Nutrient resorption from senescing plant tissues is an important determinant of the fitness of plant populations in nutrient-poor ecosystems, because it makes plants less dependent on current nutrient uptake. Moreover, it can have significant “afterlife” effects through its impact on litter chemistry and litter decomposability. Little is known about the effects of climate change on nutrient resorption. We studied the effects of climate change treatments (including winter snow addition, and spring and/or summer warming) on nutrient resorption of four dominant species in a nutrient-poor subarctic peatland. These species were Betula nana (woody deciduous), Vaccinium uliginosum (woody deciduous), Calamagrostis lapponica (graminoid) and Rubus chamaemorus (forb). After five years of treatments both mature and senesced leaf N concentrations showed a small but significant overall reduction in response to the climate treatments. However, the effects were species-specific. For example, in the controls the N concentration in senesced leaves of Calamagrostis (3.0 +/- 0.2 mg N g(-1)) was about four times lower than for Rubus (11.2 +/- 0.2 mg N g(-1)). There were no significant treatment effects on N resorption efficiency (% of the N pool in mature leaves that is resorbed during senescence). The nitrogen resorption efficiency of Calamagrostis (about 80%) was higher than in the other three species (about 60%). Thus, climate change has only a minor impact on nutrient resorption parameters. However, given the substantial interspecific differences in these parameters, substantial changes in plant-soil feedbacks may be expected as a result of the observed changes in the species composition of high-latitude vegetation. These changes are species-specific and thus difficult to predict.
Soil microbes constitute an important control on nitrogen (N) turnover and retention in arctic ecosystems where N availability is the main constraint on primary production. Ectomycorrhizal (ECM) symbioses may facilitate plant competition for the specific N pools available in various arctic ecosystems. We report here our study on the N uptake patterns of coexisting plants and microbes at two tundra sites with contrasting dominance of the circumpolar ECM shrub Betula nana. We added equimolar mixtures of glycine-N, NH4+–N and NO3−–N, with one N form labelled with 15N at a time, and in the case of glycine, also labelled with 13C, either directly to the soil or to ECM fungal ingrowth bags. After 2 days, the vegetation contained 5.6, 7.7 and 9.1% (heath tundra) and 7.1, 14.3 and 12.5% (shrub tundra) of the glycine-, NH4+- and NO3−–15N, respectively, recovered in the plant–soil system, and the major part of 15N in the soil was immobilized by microbes (chloroform fumigation-extraction). In the subsequent 24 days, microbial N turnover transferred about half of the immobilized 15N to the non-extractable soil organic N pool, demonstrating that soil microbes played a major role in N turnover and retention in both tundra types. The ECM mycelial communities at the two tundras differed in N-form preferences, with a higher contribution of glycine to total N uptake at the heath tundra; however, the ECM mycelial communities at both sites strongly discriminated against NO3−. Betula nana did not directly reflect ECM mycelial N uptake, and we conclude that N uptake by ECM plants is modulated by the N uptake patterns of both fungal and plant components of the symbiosis and by competitive interactions in the soil. Our field study furthermore showed that intact free amino acids are potentially important N sources for arctic ECM fungi and plants as well as for soil microorganisms.
Plant species and growth forms differ widely in litter chemistry, which affects decay and may have important consequences for plant growth via e.g. the release of nutrients and growth-inhibitory compounds. We investigated the overall short-term (9.5 months) and medium-term (21.5 months) feedback effects of leaf litter quality and quantity on plant production, and tested whether growth forms can be used to generalise differences among litter species. Leaf litter effects of 21 sub-arctic vascular peatland species on Poa alpina test plants changed clearly with time. Across all growth forms, litter initially reduced plant biomass compared with untreated plants, particularly litters with a high decomposition rate or low initial lignin/P ratio. In the second year, however, litter effects were neutral or positive, and related to initial litter N concentration (positive), C/N, polyphenol/N and polyphenol/P ratios (all negative), but not to decomposability. Differences in effect size among several litter species were large, while differences in response to increasing litter quantities were not significant or of similar magnitude to differences in response to three contrasting litter species. Growth forms did not differ in initial litter effects, but second-year plant production showed a trend (P < 0.10) for differences in response to litters of different growth forms: evergreen shrubs < graminoids or deciduous shrubs < forbs. While long-persisting negative litter effects were predominant across all growth forms, our data indicate that even within nutrient-constrained ecosystems such as northern peatlands, vascular plant species, and possibly growth forms, differ in litter feedbacks to plant growth. Differences in the composition of undisturbed plant communities or species shifts induced by external disturbance, such as climate change, may therefore feedback strongly to plant biomass production and probably nutrient cycling rates in northern peatlands.
Environmental changes are likely to alter the chemical composition of plant tissues, including content and concentrations of secondary compounds, and thereby affect the food sources of herbivores. After 10 years of experimental increase of temperature, nutrient levels and light attenuation in a sub-arctic, alpine ecosystem, we investigated the effects on carbon based secondary compounds (CBSC) and nitrogen in one dominant deciduous dwarf shrub, Salix herbacea x polaris and two dominant evergreen dwarf shrubs, Cassiope tetragona and Vaccinium vitis-idaea throughout one growing season. The main aims were to compare the seasonal course and treatment effects on CBSC among the species, life forms and leaf cohorts and to examine whether the responses in different CBSC were consistent across compounds. The changes in leaf chemistry both during the season and in response to the treatments were higher in S. herbacea x polaris than in the corresponding current year’s leaf cohort of the evergreen C. tetragona. The changes were also much higher than in the 1-year-old leaves of the two evergreens probably due to differences in dilution and turnover of CBSC in growing and mature leaves paired with different rates of allocation. Most low molecular weight phenolics in the current year’s leaves decreased in all treatments. Condensed tannins and the tannin-to-N ratio, however, either increased or decreased, and the strength and even direction of the responses varied among the species and leaf cohorts, supporting views of influential factors additional to resource-based or developmental controls, as e.g. species specific or genetic controls of CBSC. The results indicate that there is no common response to environmental changes across species and substances. However, the pronounced treatment responses imply that the quality of the herbivore forage is likely to be strongly affected in a changing arctic environment, although both the direction and strength of the responses will be different among plant species, tissue types and substances.
The functional response is a key element of predator-prey interactions. Basic functional response theory explains foraging behavior of individual predators, but many empirical studies of free-ranging predators have estimated functional responses by using population-averaged data. We used a novel approach to investigate functional responses of an avian predator (the rough legged-buzzard Buteo lagopus Pontoppidan, 1763) to intra-annual spatial variation in rodent density in subarctic Sweden, using breeding pairs as the sampling unit. The rough-legged buzzards responded functionally to Norwegian lemmings (Lemmus lemmus L. 1758), grey-sided voles (Myodes rufocanus Sundevall, 1846) and field voles (Microtus agrestis L. 1761), but different rodent prey were not utilised according to relative abundance. The functional response to Norwegian lemmings was a steep type II curve and a more shallow type III response to grey-sided voles. The different shapes of these two functional responses were likely due to combined effects of differences between lemmings and grey-sided voles in habitat utilisation, anti-predator behaviour and size-dependent vulnerability to predation. Diet composition changed less than changes in relative prey abundance, indicating negative switching, with high disproportional use of especially lemmings at low relative densities. Our results suggest that lemmings and voles should be treated separately in future empirical and theoretical studies in order to better understand the role of predation in this study system.
The carbon charging of pines across the treeline ecotone of three different climatic zones (Mexico 19degreesN Pinus hartwegii, Swiss Alps 46degreesN P. cembra and northern Sweden 68degreesN P. sylvestris) was analyzed, to test whether a low-temperature-driven carbon shortage can explain high-elevation tree limits, and whether the length of the growing season affects the trees’ carbon balance. We quantified the concentrations of non-structural carbohydrates (NSC) and lipids (acylglycerols) in all tree organs at three dates during the growing seasons across elevational transects from the upper end of the closed, tall forest (timberline) to the uppermost location where groups of trees greater than or equal to3 m in height occur (treeline). Mean ground temperatures during the growing season at the treelines were similar (6.1 +/- 0.7degreesC) irrespective of latitude. Across the individual transects, the concentrations of NSC and lipids increased with elevation in all organs. By the end of the growing season, all three species had very similar total mobile carbon (TMC) concentrations at the treeline (ca. 6% TMC in the aboveground dry biomass), suggesting no influence of the length of the growing season on tree carbon charging. At a temperate lowland reference site P. sylvestris reached only ca. 4% TMC in the aboveground dry biomass, with the 2% difference largely explained by higher lipid concentrations of treeline pines. We conclude that carbon availability is unlikely to be the cause of the altitudinal tree limit. It seems rather that low temperatures directly affect sink activity at the treeline, with surplus carbon stored in osmotically inactive compounds.
Hemiparasitic angiosperms concentrate nutrients in their leaves and also produce high quality litter, which can decompose faster and release more nutrients than that of surrounding species. The impact of these litters on plant growth may be particularly important in nutrient-poor communities where hemiparisites can be abundant, such as the sub-Arctic. We tested the hypothesis that plant growth is enhanced by the litter of the hemiparasite Bartsia alpina, in comparison with litter of co-occurring dwarf shrub species, using a pot based bioassay approach. Growth of Betula nana and Poa alpina was up to 51% and 41% greater, respectively, in the presence of Bartsia alpina litter than when grown with dwarf shrub litter (Vaccinium uliginosum, Betula nana and Empetrum nigrum subsp. hermaphroditum). The nutrient concentrations of Betula nana plants grown with Bartsia alpina litter were almost double those of plants grown with dwarf shrub litter, and a significantly greater proportion of biomass was allocated to shoots rather than roots, strongly suggesting that nutrient availability was higher where Bartsia alpina litter was present. The presence of litter from dwarf shrubs, or the moss Hylocomium splendens, did not reduce the positive effect of Bartsia alpina litter on plant growth. E. nigrum litter did not appear to affect plant growth substantially differently from litter of other dwarf shrub species, despite earlier reports. of its allelopathic action. The enhanced nutrient uptake and growth of plants in the presence of Bartsia alpina (and potentially other hemiparasitic species) litter could have important implications for communities in which it occurs, including enhanced survival of seedlings of co-occurring species and increased resource patchiness.
We studied the population structure of a high arctic breeding wader bird species, the White-rumped Sandpiper Calidris fuscicollis. Breeding adults, chicks and juveniles were sampled at seven localities throughout the species' breeding range in arctic Canada in 1999. The mitochondrial control region was analysed by DNA sequencing, feathers were analysed for carbon isotope ratios (C-13/C-12) by isotope ratio mass spectrometry, and morphological measurements were analysed using principal component analyses. taking the effect of sex into account (identified by molecular genetic methods). In general. our results support the notion that the White-rumped Sandpiper is a monotypic species with no subspecies, and most of the morphological and genetic variation occurs within sites. Nevertheless. some differences between sites were found. Birds from the two northernmost sites (Ellesmere and Devon Islands) had relatively longer bill and wing and shorter tarsus than birds sampled further south, possibly reflecting genetic differences between populations. The carbon isotope ratios were higher at the easternmost site (Baffin Island), revealing differences in the isotope content of the food. The mtDNA sequences showed no significant differentiation between sites and no pattern of isolation-by-distance was found. Based on the mtDNA variation, the species was estimated to have a long-term effective population size of approximately 9,000 females. The species shows no clear evidence of any population expansion or decline. Our results indicate that carbon isotope ratios, and possibly also certain mtDNA haplotypes, may be useful as tools for identifying the breeding origin of White-rumped Sandpipers on migration and wintering sites.