Landscape patterns of free amino acids in arctic tundra soils

Concentrations of free amino acids were measured in soils from four major ecosystem types in arctic Alaska. Total free amino acid concentrations were several-fold higher than ammonium (the major form of inorganic nitrogen) in water extracts of soils. The dominant free amino acids in these soils were glycine, aspartic acid, glutamic acid, serine, and arginine. Concentrations of total amino acids ranged 5-fold across communities, being highest in tussock tundra and lowest in wet meadows. Incubation experiments indicate that the turnover of amino acids is rapid, which suggests high rates of gross nitrogen mineralization in these soils. The high concentrations and dynamic nature of soil free amino acids suggest that this nitrogen pool is a significant component of nitrogen cycling in these tundra ecosystems.     

Can antibrowsing defense regulate the spread of woody vegetation in arctic tundra?

Global climate warming is projected to promote the increase of woody plants, especially shrubs, in arctic tundra. Many factors may affect the extent of this increase, including browsing by mammals. We hypothesize that across the Arctic the effect of browsing will vary because of regional variation in antibrowsing chemical defense. Using birch (Betula) as a case study, we propose that browsing is unlikely to retard birch expansion in the region extending eastward from the Lena River in central Siberia across Beringia and the continental tundra of central and eastern Canada where the more effectively defended resin birches predominate. Browsing is more likely to retard birch expansion in tundra west of the Lena to Fennoscandia, Iceland, Greenland and South Baffin Island where the less effectively defended non‐resin birches predominate. Evidence from the literature supports this hypothesis. We further suggest that the effect of warming on the supply of plant‐available nitrogen will not significantly change either this pan‐Arctic pattern of variation in antibrowsing defense or the resultant effect that browsing has on birch expansion in tundra. However, within central and east Beringia warming‐caused increases in plant‐available nitrogen combined with wildfire could initiate amplifying feedback loops that could accelerate shrubification of tundra by the more effectively defended resin birches. This accelerated shrubification of tundra by resin birch, if extensive, could reduce the food supply of caribou causing population declines. We conclude with a brief discussion of modeling methods that show promise in projecting invasion of tundra by woody plants.     

Disentangling vegetation diversity from climate–energy and habitat heterogeneity for explaining animal geographic patterns

Broad‐scale animal diversity patterns have been traditionally explained by hypotheses focused on climate–energy and habitat heterogeneity, without considering the direct influence of vegetation structure and composition. However, integrating these factors when considering plant–animal correlates still poses a major challenge because plant communities are controlled by abiotic factors that may, at the same time, influence animal distributions. By testing whether the number and variation of plant community types in Europe explain country‐level diversity in six animal groups, we propose a conceptual framework in which vegetation diversity represents a bridge between abiotic factors and animal diversity. We show that vegetation diversity explains variation in animal richness not accounted for by altitudinal range or potential evapotranspiration, being the best predictor for butterflies, beetles, and amphibians. Moreover, the dissimilarity of plant community types explains the highest proportion of variation in animal assemblages across the studied regions, an effect that outperforms the effect of climate and their shared contribution with pure spatial variation. Our results at the country level suggest that vegetation diversity, as estimated from broad‐scale classifications of plant communities, may contribute to our understanding of animal richness and may be disentangled, at least to a degree, from climate–energy and abiotic habitat heterogeneity.     

Downslope fertilizer movement in arctic tussock tundra

Slow release fertilizer pellets (NPK) were spread in June and July 1985, on the frost surface under an 8–20 cm thick layer of thawed tundra in the northern foothills of the Brooks Range, Alaska. The fertilizer was applied at a rate of 3.8 kg N, 1.2 kg P₂O₅, and 2.4 kg K to two, 10 × 0.75 m tussock bands. These bands were placed parallel to the contour lines in the middle of a uniform 10° slope. The purpose of the experiment was to test the hypothesis that fertilizer moves at a measurable rate downslope, affecting the vegetation in its course. The results support this hypothesis. One year after fertilizer application, plants collected 2 to 6 m downslope from the fertilizer band had significantly larger leaves and significantly higher nitrogen and phosphorus concentrations than those upslope from fertilizer application. It is concluded that nutrient-releasing tundra perturbations affect the downslope tundra vegetation.     

Open tundra persist,but arctic features decline—Vegetation changes in the warming Fennoscandian tundra

In the forest‐tundra ecotone of the North Fennoscandian inland, summer and winter temperatures have increased by two to three centigrades since 1965, which is expected to result in major vegetation changes. To document the expected expansion of woodlands and scrublands and its impact on the arctic vegetation, we repeated a vegetation transect study conducted in 1976 in the Darju, spanning from woodland to a summit, 200 m above the tree line. Contrary to our expectations, tree line movement was not detected, and there was no increase in willows or shrubby mountain birches, either. Nevertheless, the stability of tundra was apparent. Small‐sized, poorly competing arctic species had declined, lichen cover had decreased, and vascular plants, especially evergreen ericoid dwarf shrubs, had gained ground. The novel climate seems to favour competitive clonal species and species thriving in closed vegetation, creating a community hostile for seedling establishment, but equally hostile for many arctic species, too. Preventing trees and shrubs from invading the tundra is thus not sufficient for conserving arctic biota in the changing climate. The only dependable cure is to stop the global warming.     

Seasonal root growth in the arctic tussock tundra

Summary The quantity of growing root tips per unit of soil volume was analyzed in a central Alaskan tussock tundra site. By June 10, the aboveground fraction of the vegetation had initiated the flush of spring growth and flowering while less than 5 active root tips cm^-3 were found. By June 25 this value had increased to 30 root tips cm^-3. Similar values in July were followed by a complete cessation of root growth after the first week of August. By then, leaf senescence had also become visible. In the spring, low root temperatures were responsible for the time lag between shoot growth initiation and the beginning of root growth. In early August, root growth stopped in spite of adequate soil temperatures and accumulated carbohydrate for root growth. It is proposed that use of reserve carbohydrate for root growth in August would compromise the flush of spring growth in the following year.     

Fading indirect effects in a warming arctic tundra

Indirect interactions in food webs can strongly influence the net effect of global change on ecological communities yet they are rarely quantified and hence remain poorly understood. Using a 22-year time series, we investigated climate-induced and predator-mediated indirect effects on grazing intensity in the tundra food web of Bylot Island, which experienced a warming trend over the last two decades. We evaluated the relative effects of environmental parameters on the proportion of plant biomass grazed by geese in wetlands and examined the temporal changes in the strength of these cascading effects. Migrating geese are the dominant herbivores on Bylot Island and can consume up to 60% of the annual production of wetland graminoids. Spring North Atlantic Oscillation, mid-summer temperatures and summer abundance of lemmings （prey sharing predators with geese） best-explained annual variation in grazing intensity. Goose grazing impact increased in years with high temperatures and high lemming abundance. However, the strength of these indirect effects on plants changed over time. Grazing intensity was weakly explained by environmental factors in recent years, which were marked by a sharp increase in plant primary production and steady decrease in grazing pressure. Indirect effects do not seem to be reversing the direct positive effect of warming on wetland plants. We suggest that cascading effects on plants may lag considerably behind direct effects in vertebrate dominated arctic communities, especially where key herbivore populations are strongly affected by factors outside of the Arctic [Current Zoology 60 （2）： 189-202, 2014].     

Aboveground biomass allocation,leaf growth,and photosynthesis patterns in tundra plant forms in arctic Alaska

Summary Tundra plant growth forms can generally be characterized as consisting predominantly of low-growing perennial grasses and sedges, perennial herbaceous forbs, dwarf deciduous shrubs, and dwarf evergreen shrubs. Gross aboveground carbon allocation, leaf growth, and photosynthesis pattern studies were initiated to develop a quantitative understanding of the functional importance of these particular tundra growth forms. Photosynthetic capacities of 13 species were determined under standardized exposure conditions using a¹⁴CO₂ field system and ranged between 5 and 47 mg CO₂·g dry wt^-1·h^-1. These results, in conjunction with detailed leaf growth determinations, support the generalization that species with an evergreen growth form have lower photosynthetic capacities than species with a perennial graminoid, forb, or deciduous shrub growth form. However, these low photosynthetic capacities in evergreen shrubs are associated with relatively extended leaf longevities. Conversely, deciduous shrub forms exhibited high photosynthetic capacities, but were offset by relatively short leaf longevity periods. The perennial grasses, sedges, and forbs showed patterns intermediate to these. As a result, it appears that among tundra species of different growth form, photosynthetic capacity is inversely related to leaf longevity.     

Production and export of dissolved C in arctic tundra mesocosms: the roles of vegetation and water flow

To better understand carbon (C) cycling in arctic tundra we measureddissolved C production and export rates in mesocosms of three tundra vegetationtypes: tussock, inter-tussock and wet sedge. Three flushing frequencies wereused to simulate storm events and determine potential mass export of dissolved Cunder increased soil water flow scenarios. Dissolved C production and exportrates differed between vegetation types (inter-tussock < tussock < wetsedge). In the absence of flushing, dissolved organic C (DOC) dominatedproduction in tussock and inter-tussock soils but was consumed in wet sedgesoils (8.3, 32.7, and –0.4 g C g soil^–1day^–1). Soil water dissolved C concentrations declined over time when flushedat high and medium frequencies but were variable at low flushing frequency.Total yield of dissolved C and DOC increased with increased flushing frequency.The ratio of DOC to dissolved inorganic C exported dropped with increasedflushing under tussock but not inter-tussock or wet sedge vegetation. Massexport per liter of water added declined as flushing frequency increased intussock and inter-tussock mesocosms. Export and production of dissolved C werestrongly correlated with above ground biomass, but not with photosynthetic ratesor below ground biomass. DOC quality was examined by measuring production ofToolik Lake bacteria fed mesocosm soil water. When normalized for DOCconcentration, wet sedge soil water supported significantly higher bacterialproduction. Our results indicate that arctic tundra soils have high potentialsfor dissolved C export, that water flow and vegetation type mainly controldissolved C export, and that responses of aquatic microbes to terrestrial inputsdepend on the vegetation type in the watershed.     

Plant–herbivore interactions in Alaskan arctic tundra change with soil nutrient availability

Herbivores in nutrient‐limited systems such as arctic tundra have been suggested to play a minor role in controlling plant growth simply because they are relatively few in number. However, theory predicts that as net primary productivity (NPP) increases because of greater inputs of nutrients or energy, herbivores may have greater effects on plant growth. This prediction has not been tested in the context of climate warming in arctic tundra, which may increase soil nutrient availability and thus NPP. We examined a long‐term experiment that excluded small and large mammalian herbivores and increased soil nutrients in two arctic Alaskan tundra communities: dry heath (DH) and moist acidic tussock (MAT). In the ninth year of manipulations, we measured weekly growth of three plant species of three growth forms: tussock‐forming graminoid, rhizomatous graminoid, and dwarf deciduous shrub, in each community. All species grew better when fertilized. In DH, this increase in growth was exaggerated when plants were protected from herbivores, confirming that herbivory had a negative effect on plant growth under increased nutrient conditions, but was unimportant under ambient soil conditions. However, in MAT, the importance of herbivory differed among species with fertilization. The tussock‐forming sedge at MAT, Eriophorum vaginatum, grew better and flowered more when fenced under both ambient and amended nutrients compared to plants exposed to herbivores. This species decreases in abundance in long‐term fertilized plots when mammals are present, and our results suggest that herbivory may be accounting for at least some of that loss, in addition to shifts in competitive relationships. Although we only focused on individual plants here rather than the entire community, our results suggest that under the increased soil nutrient conditions expected with continued climate warming in the Arctic, herbivores may become more important in affecting several abundant tundra plant populations, and should not be ignored.     

Mycorrhizal colonization mediated by species interactions in arctic tundra

The Alaskan tussock tundra is a strongly nutrient-limited ecosystem, where almost all vascular plant species are mycorrhizal. We established a long-term removal experiment to document effects of arctic plant species on ecto- and ericoid mycorrhizal fungi and to investigate whether species interactions and/or nutrient availability affect mycorrhizal colonization. The treatments applied were removal of Betula nana (Betulaceae, dominant deciduous shrub species), removal of Ledum palustre (Ericaceae, dominant evergreen shrub species), control (no removal), and each of these three treatments with the addition of fertilizer. After 3 years of Ledum removal and fertilization, we found that overall ectomycorrhizal colonization in Betula was significantly reduced. Changes in ectomycorrhizal morphotype composition in removal and fertilized treatments were also observed. These results suggest that the effect of Ledum on Betula 's mycorrhizal roots is due to sequestration of nutrients by Ledum, leading to reduced nutrient availability in the soil. In contrast, ericoid mycorrhizal colonization was not affected by fertilization, but the removal of Betula and to a lower degree of Ledum resulted in a reduction of ericoid mycorrhizal colonization suggesting a direct effect of these species on ericoid mycorrhizal colonization. Nutrient availability was only higher in fertilized treatments, but caution should be taken with the interpretation of these data as soil microbes may effectively compete with the ion exchange resins for the nutrients released by plant removal in these nutrient-limited soils.     

Factors determining plant species richness in Alaskan arctic tundra

Abstract. We studied the relationship between plant N:P ratio, soil characteristics and species richness in wet sedge and tussock tundra in northern Alaska at seven sites. We also collected data on soil characteristics, above‐ground biomass, species richness and composition. The N:P ratio of the vegetation did not show any relationship with species richness. The N:P ratio of the soil was related with species richness for both vegetation types. Species richness in the tussock tundra was most strongly correlated with soil calcium content and soil pH, with a strong correlation between these two factors. N:P ratio of the soil was also correlated with soil pH. Other factors correlated with species richness were soil moisture and Sphagnum cover. Organic matter content was the factor most strongly correlated with species richness in the wet sedge vegetation. N:P ratio of the soil was strongly correlated with organic matter content. We conclude that N:P ratio in the vegetation is not an important factor determining species richness in arctic tundra and that species richness in arctic tundra is mainly determined by pH and flooding. In tussock tundra the pH, declining with soil age, in combination with Sphagnum growth strongly decreases species richness, while in wet sedge communities flooding over long periods of time creates less favourable conditions for species richness.     

Diurnal patterns of gas‐exchange and metabolic pools in tundra plants during three phases of the arctic growing season

Arctic tundra plant communities are subject to a short growing season that is the primary period in which carbon is sequestered for growth and survival. This period is often characterized by 24‐h photoperiods for several months a year. To compensate for the short growing season tundra plants may extend their carbon uptake capacity on a diurnal basis, but whether this is true remains unknown. Here, we examined in situ diurnal patterns of physiological activity and foliar metabolites during the early, mid, and late growing season in seven arctic species under light‐saturated conditions. We found clear diurnal patterns in photosynthesis and respiration, with midday peaks and midnight lulls indicative of circadian regulation. Diurnal patterns in foliar metabolite concentrations were less distinct between the species and across seasons, suggesting that metabolic pools are likely governed by proximate external factors. This understanding of diurnal physiology will also enhance the parameterization of process‐based models, which will aid in better predicting future carbon dynamics for the tundra. This becomes even more critical considering the rapid changes that are occurring circumpolarly that are altering plant community structure, function, and ultimately regional and global carbon budgets.     

Relationships between geomorphology and vegetation patterns in subantarctic Andean tundra of Tierra del Fuego

We analysed vegetation patterns in relation to geomorphology in subantarctic Andean tundra (Tierra del Fuego, southern Argentina). Habitat variation associated with geomorphology played a major role in structuring vegetation. On coarse-grained debris of talus cones, the vegetation is dominated by the epilithic fruticose lichen Neuropogon aurantiaco-ater, while vascular-plant cover is very low because of insufficient space available for seedling establishment, water shortage and, presumably, low nutrient status. Turf-banked lobes on moderately inclined slopes subject to downslope creep are characterised by vascular species forming large cushions (especially Azorella lycopodioides) which provide suitable microhabitats for several plants. Steep morainic slopes are extremely poor in species and have Marsippospermum reichei and Nardophyllum bryoides as characteristic species. The latter appears well adapted to withstand mechanical damage owing to its clonal creeping growth form. Screes represent intermediate habitats where the ecotonal species Bolax gummifera colonises debris and/or basal rock and may enhance the establishment of vascular species.     

Seasonal geochemistry of an arctic tundra drainage basin

The snow melt food at Imnavait Creek takes place sometime between 12 May and 2 June and constitutes the single most important hydrological and geochemical event. Three years of study indicate this event spans 7 to 10 days and that peak discharge can be expected to be between 0.6 and 0.9 cu. mes. Ion concentrations peak during the first 15% of the event while pH is at a minimum. In all cases, ion concentrations in the spring runoff are 4 to 9 times those of the snow pack, Precipitaion, including dryfall, contributes significant amounts of Ca, Mg, K, Na, Cl and SO₄. Postassium is present in surface waters only during melt-off and for a short time after. Calcium. Mg. suspended solids and electrical conductivity all reach broad, poorly defined peaks in mid-summer. Only pH shows a significant relationship to discharge. On a seasonal basis a substantial charge imbalance favoring cations occurs. It seems probable that the, as yet, unmeasured negative charge is associated with organic anions. No seasonal trends were recorded for Mg, K or Mn in subsurface flow in the surrounding slopes. Calcium. Fe and Al showed a late season peak, and the concentration of Na and Si decreased as the melt season progressed.     

Population dynamics of tundra voles in relation to configuration of willow thickets in southern arctic tundra

The areal extent and configuration of thickets of willow shrubs are currently changing in the Arctic both as an effect of global warming and changed browsing pressure of reindeer. These changes have been predicted to impact the distribution and abundance of wildlife species relying on willow thickets as habitat. We assessed the relation between variables quantifying willow thicket configuration and population dynamics of tundra voles (Microtus oeconomus) in three riparian regions in Finnmark, northern Norway, which were subject to intense browsing by semi-domesticated reindeer. The tundra vole, which exhibits 5-year population cycles in Finnmark, is the dominant small rodent species in riparian landscape elements in southern arctic tundra. In the course of a 4-year trapping study, tundra vole populations went through the cyclic phases of increase, peak and crash, however, with distinct differences between the three regions in the population dynamics. Within regions, the occupancy pattern during the increase phase was positively related to willow thicket configuration (in particular edge density and willow height) only in the region attaining the highest abundance and occupancy. However, local abundance was not clearly related to habitat features within any regions. The lack of consistency in the response of tundra vole populations to willow thicket configuration, as well as the positive relation between the degree of thicket shredding and tundra vole habitat occupancy in one of the regions, indicates that tundra voles will not be much affected by climate or browsing induced changes in the shrubbiness of the tundra in the future.     

Dispersal ability links to cross‐scale species diversity patterns across the Eurasian Arctic tundra

Aim The role of dispersal in structuring biodiversity across spatial scales is controversial. If dispersal controls regional and local community assembly, it should also affect the degree of spatial species turnover as well as the extent to which regional communities are represented in local communities. Here we provide the first integrated assessment of relationships between dispersal ability and local‐to‐regional spatial aspects of species diversity across a large geographical area. Location Northern Eurasia. Methods Using a cross‐scale analysis covering local (0.64 m²) to continental (the Eurasian Arctic biome) scales, we compared slope parameters of the dissimilarity‐to‐distance relationship in species composition and the local‐to‐regional relationship in species richness among three plant‐like groups that differ in dispersal ability: lichens with the highest dispersal ability; mosses and moss allies with intermediate dispersal ability; and seed plants with the lowest dispersal ability. Results Diversity patterns generally differed between the three groups according to their dispersal ability, even after controlling for niche‐based processes. Increasing dispersal ability is linked to decreasing spatial species turnover and an increasing ratio of local to regional species richness. All comparisons supported our expectations, except for the slope of the local‐to‐regional relationship in species richness for mosses and moss allies which was not significantly steeper than that of seed plants. Main conclusions The negative link between dispersal ability and spatial species turnover and the corresponding positive link between dispersal ability and the ratio of local‐to‐regional species richness support the idea that dispersal affects community structure and diversity patterns across spatial scales.     

Contrasting denitrifier communities relate to contrasting N2O emission patterns from acidic peat soils in arctic tundra

Cryoturbated peat circles (that is, bare surface soil mixed by frost action; pH 3–4) in the Russian discontinuous permafrost tundra are nitrate-rich ‘hotspots'' of nitrous oxide (N₂O) emissions in arctic ecosystems, whereas adjacent unturbated peat areas are not. N₂O was produced and subsequently consumed at pH 4 in unsupplemented anoxic microcosms with cryoturbated but not in those with unturbated peat soil. Nitrate, nitrite and acetylene stimulated net N₂O production of both soils in anoxic microcosms, indicating denitrification as the source of N₂O. Up to 500 and 10 μ nitrate stimulated denitrification in cryoturbated and unturbated peat soils, respectively. Apparent maximal reaction velocities of nitrite-dependent denitrification were 28 and 18 nmol N₂O g_DW⁻¹ h⁻¹, for cryoturbated and unturbated peat soils, respectively. Barcoded amplicon pyrosequencing of narG, nirK/nirS and nosZ (encoding nitrate, nitrite and N₂O reductases, respectively) yielded ≈49 000 quality-filtered sequences with an average sequence length of 444 bp. Up to 19 species-level operational taxonomic units were detected per soil and gene, many of which were distantly related to cultured denitrifiers or environmental sequences. Denitrification-associated gene diversity in cryoturbated and in unturbated peat soils differed. Quantitative PCR (inhibition-corrected per DNA extract) revealed higher copy numbers of narG in cryoturbated than in unturbated peat soil. Copy numbers of nirS were up to 1000 × higher than those of nirK in both soils, and nirS nirK⁻¹ copy number ratios in cryoturbated and unturbated peat soils differed. The collective data indicate that the contrasting N₂O emission patterns of cryoturbated and unturbated peat soils are associated with contrasting denitrifier communities.     

Controls on production of bryophytes in an arctic tundra stream

1. Two bryophyte taxa (Hygrohypnum spp. and, to a lesser extent, Fontinalis neomexicana) were abundant in riffles within phosphorus-fertilized reaches of the Kuparuk River (North Slope, Alaska), but were much less common in fertilized pools and virtually absent in unfertilized reaches of the river. We conducted field experiments using stems and clumps of both species and artificial bryophytes to test the hypotheses that bryophyte growth was strongly limited by low phosphorus concentrations in unfertilized reaches, and limited by epiphytes in fertilized pools. 2. Stem tips of Hygrohypnum spp. did not elongate when grown in unfertilized pool and riffle environments. In fertilized reaches, Hygrohypnum elongated significantly, although there was no significant difference in elongation of stem tips placed in pools [2.5 ± 0.9 cm (SD)] as opposed to riffles (2.8 ± 0.9 cm) for 32 days. 3. Stem tips of F. neomexicana elongated significantly in all sites. There was a significant difference in elongation of stem tips in control and fertilized riffles (2.1 ± 1.1 and 4.7 ± 0.1 cm, respectively) but not in tips grown in control and fertilized pools (2.8 ± 0.8 and 2.7 ± 0.9 cm, respectively). 4. Biomass increments in clumps of these same species followed similar patterns except in fertilized pools. Hygrohypnum spp. lost weight in control riffle environments and did not grow in pools, but accumulated 181 ± 44 and 335 ± 200% of initial biomass in fertilized riffles in 1992 (over 32 days) and 1993 (over 44 days), respectively. F. neomexicana accumulated 38 ± 39 and 98 ± 47% of initial biomass in 1992 in unfertilized and fertilized riffles, respectively. Total phosphorus concentrations of both bryophytes in 1992 were significantly greater when grown in fertilized riffles than control riffles. 5. Artificial mosses (untwisted, natural fibre rope) and clumps of Hygrohypnum spp. were used to assess effects of flow regime on derrital and epiphyte accumulation in the fertilized zone. Epiphyte and detrital mass was 4–4.5 times greater on average on artificial mosses in slow-flowing pool environments than in fast-flowing riffle environments. Epiphyte chlorophyll a was 4 times greater on Hygrohypnum clumps in pools than in riffles. This difference was probably brought about by increased detrital deposition and reduced grazing by invertebrates in pools. It is likely that both Hygrohypnum spp. and F. neomexicana could grow throughout the river, but are limited strongly by low phosphorus concentrations in unfertilized reaches and secondarily by detritus accumulation and interference competition with epiphytic algae in fertilized pools.