Small-scale plant species distribution in snowbeds and its sensitivity to climate change

Alpine snowbeds are characterized by a long-lasting snow cover and low soil temperature during the growing season. Both these key abiotic factors controlling plant life in snowbeds are sensitive to anthropogenic climate change and will alter the environmental conditions in snowbeds to a considerable extent until the end of this century. In order to name winners and losers of climate change among the plant species inhabiting snowbeds, we analyzed the small-scale species distribution along the snowmelt and soil temperature gradients within alpine snowbeds in the Swiss Alps. The results show that the date of snowmelt and soil temperature were relevant abiotic factors for small-scale vegetation patterns within alpine snowbed communities. Species richness in snowbeds was reduced to about 50% along the environmental gradients towards later snowmelt date or lower daily maximum temperature. Furthermore, the occurrence pattern of the species along the snowmelt gradient allowed the establishment of five species categories with different predictions of their distribution in a warmer world. The dominants increased their relative cover with later snowmelt date and will, therefore, lose abundance due to climate change, but resist complete disappearance from the snowbeds. The indifferents and the transients increased in species number and relative cover with higher temperature and will profit from climate warming. The snowbed specialists will be the most suffering species due to the loss of their habitats as a consequence of earlier snowmelt dates in the future and will be replaced by the avoiders of late-snowmelt sites. These forthcoming profiteers will take advantage from an increasing number of suitable habitats due to an earlier start of the growing season and increased temperature. Therefore, the characteristic snowbed vegetation will change to a vegetation unit dominated by alpine grassland species. The study highlights the vulnerability of the established snowbed vegetation to climate change and requires further studies particularly about the role of biotic interactions in the predicted invasion and replacement process.     

Responses of flowering phenology of snowbed plants to an experimentally imposed extreme advanced snowmelt

In snowbed habitats, characterized by a long-lasting snow cover, the timing of snowmelt can be included among the major factors controlling plant phenology. Nevertheless, only a few ecological studies have tested the responses of flowering phenology of species growing in very late snow-free habitats to an advanced snowmelt (AS) date. The aim of this study was to determine the impacts of an extremely earlier melt-out of snow on flowering phenology of vascular plant species inhabiting an alpine snowbed. The study was conducted in the high Gavia Valley (Italy, 2,700 m a.s.l.). On 30th May 2012, we removed manually the snow cover and set up an experiment with 5 AS and 5 control plots. Phenological observations of the most abundant vascular species were conducted every 4–6 days. Moreover, we calculated cumulative soil temperature and recorded the mortality of reproductive structures of three species. For several species flowering occurred earlier, and the prefloration period was extended in the AS treatment in comparison with the control. For the majority of species, cumulative soil temperatures in the AS treatment and the control were comparable, confirming that temperature exerts the main control on the flowering of the species inhabiting snowbeds. Earlier flowering species resulted more affected by an AS date in comparison with later flowering species. The mortality of reproductive structures did not increase in the AS treatments in comparison with the control suggesting that few and weak frost events in late spring do not affect the survival of reproductive structures of the species studied.     

Foliar freezing resistance of Australian alpine plants over the growing season

We assessed the freezing resistance of leaves ex situ of 25 Australian alpine plant species. We compared the freezing resistance of forb, graminoid and shrub species from three alpine summits of different altitudes; from a low altitude site just above treeline, to a fully alpine tundra site. Foliar freezing resistance (LT₅₀) in spring varied from ?5.9°C to ?18.7°C and standardized LT₅₀ values within species were significantly related to site altitude. Additionally, when comparing all the species in the study, freezing resistance was significantly related to site; the LT₅₀ of samples from a low‐altitude summit (1696 m) were significantly lower than those of samples from mid‐ (1805 m) and high‐altitude (1860 m) summits. The LT₅₀ of juvenile foliage did not differ significantly from that of adult foliage. Shrubs were highly resistant to freezing. At the highest summit, we examined the course of seasonal freezing resistance from early summer to early autumn across three alpine plant communities that differed in the time of natural snowmelt; from sheltered (snowpatch) to exposed (open heath). No differences in freezing resistance over the growing season were detected for exposed or sheltered communities and there were no consistent trends indicating frost hardening over the growing season. Overall, the common Australian alpine species we investigated appear well adapted to freezing conditions throughout the snow‐free growing season. We have no evidence to suggest that freezing temperatures soon after snowmelt in spring are especially damaging to the alpine plants at these summits.     

Increased spring freezing vulnerability for alpine shrubs under early snowmelt

Alpine dwarf shrub communities are phenologically linked with snowmelt timing, so early spring exposure may increase risk of freezing damage during early development, and consequently reduce seasonal growth. We examined whether environmental factors (duration of snow cover, elevation) influenced size and the vulnerability of shrubs to spring freezing along elevational gradients and snow microhabitats by modelling the past frequency of spring freezing events. We sampled biomass and measured the size of Salix herbacea, Vaccinium myrtillus, Vaccinium uliginosum and Loiseleuria procumbens in late spring. Leaves were exposed to freezing temperatures to determine the temperature at which 50 % of specimens are killed for each species and sampling site. By linking site snowmelt and temperatures to long-term climate measurements, we extrapolated the frequency of spring freezing events at each elevation, snow microhabitat and per species over 37 years. Snowmelt timing was significantly driven by microhabitat effects, but was independent of elevation. Shrub growth was neither enhanced nor reduced by earlier snowmelt, but decreased with elevation. Freezing resistance was strongly species dependent, and did not differ along the elevation or snowmelt gradient. Microclimate extrapolation suggested that potentially lethal freezing events (in May and June) occurred for three of the four species examined. Freezing events never occurred on late snow beds, and increased in frequency with earlier snowmelt and higher elevation. Extrapolated freezing events showed a slight, non-significant increase over the 37-year record. We suggest that earlier snowmelt does not enhance growth in four dominant alpine shrubs, but increases the risk of lethal spring freezing exposure for less freezing-resistant species.     

Direct and indirect control by snow cover over decomposition in alpine tundra along a snowmelt gradient

We assessed direct and indirect effects of snow cover on litter decomposition and litter nitrogen release in alpine tundra. Direct effects are driven by the direct influence of snow cover on edaphoclimatic conditions, whereas indirect effects result from the filtering effect of snow cover on species’ abundance and traits. We compared the in situ decomposition of leaf litter from four dominant plant species (two graminoids, two shrubs) at early and late snowmelt locations using a two-year litter-bag experiment. A seasonal experiment was also performed to estimate the relative importance of winter and summer decomposition. We found that growth form (graminoids vs. shrubs) are the main determinants of decomposition rate. Direct effect of snow cover exerted only a secondary influence. Whatever the species, early snowmelt locations showed consistently reduced decomposition rates and delayed final stages of N mineralization. This lower decomposition rate was associated with freezing soil temperatures during winter. The results suggest that a reduced snow cover may have a weak and immediate direct effect on litter decomposition rates and N availability in alpine tundra. A much larger impact on nutrient cycling is likely to be mediated by longer term changes in the relative abundance of lignin-rich dwarf shrubs.     

Arthropod community composition along snowmelt gradients in snowbeds in the Snowy Mountains of south‐eastern Australia

Environmental gradients drive variation in community composition across a range of spatial scales. In alpine regions, areas of long‐lasting snow (‘snow patches’) create snowmelt gradients that drive considerable change in vegetation structure and composition over small spatial scales. This study examined whether there is parallel variation in arthropod communities using snowmelt gradients in the Australian Alps. Mites (Acarina) were the most common arthropods in snow patches, followed by springtails while, among the insects, the orders Hymenoptera (primarily Formicidae), Diptera, Coleoptera (primarily Carabidae) and Hemiptera (primarily Cicadellidae) dominated. Along the snowmelt gradient, arthropod assemblages changed from having equal proportions of predators and herbivores in early‐melting zones to being predator‐dominated in late‐melting zones, particularly early in the growing season. This followed a transition in vegetation cover and composition and was driven by higher numbers of predacious carabid beetles in later‐melting zones. Overall, however, our results suggest that snowbed arthropod communities in the Australian alpine zone are more sensitive to short‐term effects, such as time since snowmelt, than to differences in vegetation structure and composition or long‐term patterns of snowmelt. Continued advancement of snowmelt timing due to warmer spring temperatures is therefore likely to have more impact on the seasonality of snowbed arthropod communities than on the overall community composition.     

Evergreen alpine shrubs have high freezing resistance in spring,irrespective of snowmelt timing and exposure to frost: an investigation from the Snowy Mountains,Australia

Over winter, alpine plants are protected from low-temperature extremes by a blanket of snow. Climate change predictions indicate an overall reduction in snowpack and an earlier thaw; a situation which could expose the tips of shrubs which extend above the snowpack to freezing events in early spring, and cause foliar frost damage during the onset of physiological activity. We assessed the photosynthetic responses of freezing-damaged shrub leaves from an assay of freezing temperatures in the Snowy Mountains in south-eastern Australia, using chlorophyll fluorometery ex situ. We sampled leaves that were exposed early during the spring thaw and leaves that were buried in snow for up to two extra weeks, from four evergreen shrub species at monthly intervals following the period of snowmelt. Freezing resistance (estimated from LT₅₀) was poorest at the earliest spring sampling time, in both exposed above-snow and protected below-snow foliage in all species. Protected foliage in early spring had lower freezing resistance than exposed foliage, but not significantly so. By the third sampling time, freezing resistance was significantly better in the lower protected foliage (LT₅₀ of ? 14) compared with the upper exposed foliage (LT₅₀ of ? 10) in one species. Over the course of spring, freezing resistance improved significantly in all species, with LT₅₀ values of between ? 10 and ? 15 °C by the third sampling time, which is lower than the minimum air temperatures recorded at that time (> ? 5 °C). The results indicate that the dominant evergreen shrub species in this area may only be susceptible to freezing events very early in spring, before a period of frost-hardening occurs after snowmelt. Later in spring, these alpine shrubs appear frost hardy, thus further perpetuating the positive feedbacks surrounding shrub expansion in alpine areas.     

Advanced snowmelt causes shift towards positive neighbour interactions in a subarctic tundra community

Positive and negative species interactions are important factors in structuring vegetation communities. Studies in many ecosystems have focussed on competition; however, facilitation has often been found to outweigh competition under harsh environmental conditions. The balance between positive and negative species interactions is known to shift along spatial, temporal and environmental gradients and thus is likely to be affected by climate change. Winter temperature and precipitation patterns in Interior Alaska are rapidly changing and could lead to warmer winters with a shallow, early melting snow cover in the near future. We conducted snow manipulation and neighbour removal experiments to test whether the relative importance of positive and negative species interactions differs between three winter climate scenarios in a subarctic tundra community. In plots with ambient, manually advanced or delayed snowmelt, we assessed the relative importance of neighbours for survival, phenology, growth and reproduction of two dwarf shrub species. Under ambient conditions and after delayed snowmelt, positive and negative neighbour effects were generally balanced, but when snowmelt was advanced we found overall facilitative neighbour effects on survival, phenology, growth and reproduction of Empetrum nigrum, the earlier developing of the two target species. As earlier snowmelt was correlated with colder spring temperatures and a higher number of frosts, we conclude that plants experienced harsher environmental conditions after early snowmelt and that neighbours could have played an important role in ameliorating the physical environment at the beginning of the growing season.     

Demography of an endangered endemic rupicolous cactus

The snow cover extent is an important factor for the structure and composition of arctic and alpine tundra communities. Over the last few decades, snowmelt in many arctic and alpine regions has advanced, causing the growing season to start earlier and last longer. In a field experiment in subarctic tundra in Interior Alaska, I manipulated the timing of snowmelt and measured the response in mortality, phenology, growth, and reproduction of the eight dominant plant species. I then tested whether the phenological development of these species was controlled by snowmelt date or by temperature (in particular growing degree days, GDD). In order to expand our understanding of plant sensitivity to snowmelt timing, I explored whether the response patterns can be generalized with regard to the temporal niche of each species. Differences in the phenology between treatments were only found for the first stages of the phenological development (=phenophases). The earlier the temporal niche (i.e., the sooner after snowmelt a species develops) the more its phenology was sensitive to snowmelt. Later phenophases were mostly controlled by GDD, especially in late-developing species. In no species did an earlier snowmelt and a longer growing season directly enhance plant fitness or fecundity, in spite of the changes in the timing of plant development. In conclusion, the temporal niche of a species’ phenological development could be a predictor of its response to snowmelt timing. However, only the first phenophases were susceptible to changes in snowmelt, and no short-term effects on plant fitness were found.     

长白山高山苔原雪斑大白花地榆群落土壤氮素动态与生产力的关系

在高山苔原冬季积雪覆盖的群落生长季短, 但明显比周围群落生长茂盛。为了说明雪斑地段群落生长机理, 对长白山苔原雪斑土壤氮素动态以及大白花地榆(Sanguisorba sitchensis)群落生产力进行了连续测定。雪斑群落土壤冬季相对温暖, 最低日平均温度-1.4 ℃, 裸露地段-16.9 ℃, 全年水分条件充足; 积雪期凋落物分解和氮矿化均在进行, 土壤具有很高的氮素含量及矿化速率。大白花地榆地上部分净初级生产力为4 046 kg·hm^-2·a^-1。正是独特的水热条件和养分条件, 以及具有很大的叶面积同化器官, 高山苔原雪斑地段的大白花地榆群落才得以维持生存并表现出很高的生产力水平。     

Intraspecific variations in seedling emergence and survival of Potentilla matsumurae (Rosaceae) between alpine fellfield and snowbed habitats

Potentilla matsumurae has a wide distribution from wind-blown fellfields to snowbeds in alpine regions of Japan. The environmental factors influencing seedling establishment differ between the fellfield and snowbed habitats; plants growing in each habitat may therefore have different germination strategies. Using a reciprocal sowing experiment, patterns of seedling emergence and survivorship were examined in both habitat types in the Taisetsu Mountains, Japan. Seeds derived from a fellfield population germinated earlier than did those derived from a snowbed population at both habitats, and the germination of fellfield seeds continued throughout the growing season. The timing of seedling emergence greatly affected subsequent survival at the fellfield. Seedlings that emerged in the first half of the growing season had low survivorship during the first year because of frost and drought damage, but the remaining seedlings had high survivorship during the winter; seedlings that emerged in the latter half of the growing season showed the opposite trend. At the snowbed, seedling survival was high throughout the growing season. Germination experiments in the laboratory highlighted a difference in the sensitivity of seeds from the fellfield and snowbed populations to fluctuating temperatures. These results indicate that intraspecific variation in emergence and survivorship may occur over a small scale in an alpine environment.     

Habitat-specific responses in the flowering phenology and seed set of alpine plants to climate variation: implications for global-change impacts

The timing of the snowmelt is a crucial factor in determining the phenological schedule of alpine plants. A long-term monitoring of snowmelt regimes in a Japanese alpine area revealed that the onset of the snowmelt season has been accelerated during the last 17 years in early snowmelt sites but that such a trend has not been detected in late snowmelt sites. This indicates that the global warming effect on the snowmelt pattern may be site-specific. The flowering phenology of fellfield plants in an exposed wind-blown habitat was consistent between an unusually warm year (1998) and a normal year (2001). In contrast, the flowering occurrence of snowbed plants varied greatly between the years depending on the snowmelt time. There was a large number of flowering species in the fellfield community from mid- to late to late June and from mid- to late July. The flowering peak of an early-melt snowbed plant community was in the middle of the flowering season and that of a late-melt snowbed community was in the early flowering season. These habitat-specific phenological patterns were consistent between 1998 and 2001. The effects of the variation in flowering timing on seed-set success were evaluated for an entomophilous snowbed herb, Peucedanum multivittatum, along the snowmelt gradient during a 5-year period. When flowering occurred prior to early August, mean temperature during the flowering season positively influenced the seed set. When flowering occurred later than early August, however, the plants enjoyed high seed-set success irrespective of temperature conditions if frost damage was absent. These observations are probably explained based on the availability of pollinators, which depends not only on ambient temperature but also on seasonal progress. These results suggest that the effects of climate change on biological interaction may vary depending on the specific habitat in the alpine ecosystem in which diverse snowmelt patterns create complicated seasonality for plants within a very localized area.     

高山林线交错带高山杜鹃的凋落物分解

凋落物分解是维持生态系统生产力、养分循环、土壤有机质形成的关键生态过程。高山林线交错带是陆地生态系统中对气候变化响应的敏感区域。季节变化和海拔梯度上的植被类型差异可能会影响该区域凋落物的分解,进而对高山生态系统的碳氮循环产生重要影响。采用凋落物分解袋的方法,研究了川西高山林线交错带优势种高山杜鹃(Rhododendron lapponicum)凋落叶在雪被期和生长季的分解特征。结果显示:(1)季节变化和植被类型对高山杜鹃凋落物的分解均具有显著影响(P0.05),凋落叶的质量损失主要发生在生长季且在高山林线最大,暗针叶林中雪被期的质量损失略高于生长季,但差异不显著;(2)林线交错带上高山杜鹃凋落叶分解缓慢,一年干物质失重率为9.62%,拟合分解系数k为0.145;(3)高山杜鹃凋落叶的质量变化主要体现在纤维素降解显著且集中在雪被期,木质素无明显降解,在高山林线上C/N、C/P、木质素/N变化幅度较小且C、N、P的释放表现得稳定而持续。结果表明,季节性雪被对林线交错带内高山杜鹃分解的影响不仅局限在雪被期内,雪被融化期间频繁的冻融作用和雪融水淋洗作用可能会促进高山杜鹃凋落物在生长季初期的分解。总的来看,在气候变暖的情景下,雪被的缩减、生长季的延长和高山杜鹃群落的扩张可能加速高山林线交错带高山杜鹃凋落物的分解。     

Will loss of snow cover during climatic warming expose New Zealand alpine plants to increased frost damage?

If snow cover in alpine environments were reduced through climatic warming, plants that are normally protected by snow-lie in winter would become exposed to greater extremes of temperature and solar radiation. We examined the annual course of frost resistance of species of native alpine plants from southern New Zealand that are normally buried in snowbanks over winter (Celmisia haastii and Celmisia prorepens) or in sheltered areas that may accumulate snow (Hebe odora) and other species, typical of more exposed areas, that are relatively snow-free (Celmisia viscosa, Poa colensoi, Dracophyllum muscoides). The frost resistance of these principal species was in accord with habitat: those from snowbanks or sheltered areas showed the least frost resistance, whereas species from exposed areas had greater frost resistance throughout the year. P. colensoi had the greatest frost resistance (−32.5°C). All the principal species showed a rapid increase in frost resistance from summer to early winter (February–June) and maximum frost resistance in winter (July–August). The loss of resistance in late winter to early summer (August–December) was most rapid in P. colensoi and D. muscoides. Seasonal frost resistance of the principal species was more strongly related to daylength than to temperature, although all species except C. viscosa were significantly related to temperature when the influence of daylength was accounted for. Measurements of chlorophyll fluorescence indicated that photosynthetic efficiency of the principal species declined with increasing daylength. Levels of frost resistance of the six principal alpine plant species, and others measured during the growing season, were similar to those measured in tropical alpine areas and somewhat more resistant than those recorded in alpine areas of Europe. The potential for frost damage was greatest in spring. The current relationship of frost resistance with daylength is sufficient to prevent damage at any time of year. While warmer temperatures might lower frost resistance, they would also reduce the incidence of frosts, and the incidence of frost damage is unlikely to be altered. The relationship of frost resistance with daylength and temperature potentially provides a means of predicting the responses of alpine plants in response to global warming.     

Site-specific factors influence the richness and phenology of snowbed plants in the Pyrenees

Although the timing of snowmelt and growth temperatures appear to be the main factors that influence the species richness and phenology of snowbed plants, site-specific characteristics may also play a role in modifying the effects of the timing of snowmelt and temperature. In this study, the effects of site-specific factors (microtopography and snow origin) on species richness and plant phenology were evaluated in 72 plots in two snowbeds in the Andorran Pyrenees. Snowmelt patterns influenced the spatial distribution of species richness and abundance. Site-specific factors had significant effects on the responses of species (shortening or lengthening the duration of the phenophase) and on the extent to which the timing of snowmelt influenced leaf expansion and flowering. Notably, the highest rates of leaf expansion occurred on late snowmelt isoclines, where, nevertheless, the time taken to reach peak flowering was significantly longer than on the early snowmelt isoclines. The results of this study highlight the fact that, in addition to the effects of interannual variability in climate, site-specific factors have a significant effect on the phenology and reproductive success of the commonest plants in the snowbed communities of the Pyrenees.     

Snowbeds are more affected than other subalpine–alpine plant communities by climate change in the Swiss Alps

While the upward shift of plant species has been observed on many alpine and nival summits, the reaction of the subalpine and lower alpine plant communities to the current warming and lower snow precipitation has been little investigated so far. To this aim, 63 old, exhaustive plant inventories, distributed along a subalpine–alpine elevation gradient of the Swiss Alps and covering different plant community types (acidic and calcareous grasslands; windy ridges; snowbeds), were revisited after 25–50 years. Old and recent inventories were compared in terms of species diversity with Simpson diversity and Bray–Curtis dissimilarity indices, and in terms of community composition with principal component analysis. Changes in ecological conditions were inferred from the ecological indicator values. The alpha‐diversity increased in every plant community, likely because of the arrival of new species. As observed on mountain summits, the new species led to a homogenization of community compositions. The grasslands were quite stable in terms of species composition, whatever the bedrock type. Indeed, the newly arrived species were part of the typical species pool of the colonized community. In contrast, snowbed communities showed pronounced vegetation changes and a clear shift toward dryer conditions and shorter snow cover, evidenced by their colonization by species from surrounding grasslands. Longer growing seasons allow alpine grassland species, which are taller and hence more competitive, to colonize the snowbeds. This study showed that subalpine–alpine plant communities reacted differently to the ongoing climate changes. Lower snow/rain ratio and longer growing seasons seem to have a higher impact than warming, at least on plant communities dependent on long snow cover. Consequently, they are the most vulnerable to climate change and their persistence in the near future is seriously threatened. Subalpine and alpine grasslands are more stable, and, until now, they do not seem to be affected by a warmer climate.     

THE ANNUAL CARBOHYDRATE CYCLE OF ALPINE PLANTS AS RELATED TO GROWTH

Mooney, H. A., and W. D. Billings. (Duke U., Durham, N. C.) The annual carbohydrate cycle of alpine plants as related to growth. Amer. Jour. Bot. 47(7): 594–598. Illus. 1960.—Analyses were made of the carbohydrate content of roots, rhizomes, and shoots of certain plants in the alpine tundra region of the Medicine Bow Mountains, Wyoming, from the beginning of one growing season until the start of the following one. Principal species investigated were Saxifraga rhomboidea, Polygonum bistortoides, and Geum turbinatum. Results were correlated with phenological events as observed in the field. Growth of these plants was found to be very rapid, commencing in some instances under a cover of old snow. The underground organs contained relatively large amounts of carbohydrate reserves. A great part of this stored carbohydrate was utilized in growth prior to snowmelt and during the grand period of shoot growth immediately following snowmelt. In Polygonum, 50% of the rhizome reserves was used in a 1-wk. period in early growth. Except for this short period of rapid depletion in rhizomes and roots during early growth, high carbohydrate levels were maintained both in the shoot and in the underground organs during most of the growing season. Generally, the lowest carbohydrate reserve level in both root and shoot occurred before flowering, a relatively high level was maintained in the shoot from flowering until after fruiting, while peak storage in underground parts was reached at the start of fall dormancy. The carbohydrate cycle in these alpine plants is quite similar to that in certain arctic plants.     

Substantial compositional turnover of fungal communities in an alpine ridge‐to‐snowbed gradient

The main gradient in vascular plant, bryophyte and lichen species composition in alpine areas, structured by the topographic gradient from wind‐exposed ridges to snowbeds, has been extensively studied. Tolerance to environmental stress, resulting from wind abrasion and desiccation towards windswept ridges or reduced growing season due to prolonged snow cover towards snowbeds, is an important ecological mechanism in this gradient. The extent to which belowground fungal communities are structured by the same topographic gradient and the eventual mechanisms involved are less well known. In this study, we analysed variation in fungal diversity and community composition associated with roots of the ectomycorrhizal plant Bistorta vivipara along the ridge‐to‐snowbed gradient. We collected root samples from fifty B. vivipara plants in ten plots in an alpine area in central Norway. The fungal communities were analysed using 454 pyrosequencing analyses of tag‐encoded ITS1 amplicons. A distinct gradient in the fungal community composition was found that coincided with variation from ridge to snowbeds. This gradient was paralleled by change in soil content of carbon, nitrogen and phosphorus. A large proportion (66%) of the detected 801 nonsingleton operational taxonomic units (OTUs) were ascomycetes, while basidiomycetes dominated quantitatively (i.e. with respect to number of reads). Numerous fungal OTUs, many with taxonomic affinity to Sebacinales, Cortinarius and Meliniomyces, showed distinct affinities either to ridge or to snowbed plots, indicating habitat specialization. The compositional turnover of fungal communities along the gradient was not paralleled by a gradient in species richness.     

Il Contributo di Maria Selebam de Cattani Agli Studi Delle Alghe Marine e di Certe Sue Raccolte Conservate a Venezia

Abstract

Phytosociology, ecology and chorology of snowbed vegetation in Croatia were researched. Snowbed stands, found only in freezing ravines and dolines of the Liburnian karst and Velebit Mountains (NW Dinaric Alps, NW Croatia), were classified into the association Drepanoclado-Heliospermetum (Salicion retusae, Arabidetalia caeruleae, Thlaspietea rotundifolii). Those stands, exposed mostly to the north and shaded for the majority of the growing season, were developed on more or less settled periglaciar screes and boulders in the coldest parts (bottoms, smaller ditches) of the dolines with long-lasting snow cover. Due to lower altitude of Croatian mountains, snowbed vegetation could be found only azonally, in freezing ravines surrounded by altimontane and subalpine (fir-)beech and spruce forests, where these stands manage to thrive due to specific microclimatic conditions. Snowbeds host some rare, endangered and/or protected plant species in Croatia. Vulnerability of the flora and vegetation of snowbeds in Croatia is briefly discussed.     

Counterbalancing effects of competition for resources and facilitation against grazing in alpine snowbed communities

Alpine snowbeds are habitats where the major limiting factors for plant growth are herbivory and a small time window for growth due to late snowmelt. Despite these limitations, snowbed vegetation usually forms a dense carpet of palatable plants due to favourable abiotic conditions for plant growth within the short growing season. These environmental characteristics make snowbeds particularly interesting to study the interplay of facilitation and competition. We hypothesised an interplay between resource competition and facilitation against herbivory. Further, we investigated whether these predicted neighbour effects were species‐specific and/or dependent on ontogeny, and whether the balance of positive and negative plant–plant interactions shifted along a snowmelt gradient. We determined the neighbour effects by means of neighbour removal experiments along the snowmelt gradient, and linear mixed model analyses. The results showed that the effects of neighbour removal were weak but generally consistent among species and snowmelt dates, and depended on whether biomass production or survival was considered. Higher total biomass and increased fruiting in removal plots indicated that plants competed for nutrients, water, and light, thereby supporting the hypothesis of prevailing competition for resources in snowbeds. However, the presence of neighbours reduced herbivory and thereby also facilitated survival. For plant growth the facilitative effects against herbivores in snowbeds counterbalanced competition for resources, leading to a weak negative net effect. Overall the neighbour effects were not species‐specific and did not change with snowmelt date. Our finding of counterbalancing effects of competition and facilitation within a plant community is of special theoretical value for species distribution models and can explain the success of models that give primary importance to abiotic factors and tend to overlook interrelations between biotic and abiotic effects on plants.