Amy E. Miller

Microbial Community Succession in an Unvegetated, Recently Deglaciated Soil

Primary succession is a fundamental process in macroecosystems; however, if and how soil development influences microbial community structure is poorly understood. Thus, we investigated changes in the bacterial community along a chronosequence of three unvegetated, early successional soils (∼20-year age gradient) from a receding glacier in southeastern Peru using molecular phylogenetic techniques. We found that evenness, phylogenetic diversity, and the number of phylotypes were lowest in the youngest soils, increased in the intermediate aged soils, and plateaued in the oldest soils. This increase in diversity was commensurate with an increase in the number of sequences related to common soil bacteria in the older soils, including members of the divisions Acidobacteria, Bacteroidetes, and Verrucomicrobia. Sequences related to the Comamonadaceae clade of the Betaproteobacteria were dominant in the youngest soil, decreased in abundance in the intermediate age soil, and were not detected in the oldest soil. These sequences are closely related to culturable heterotrophs from rock and ice environments, suggesting that they originated from organisms living within or below the glacier. Sequences related to a variety of nitrogen (N)-fixing clades within the Cyanobacteria were abundant along the chronosequence, comprising 6–40% of phylotypes along the age gradient. Although there was no obvious change in the overall abundance of cyanobacterial sequences along the chronosequence, there was a dramatic shift in the abundance of specific cyanobacterial phylotypes, with the intermediate aged soils containing the greatest diversity of these sequences. Most soil biogeochemical characteristics showed little change along this ∼20-year soil age gradient; however, soil N pools significantly increased with soil age, perhaps as a result of the activity of the N-fixing Cyanobacteria. Our results suggest that, like macrobial communities, soil microbial communities are structured by substrate age, and that they, too, undergo predictable changes through time.

Niche complementarity due to plasticity in resource use: plant partitioning of chemical N forms

Niche complementarity, in which coexisting species use different forms of a resource, has been widely invoked to explain some of the most debated patterns in ecology, including maintenance of diversity and relationships between diversity and ecosystem function. However, classical models assume resource specialization in the form of distinct niches, which does not obviously apply to the broadly overlapping resource use in plant communities. Here we utilize an experimental framework based on competition theory to test whether plants partition resources via classical niche differentiation or via plasticity in resource use. We explore two alternatives: niche preemption, in which individuals respond to a superior competitor by switching to an alternative, less-used resource, and dominant plasticity, in which superior competitors exhibit high resource use plasticity and shift resource use depending on the competitive environment. We determined competitive ability by measuring growth responses with and without neighbors over a growing season and then used 15N tracer techniques to measure uptake of different nitrogen (N) forms in a field setting. We show that four alpine plant species of differing competitive abilities have statistically indistinguishable uptake patterns (nitrate > ammonium > glycine) in their fundamental niche (without competitors) but differ in whether they shift these uptake patterns in their realized niche (with competitors). Competitively superior species increased their uptake of the most available N form, ammonium, when in competition with the rarer, competitively inferior species. In contrast, the competitively inferior species did not alter its N uptake pattern in competition. The existence of plasticity in resource use among the dominant species provides a mechanism that helps to explain the manner by which plant species with broadly overlapping resource use might coexist.

Alpine plants show species-level differences in the uptake of organic and inorganic nitrogen

As an estimate of species-level differences in the capacity to take up different forms of N, we measured plant uptake of 15N-NH4+, 15N-NO3− and 15N, [1]-13C glycine within a set of herbaceous species collected from three alpine community types. Plants grown from cuttings in the greenhouse showed similar growth responses to the three forms of N but varied in the capacity to take up NH4+, NO3− and glycine. Glycine uptake ranged from approximately 42% to greater than 100% of NH4+ uptake; however, four out of nine species showed significantly greater uptake of either NH4+ or NO3− than of glycine. Relative concentrations of exchangeable N at the sites of plant collection did not correspond with patterns of N uptake among species; instead, species from the same community varied widely in the capacity to take up NH4+, NO3−, and glycine, suggesting the potential for differentiation among species in resource (N) use.

PLANT UPTAKE OF INORGANIC AND ORGANIC NITROGEN: NEIGHBOR IDENTITY MATTERS

The importance of interspecific competition as a cause of resource partitioning among species has been widely assumed but rarely tested. Using neighbor removals in combination with 15N tracer additions in the field, we examined variation among three alpine species in the uptake of 15N-NH4+, 15N-NO3-, and 15N-13C-[2]-glycine in intact neighborhoods, when paired with a specific neighbor, and when all neighbors were removed. Species varied in the capacity to take up 15N-labeled NH4+, NO3-, and glycine in intact neighborhoods and in interspecific pairs. When interspecific neighbor pairs were compared with no neighbor controls, neighbors reduced 15N uptake in target species by as much as 50%, indicating competition for N. Furthermore, neighbor identity influenced the capacity of species to take up different forms of N. Thus, competition within interspecific neighbor pairs often caused reduced uptake of a particular form of N, as well as shifts to uptake of an alternative form of N. Such shifts in resource use as a result of competition are an implicit assumption in studies of resource partitioning but have rarely been documented. Our study suggests that plasticity in the uptake of different forms of N may be a mechanism by which cooccurring plants reduce competition for N.

Climate variability and spruce beetle (Dendroctonus rufipennis) outbreaks in south‐central and southwest Alaska

We used tree ring data (AD 1601-2007) to examine the occurrence of and climatic influences on spruce beetle (Dendroctonus rufipennis) outbreaks in south-central and southwest Alaska and found evidence of regional-scale outbreaks dating from the mid-1700s, related to climate variability at multiple temporal scales. Over interannual time scales (approximately 1-3 years), El Niño years, combined with severe late-summer drought, appeared to contribute significantly to spruce beetle outbreaks in the study area. Over multidecadal time scales (up to approximately 40 years), cool-phase Pacific Decadal Oscillation (PDO) conditions tended to precede beetle outbreaks, regardless of the phase of El Niño-Southern Oscillation (ENSO). All sites showed low-severity disturbances attributed to spruce beetle damage, most notably during the 1810s. During other major periods of disturbance (i.e., 1870s, 1910s, 1970s), the effects of spruce beetle outbreaks were of moderate or higher severity. The highly synchronized timing of spruce beetle outbreaks at interannual to multidecadal scales, and particularly the association between cool-phase PDO conditions and beetle disturbance, suggests that climate (i.e., temperature, precipitation) is a primary driver of outbreaks in the study area. Our disturbance chronologies (mid-1700s to present) suggest that recent irruptions (1990s to present) in south-central and southwest Alaska are within the historical geographic range, but that outbreaks since the 1990s show greater spatiotemporal synchrony (i.e., more sites record high-severity infestations) than at any other time in the past approximatly 250 years.

Nitrogen preferences and plant-soil feedbacks as influenced by neighbors in the alpine tundra

Plant resource partitioning of chemical forms of nitrogen (N) may be an important factor promoting species coexistence in N-limited ecosystems. Since the microbial community regulates N-form transformations, plant partitioning of N may be related to plant–soil feedbacks. We conducted a 15N tracer addition experiment to study the ability of two alpine plant species, Acomastylis rossii and Deschampsia caespitosa, to partition organic and inorganic forms of N. The species are codominant and associated with strong plant–soil feedbacks that affect N cycling. We manipulated interspecific interactions by removing Acomastylis or Deschampsia from areas where the species were codominant to test if N uptake patterns varied in the presence of the other species. We found that Deschampsia acquired organic and inorganic N more rapidly than Acomastylis, regardless of neighbor treatment. Plant N uptake—specifically ammonium uptake—increased with plant density and the presence of an interspecific neighbor. Interestingly, this change in N uptake was not in the expected direction to reduce niche overlap and instead suggested facilitation of ammonium use. To test if N acquisition patterns were consistent with plant–soil feedbacks, we also compared microbial rhizosphere extracellular enzyme activity in patches dominated by one or the other species and in areas where they grew together. The presence of both species was generally associated with increased rhizosphere extracellular enzyme activity (five of ten enzymes) and a trend towards increased foliar N concentrations. Taken together, these results suggest that feedbacks through the microbial community, either in response to increased plant density or specific plant neighbors, could facilitate coexistence. However, coexistence is promoted via enhanced resource uptake rather than reduced niche overlap. The importance of resource partitioning to reduce the intensity of competitive interactions might vary across systems, particularly as a function of plant-soil feedbacks.

The consequence of species loss on ecosystem nitrogen cycling depends on community compensation

Repercussions of species loss on ecosystem processes depend on the effects of the lost species as well as the compensatory responses of the remaining species in the community. We experimentally removed two co-dominant plant species and added a 15N tracer in alpine tundra to compare how species’ functional differences influence community structure and N cycling. For both of the species, production compensated for the biomass removed by the second year. However, the responses of the remaining species depended on which species was removed. These differences in compensation influenced how species loss impacted ecosystem processes. After the removal of one of the co-dominant species, Acomastylis rossii, there were few changes in the relative abundance of the remaining species, and differences in functioning could be predicted based on effects associated with the removed species. In contrast, the removal of the other co-dominant, Deschampsia caespitosa, was associated with subsequent changes in community structure (species relative abundances and diversity) and impacts on ecosystem properties (microbial biomass N, dissolved organic N, and N uptake of subordinate species). Variation in compensation may contribute to the resulting effects on ecosystem functioning, with the potential to buffer or accelerate the effects of species loss.

Deriving Snow Cover Metrics for Alaska from MODIS

Moderate Resolution Imaging Spectroradiometer (MODIS) daily snow cover products provide an opportunity for determining snow onset and melt dates across broad geographic regions; however, cloud cover and polar darkness are limiting factors at higher latitudes. This study presents snow onset and melt dates for Alaska, portions of western Canada and the Russian Far East derived from Terra MODIS snow cover daily 500 m grid data (MOD10A1) and evaluates our method for filling data gaps caused by clouds or polar darkness. Pixels classified as cloud or no data were reclassified by: spatial filtering using neighboring pixel values; temporal filtering using pixel values for days before/after cloud cover; and snow-cycle filtering based on a time series assessment of a pixel’s position within snow accumulation, cover or melt periods. During the 2012 snow year, these gap-filling methods reduced cloud pixels from 27.7% to 3.1%. A total of 12 metrics (e.g., date of first and last snow, date of persistent snow cover and periods of intermittence) for each pixel were calculated by snow year. A comparison of MODIS-derived snow onset and melt dates with in situ observations from 244 weather stations generally showed an early bias in MODIS-derived dates and an effect of increasing cloudiness exacerbating bias. Our results show that mean regional duration of seasonal snow cover is 179–311 days/year and that snow cover is often intermittent, with 41% of the area experiencing ≥2 snow-covered periods during a snow season. Other regional-scale patterns in the timing of snow onset and melt are evident in the yearly 500 m gridded products publically available at http://static.gina.alaska.edu/NPS_products/MODIS_snow/.

Tree‐ring isotopes reveal drought sensitivity in trees killed by spruce beetle outbreaks in south‐central Alaska

Increasing temperatures have resulted in reduced growth and increased tree mortality across large areas of western North American forests. We use tree-ring isotope chronologies (δ13 C and δ18 O) from live and dead trees from four locations in south-central Alaska, USA, to test whether white spruce trees killed by recent spruce beetle (Dendroctonus rufipennis Kirby) outbreaks showed evidence of drought stress prior to death. Trees that were killed were more sensitive to spring/summer temperature and/or precipitation than trees that survived. At two of our sites, we found greater correlations between the δ13 C and δ18 O chronologies and spring/summer temperatures in dead trees than in live trees, suggesting that trees that are more sensitive to temperature-induced drought stress are more likely to be killed. At one site, the difference between δ13 C in live and dead trees was related to winter/spring precipitation, with dead trees showing stronger correlations between δ13 C and precipitation, again suggesting increased water stress in dead trees. At all sites where δ18 O was measured, δ18 O chronologies showed the greatest difference in climate response between live and dead groups, with δ18 O in live trees correlating more strongly with late winter precipitation than dead trees. Our results indicate that sites where trees are already sensitive to warm or dry early growing-season conditions experienced the most beetle-kill, which has important implications for forecasting future mortality events in Alaska.