbivore(s

Intraspecific trait variability is a key feature underlying high Arctic plant community resistance to climate warming

In the high Arctic, plant community species composition generally responds slowly to climate warming, whereas less is known about the community functional trait responses and consequences for ecosystem functioning. The slow species turnover and large distribution ranges of many Arctic plant species suggest a significant role of intraspecific trait variability in functional responses to climate change. Here we compare taxonomic and functional community compositional responses to a long-term (17-year) warming experiment in Svalbard, Norway, replicated across three major high Arctic habitats shaped by topography and contrasting snow regimes. We observed taxonomic compositional changes in all plant communities over time. Still, responses to experimental warming were minor and most pronounced in the drier habitats with relatively early snowmelt timing and long growing seasons (Cassiope and Dryas heaths). The habitats were clearly separated in functional trait space, defined by 12 size- and leaf economics-related traits, primarily due to interspecific trait variation. Functional traits also responded to experimental warming, most prominently in the Dryas heath and mostly due to intraspecific trait variation. Leaf area and mass increased and leaf δ15N decreased in response to the warming treatment. Intraspecific trait variability ranged between 30% and 71% of the total trait variation, reflecting the functional resilience of those communities, dominated by long-lived plants, due to either phenotypic plasticity or genotypic variation, which most likely underlies the observed resistance of high Arctic vegetation to climate warming. We further explored the consequences of trait variability for ecosystem functioning by measuring peak season CO2 fluxes. Together, environmental, taxonomic, and functional trait variables explained a large proportion of the variation in net ecosystem exchange (NEE), which increased when intraspecific trait variation was accounted for. In contrast, even though ecosystem respiration and gross ecosystem production both increased in response to warming across habitats, they were mainly driven by the direct kinetic impacts of temperature on plant physiology and biochemical processes. Our study shows that long-term experimental warming has a modest but significant effect on plant community functional trait composition and suggests that intraspecific trait variability is a key feature underlying high Arctic ecosystem resistance to climate warming.publishedVersio

Herbivory and warming have opposing short-term effects on plant-community nutrient levels across high-Arctic tundra habitats

Environmental changes can rapidly alter standing biomass in tundra plant communities; yet, to what extent can they modify plant-community nutrient levels? Nutrient levels and their changes can affect biomass production, nutrient cycling rates and nutrient availability to herbivores. We examined how environmental perturbations alter Arctic plant-community leaf nutrient concentrations (percentage of dry mass, i.e. resource quality) and nutrient pools (absolute mass per unit area, i.e. resource quantity). We experimentally imposed two different types of environmental perturbations in a high-Arctic ecosystem in Svalbard, spanning three habitats differing in soil moisture and plant-community composition. We mimicked both a pulse perturbation (a grubbing event by geese in spring) and a press perturbation (a constant level of summer warming). After 2 years of perturbations, we quantified peak-season nitrogen and phosphorus concentrations in 1268 leaf samples from the most abundant vascular plant species. We derived community-weighted nutrient concentrations and total amount of nutrients (pools) for whole plant communities and individual plant functional types (PFTs). Spring grubbing increased plant-community nutrient concentrations in mesic (+13%) and wet (+8%), but not moist, habitats, and reduced nutrient pools in all habitats (moist: −49%; wet, mesic: −31% to −37%). Conversely, summer warming reduced plant-community nutrient concentrations in mesic and moist (−10% to −12%), but not wet, habitats and increased nutrient pools in moist habitats (+50%). Fast-growing PFTs exhibited nutrient-concentration responses, while slow-growing PFTs generally did not. Grubbing enhanced nutrient concentrations of forbs and grasses in wet habitats (+20%) and of horsetails and grasses in mesic habitats (+19–23%). Conversely, warming decreased nutrient concentrations of horsetails in wet habitats (−15%) and of grasses, horsetails and forbs in moist habitats (−12% to −15%). Nutrient pools held by each PFT were less affected, although the most abundant PFTs responded to perturbations. Synthesis. Arctic plant-community nutrient levels can be rapidly altered by environmental changes, with consequences for short-term process rates and plant-herbivore interactions. Community-level responses in nutrient concentrations and pools were opposing and differed among habitats and PFTs. Our findings have implications for how we understand herbivory- and warming-induced shifts in the fine-scaled distribution of resource quality and quantity within and across tundra habitats

Definition of sampling units begets conclusions in ecology: The case of habitats for plant communities

In ecology, expert knowledge on habitat characteristics is often used to define sampling units such as study sites. Ecologists are especially prone to such approaches when prior sampling frames are not accessible. Here we ask to what extent can different approaches to the definition of sampling units influence the conclusions that are drawn from an ecological study? We do this by comparing a formal versus a subjective definition of sampling units within a study design which is based on well-articulated objectives and proper methodology. Both approaches are applied to tundra plant communities in mesic and snowbed habitats. For the formal approach, sampling units were first defined for each habitat in concave terrain of suitable slope using GIS. In the field, these units were only accepted as the targeted habitats if additional criteria for vegetation cover were fulfilled. For the subjective approach, sampling units were defined visually in the field, based on typical plant communities of mesic and snowbed habitats. For each approach, we collected information about plant community characteristics within a total of 11 mesic and seven snowbed units distributed between two herding districts of contrasting reindeer density. Results from the two approaches differed significantly in several plant community characteristics in both mesic and snowbed habitats. Furthermore, differences between the two approaches were not consistent because their magnitude and direction differed both between the two habitats and the two reindeer herding districts. Consequently, we could draw different conclusions on how plant diversity and relative abundance of functional groups are differentiated between the two habitats depending on the approach used. We therefore challenge ecologists to formalize the expert knowledge applied to define sampling units through a set of well-articulated rules, rather than applying it subjectively. We see this as instrumental for progress in ecology as only rules based on expert knowledge are transparent and lead to results reproducible by other ecologists

Hiding in the background: community-level patterns in invertebrate herbivory across the tundra biome

Invertebrate herbivores depend on external temperature for growth and metabolism. Continued warming in tundra ecosystems is proposed to result in increased invertebrate herbivory. However, empirical data about how current levels of invertebrate herbivory vary across the Arctic is limited and generally restricted to a single host plant or a small group of species, so predicting future change remains challenging. We investigated large-scale patterns of invertebrate herbivory across the tundra biome at the community level and explored how these patterns are related to long-term climatic conditions and year-of-sampling weather, habitat characteristics, and aboveground biomass production. Utilizing a standardized protocol, we collected samples from 92 plots nested within 20 tundra sites during summer 2015. We estimated the community-weighted biomass lost based on the total leaf area consumed by invertebrates for the most common plant species within each plot. Overall, invertebrate herbivory was prevalent at low intensities across the tundra, with estimates averaging 0.94% and ranging between 0.02 and 5.69% of plant biomass. Our results suggest that mid-summer temperature influences the intensity of invertebrate herbivory at the community level, consistent with the hypothesis that climate warming should increase plant losses to invertebrates in the tundra. However, most of the observed variation in herbivory was associated with other site level characteristics, indicating that other local ecological factors also play an important role. More details about the local drivers of invertebrate herbivory are necessary to predict the consequences for rapidly changing tundra ecosystems