Large herbivore grazing and invertebrates in an alpine ecosystem

The literature on how plants respond to grazing and other disturbance factors have advanced greatly in recent decades, but studies of invertebrates are comparably few. We here quantify the effects of 3 levels of sheep grazing on selected invertebrates in an alpine ecosystem in Norway. We tested the hypothesis that invertebrates are more sensitive to grazing than plants (responding mainly at high density), and that primary consumers (herbivorous beetles) are more sensitive than predatory species (beetles and spiders). We captured 1218 specimens belonging to 44 beetle species and 6672 specimens belonging to 66 species of spiders. The community was dominated by few species: 5 beetle and 3 spider species made up 53.0% and 37.4% of the catch, respectively. At the local (plot) scale, most negative responses were only recorded at high sheep density, and invertebrates were thus not more responsive than the plant community. Two dominant herbivorous beetles responded to grazing while 3 dominant species of predatory beetles did not (one marginally). Spider species richness and frequency of occurrence of 2 dominant species were negatively affected by sheep grazing, suggesting variation between different taxonomic groups of predators. Further functional details than simple classification like herbivorous, predatory and litter-dwelling invertebrates seem to be required before a framework to predict responses to disturbance are robust.     

Temporal variation in density dependent body growth of a large herbivore

Temporal variation both due to density dependent and density independent processes affect performance and vital rates in large herbivores. Annual fluctuations in climate affect foraging conditions and thus body growth of large herbivores during the short growing season in alpine habitats. Also, high animal densities on summer ranges may increase competition for food and reduce body mass gain. Yet, little is known about interactive effects of density and climate on alpine summer ranges, and the time scales these processes operate on. In this fully replicated landscape‐scale experiment, we kept domestic sheep at high and low densities over nine grazing seasons in an alpine habitat, and tested the relative role of density and annual variation for lamb body mass gain during summer and whether effects of density and annual variation interacted. We found that lambs at high density gained less mass over the summer season than lambs at low density. At short time scales the density effect interacted with annual fluctuations in body growth. We documented a long‐term temporal trend in body mass, consistent with the hypothesis that grazing effects affect habitat differentially at high and low density over years. At high density lamb autumn body mass declined during the first three grazing seasons and then stabilized, whereas body mass slightly increased over years at low density. This long‐term trend suggests accumulative density dependent effects from a biomass or quality reduction, and hence delayed food competition at high density and possibly facilitation at low density. Our experiment provides new insight into how density dependent effects on performance of a large herbivore depend on temporal scale of observation.     

Grazing responses in herbs in relation to herbivore selectivity and plant traits in an alpine ecosystem

Herbivores shape plant communities through selective foraging. However, both herbivore selectivity and the plant’s ability to tolerate or resist herbivory may depend on the density of herbivores. In an alpine ecosystem with a long history of grazing, plants are expected to respond to both enhanced and reduced grazing pressures, and the interaction between plant traits and changes in species abundance are expected to differ between the two types of alteration of grazing regime. To understand the mechanisms behind species response, we investigated the relationship between sheep selectivity (measured in situ), plant traits and experimentally derived measures of change in species abundance as a response to the enhancement (from low to high density) or cessation (from low to zero density) of sheep grazing pressure over a six-year time period for 22 abundant herb species in an alpine habitat in south Norway. Sheep selected large, late-flowering herbs with a low leaf C/N ratio. Species that increased in abundance in response to enhanced grazing pressure were generally small and had high root/shoot ratios, thus exhibiting traits that reflect both resistance (through avoidance) and tolerance (through regrowth capacity) strategies. The abundance of selected species remained stable during the study period, and also under the enhanced grazing pressure treatment. There was, however, a tendency for selected species to respond positively to cessation of grazing, although overall responses to cessation of grazing were much less pronounced than responses to enhanced grazing. Avoidance through short stature (probably associated with increased light availability through the removal of tall competitors) as well as a certain amount of regrowth capacity appear to be the main mechanisms behind a positive response to enhanced grazing pressure in this study. The plant trait perspective clearly improves our insight into the mechanisms behind observed changes in species abundance when the disturbance regime is altered. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Seedling demography in an alpine ecosystem

Seedling establishment has long been believed to be rare on alpine tundra because of predicted life history trade-offs, the clonality of alpine species, and the harshness of the alpine climate. Contrary to the idea that seedlings are rare on alpine tundra, a 4-yr demographic study of seedlings at Niwot Ridge, Colorado, USA, found seedlings at high densities, particularly in wetter plant communities. Higher germination densities were associated with higher soil moistures both across communities and across time. Mortality of seedlings was highest in the first year and decreased in subsequent years. Species' abundances differed between seedling and adult populations. Many forbs that lacked vegetative reproduction were significantly more abundant among seedling populations, and many monocots and clonal forbs were more abundant among adult populations. In a comparison with published demographic rates, seedling recruitment and mortality rates of Niwot Ridge species fell above or within rates for a wide range of perennial species. Therefore, germination and seedling establishment stages are no more limiting to sexual reproduction in alpine plants than in other perennial plants.     

Density dependent and temporal variability in habitat selection by a large herbivore; an experimental approach

Both density dependent and density independent processes such as climate affect population dynamics in large herbivores. Understanding herbivore foraging patterns is essential to identify the underlying mechanisms behind variation in vital rates. However, very little is known about how animals vary their selection of habitat temporally, alone or in interaction with density during summer. At the foraging scale, we tested using a fully replicated experiment whether domestic sheep Ovis aries stocked at high (80  per  km²) and low (25  per  km²) densities (spatial contrasts) varied their habitat selection temporally over a four year period. We predicted reduced selection of high productivity vegetation types with increasing density, and that seasonal and annual variation in climate would affect this density dependent selection pattern by increasing competition for high quality habitats in late grazing season and in years with poor vegetation development and over time related to vegetation responses to grazing. As predicted from the Ideal free distribution model, selection of high productivity habitat decreased at high density. There was also a marked temporal variation in habitat selection. Selection of the most productive vegetation types declined towards the end of each grazing season, but increased over years both at low and high sheep density. There was only weak evidence for interactions, as selection ratio of highly productive habitats tended to increase more over years at low density as compared to high density. Limited interactive effects of density and annual variation on habitat selection during summer may explain why similar interactions in vital rates have rarely been reported for summer seasons. Our results are consistent with the view that variation in habitat selection is a central mechanism for climate and density related variation in vital rates.     

Age class,density and temporal effects on diet composition of sheep in an alpine ecosystem

Understanding diet selection is important since diet determines energy intake and therefore growth of ungulate populations. Yet very few studies have reported annual variation in diet. Density-dependent diet choice by large herbivores has been reported several times, but these studies are typically either short-term or they lack replication of the density treatment. In a landscape-scale experiment with 3 replicates of two densities (25 and 80 individuals/km²) of domestic sheep, we determined diet composition using microhistological analysis during 6 summer grazing seasons (2002–2007) in alpine habitats. We tested how age class, density and temporal variation (within season, annually, and over years) affected summer diet. There was marked evidence of additive effects of these factors on overall diet composition, but interactions were few. The interaction between density and annual variation was an important determinant of the proportion of the main forage component (Avenella flexuosa), but not of the proportions of herbs, Salix spp. and for “other” forage plants. Surprisingly, the density effect on this intermediate quality forage (A. flexuosa) was not consistent among years (both positive, negative and no effects), likely arising due to large variation in the proportion of the other forage plants. We discuss how foraging ecology can supplement the insight from life history theory in explaining variation in vital rates.     

Long-term behaviour of 15N in an alpine grassland ecosystem

Nitrogen dynamics in semi-natural environments is crucial for the development and ecological stability of these systems. The present paper shows the results of the reinvestigation of a ¹⁵N-tracer experiment, which was established in the Grossglockner massif in Austria at 2300ma.s.l. in 1974/1975. We show that large quantities of nitrogen introduced by a single pulse labelling (amounting to approximately 1.7% of the nitrogen in the system) into an alpine grassland remain in the soil–plant system, with only 55% being lost during 27–28 years. In the first 10cm of the four investigated soil profiles 40% of ¹⁵N was recovered, being mainly bound in organic forms. A simple site specific model was established on the basis of the results considering a biological, residual and labile N-pool, the latter being the source for N-losses. By the model a long mean residence time close to 100 years was derived for the remaining ¹⁵N.     

The pattern of density dependent growth inhibition in murine fibroblasts

Observations on the pattern of nuclear incorporation of ³H-TdR in long term (8-day) and short term (3-day) 3T3 cultures with local cell densities between 0.2 × 10⁴ and 6.2 × 10⁴ cells/cm² are reported. Contrary to a number of previous studies our observations indicate that density dependent inhibition is exhibited in relatively sparse cultures, commencing at 0.5 × 10⁴ cells/cm². Various possible mechanisms which could have caused the observed pattern of density-dependent regression in labelling index are discussed.     

Spatial analysis of density dependent pattern in coniferous forest stands

An investigation of spatial pattern in relatively sparse Pinus ponderosa-P. Jeffreyi stands showed that a simple Poisson model of random distribution described the pattern at 5 to 50 m scales in the denser stands examined when allowance is made for inhibition between nearest neighbors. There is evidence for a clumped distribution in large quadrats for the sparsest stands, which concurs with prior work where a mixed Poisson model was fit to the data. The technique used was innovative in that it involved digitally recording tree locations from high resolution aerial photos, which allowed for the automatic application of several statistical techniques in order to determine how pattern varies with plot density and scale. Point locations were recorded for six 11.3 ha plots in three density regions of a 340 ha study area in northeastern California, USA. The inter-event distance distribution, and one- and two-dimensional power spectra were calculated, and variable quadrat analysis was performed for the data sets. The second order and spectral analyses showed no evidence of a distinctive clumped pattern at any scale, and all analyses showed that the pattern was regular at the scale of the average inter-plant distance in the denser stands. For the sparser stands, the counts in large quadrats did not fit a Poisson distribution, but were better fit by a mixed Poisson model describing aggregated pattern.     

Performance of two alpine plant species along environmental gradients in an alpine meadow ecosystem in central Tibet

Understanding how biotic interactions and abiotic conditions affect plant performance is important for predicting changes in ecosystem function and services in variable environments. We tested how performances of Astragalus rigidulus and Potentilla fruticosa change along gradients of biotic interactions (represented by plant species richness, abundance of the dominant plant species Kobresia pygmaea, and herbivory intensity) and abiotic conditions (represented by elevation, aspect, and slope steepness) across a semi-arid landscape in central Tibet. Redundancy analyses showed that the biotic variables explained 30 and 39 % of the variation in overall performance of A. rigidulus (P = 0.03) and P. fruticosa (P = 0.01), respectively. Abiotic variables did not contribute significantly to variation among A. rigidulus populations. Plant size decreased with species richness in both species and was larger on south- rather than north-facing slopes. Reproductive effort for both species was significantly negatively related to the abundance of K. pygmaea and both species had larger reproductive effort on south- rather than north- and west-facing slopes. The proportion of biomass allocated to sexual reproduction in P. fruticosa was negatively correlated with K. pygmaea abundance and herbivory intensity. The population density of P. fruticosa was positively related to elevation, species richness, and K. pygmaea abundance. We conclude that plant performance at a local scale was more strongly related to biotic than abiotic conditions, but different components of plant performance responded differently to predictor variables and the responses were species-specific. These findings have important implications for rangeland management under changing environmental conditions.     

Seasonal shifts in habitat selection of a large herbivore and the influence of human activity

Failure to recognize factors contributing to variation in habitat models like resource selection functions (RSFs) can affect their application for projecting probabilities of occurrence, and thereby limit their relevance for conservation and management. We compared seasonal RSFs (2006–2008) for 16 adult female moose (Alces alces) with home ranges located in western Algonquin Provincial Park (APP), Ontario, Canada, to those of 14 adult females located in provincial Wildlife Management Unit (WMU) 49, 40 km west of the protected area. Wildlife and habitat management practices differed between regions: hunting was higher in WMU 49 compared to APP, and APP preserved large tracts of old growth forest rarely found in WMU 49. Seasonal RSFs projected expected similarities in moose resource use between regions (e.g., responses to wetlands and stands of eastern hemlock, Tsuga canadensis [in winter]); however, we also observed differences consistent with the hypothesis that animals, through effects of hunting, would shift habitat use seasonally and in response to roads. We further observed evidence of functional responses in habitat selection due to underlying differences in forestry practices (e.g., responses to stands of old-growth hemlock forest). Given the close proximity and shared biogeographic region between study areas, we believe that observed spatial dynamics in RSFs were ultimately reflective of divergent management strategies between areas and ensuing differences in predation and hunting mortality risk, and functional habitat.     

Vulnerability of phenological synchrony between plants and pollinators in an alpine ecosystem

The relationship between flowering phenology and abundance of bumble bees (Bombus spp.) was investigated using 2 years of phenological data collected in an alpine region of northern Japan. Abundance of Bombus species was observed along a fixed transect throughout the flowering season. The number of flowering species was closely related to the floral resources for pollinators at the community scale. In the year with typical weather, the first flowering peak corresponded to the emergence time of queen bees from hibernation, while the second flowering peak corresponded to the active period of worker bees. In the year with an unusually warm spring, however, phenological synchrony between plants and bees was disrupted. Estimated emergence of queen bees was 10 days earlier than the first flowering date owing to earlier soil thawing and warming. However, subsequent worker emergence was delayed, indicating slower colony development. The flowering season finished 2 weeks earlier in the warm-spring year in response to earlier snowmelt. A common resident species in the alpine environment, B. hypocrita sapporoensis, flexibly responded to the yearly fluctuation of flowering. In contrast, population dynamics of other Bombus species were out of synchrony with the flowering: their frequencies were highest at the end of the flowering season in the warm-spring year. Therefore, phenological mismatch between flowers and pollinators is evident during warm years, which may become more prevalent in a warmer climate. To understand the mechanism of phenological mismatch in the pollination system of the alpine ecosystem, ground temperature, snowmelt regime, and life cycle of pollinators are key factors.     

Enhanced fitness and greater herbivore resistance: implications for dandelion invasion in an alpine habitat

Several hypotheses have been proposed to explain the defense strategies of invasive plants in new ranges. In the absence of specialist herbivores, it is believed that invasive plants may allocate fewer resources to resistance and more to growth and reproduction, thus increasing tolerance to damage in the invasive genotypes. In order to test these predictions, we compared both performance (growth and reproduction) and defense strategies (tolerance and resistance) of two populations of Taraxacum officinale, one from the native range in the French Alps, and one from the introduced range in the Chilean Andes. Individuals from the introduced population demonstrated improved reproductive traits relative to those from the native population, although there was no discernible difference in biomass accumulation. Additionally, reduced tolerance was evident in the case of the former; whereas fitness traits of native plants were unaffected by damage, invasive plants reduced growth and seed output by 25 and 30% respectively following damage treatments. Increases in levels of phenols and anthocyanins, produced as a defense response to herbivory, were observed in introduced plants. Our results suggest that reallocation of resources to reproduction may be an important factor favouring invasive success of T. officinale in Chile, and that a higher investment in chemical resistance traits in this population may also be a factor in this regard.     

Vegetation pattern formation in a fog-dependent ecosystem

Vegetation pattern formation is a striking characteristic of several water-limited ecosystems around the world. Typically, they have been described on runoff-based ecosystems emphasizing local interactions between water, biomass interception, growth and dispersal. Here, we show that this situation is by no means general, as banded patterns in vegetation can emerge in areas without rainfall and in plants without functional root (the Bromeliad Tillandsia landbeckii) and where fog is the principal source of moisture. We show that a simple model based on the advection of fog-water by wind and its interception by the vegetation can reproduce banded patterns which agree with empirical patterns observed in the Coastal Atacama Desert. Our model predicts how the parameters may affect the conditions to form the banded pattern, showing a transition from a uniform vegetated state, at high water input or terrain slope to a desert state throughout intermediate banded states. Moreover, the model predicts that the pattern wavelength is a decreasing non-linear function of fog-water input and slope, and an increasing function of plant loss and fog-water flow speed. Finally, we show that the vegetation density is increased by the formation of the regular pattern compared to the density expected by the spatially homogeneous model emphasizing the importance of self-organization in arid ecosystems.     

Threatened woody flora as an ecological indicator of large herbivore introductions

Rewilding and translocations of large herbivores for conservation purposes have increased in recent times, with numerous introductions inside and outside their native range. This study aims to analyze the use of threatened plant taxa as a possible ecological indicator of large herbivore introductions. We examined the effects of a threatened large ungulate, the Barbary sheep (Ammotragus lervia), on both endangered and vulnerable woody taxa after its introduction in 1970. Contrary to our hypothesis, the herbivore impact on threatened woody species was higher than that found on widespread woody plants. The results reveal that 35.7% of the threatened species showed the highest possible level of herbivore damage in contrast to 6.5% for the widespread species. Threatened species were preferred over common plants, probably due to their greater palatability. Overall plant cover, including neighboring species, was also an important factor determining browsing damage and, thus, habitats with low ground cover should be particularly considered in conservation plans. Herbivore damage on common taxa should be taken with caution since they could mask unsustainable herbivore densities for threatened woody taxa or protected habitats. The use of threatened woody taxa through the studied ecological indicators (herbivore damage, plant preferences, habitat use and regeneration success) represented a useful tool to assess the sustainability of large herbivores introductions and to establish a priority conservation ranking for threatened plant species. These findings highlight the deleterious effects of overabundant ungulate populations regardless its origin (exotic or native) and the need of monitoring threatened woody taxa to better estimate the suitability and sustainability of large herbivore introductions.     

Rapid environmental degradation in a subarctic ecosystem influences resource use of a keystone avian herbivore

1. Environmental degradation can change resource use strategies of animals and thereby affect survival and fitness. Arctic herbivores may be especially susceptible to the effects of such environmental change because their rapid growth rates demand high-quality forage, which may be limited as environmental conditions deteriorate. We studied the consequences of a trophic cascade, driven by Lesser Snow Goose (Chen caerulescens caerulescens) overgrazing on the south-west coast of Hudson Bay, Canada, which has caused tidal marsh (TM) degradation and the reduction in high-quality forage plants, on gosling growth and resource use. 2. We compared resource use and body size of goslings that inhabited tidal and freshwater marsh (FM) to determine how current foraging strategies influence growth and to test the hypothesis that during early growth goslings require and so consume high-quality TM plants, but that during later growth they may switch to foraging in lower-quality FM. 3. To investigate gosling resource use throughout growth, we measured once a week for 28 days the body size of goslings as well as stable isotope ratios (δ(34) S, δ(15) N and δ(13) C) in multiple tissues of goslings that were collected from both TM and nearby FM. We also measured the stable isotope ratios in forage plants sampled along transects and from gosling foreguts. We used an isotope-mixing model to determine the contribution of FM plants to gosling tissues. 4. Contrary to the proposed hypothesis, goslings inhabiting FM or TM primarily consumed FM plants during early growth. Furthermore, goslings that foraged extensively in FM had similar growth rates and grew to a similar size and body mass, as goslings that foraged in the degraded TM. However, goslings that currently inhabit freshwater or TM were significantly smaller than goslings that inhabited TM in the 1980s prior to habitat degradation. 5. Consequences of smaller overall body size include decreased survival and fecundity for arctic-nesting geese. The ability of phenotypically plastic responses to sustain persistence is limited by reaction norms and the extent of environmental change. Current research is assessing whether those limits have been reached in this system.     

Time-scale effects in the interaction between a large and a small herbivore

In the short term, grazing will mainly affect plant biomass and forage quality. However, grazing can affect plant species composition by accelerating or retarding succession at longer time-scales. Few studies concerning interactions among herbivores have taken the change in plant species composition into account. In a salt-marsh system, the long-term effects of exclusion of a large herbivore (cattle) on the abundance of a small herbivore (hare) were studied. Excluding cattle grazing for 30 years resulted in large changes in vegetation composition. In general, the cover of tall-growing species increased in the absence of cattle grazing. These long-term changes negatively affected hare grazing intensity. Hares preferentially fed on Festuca rubra and negatively selected tall growing plants, such as Elymus athericus, both in cattle-grazed and long-term ungrazed areas. However, the intensity of hare grazing was not related to the cover of F. rubra. The cover of tall-growing plants (E. athericus, Atriplex prostrata and Juncus maritimus) appeared to be the best predictor and hare grazing intensity decreased sharply with an increase of the cover of tall plants. When cover of tall plants did not increase, hare grazing intensity was not affected. The study shows that the time-scale of the experiment is of prime importance in studying interactions between herbivores. Species that do not seem to influence the abundance of one another or are competing for the same resources on a short time-scale might well be facilitating each other when looking at larger time-scales while taking plant species replacement into account.     

长白山高山冻原生态系统的碳储量

对长白山高山冻原生态系统碳在植被-凋落物-土壤中的分布进行了研究。结果表明,在长白山高山冻原植被亚系统中,碳储量分布规律是典型高山冻原(TT)>石质高山冻原(ST)>草甸高山冻原(MT)>沼泽高山冻原(WT)>石海高山冻原(RT),长白山高山冻原的植被碳储量为3.3457×104t。而生物量分布规律是WT>TT>MT>ST>RT。在长白山高山冻原凋落物亚系统中,平均凋落物量是0.991 kg.hm-2.年-1;凋落物碳储量为1.5043×104t,碳储量分布规律是TT>ST>MT>RT>WT。长白山高山冻原土壤(0~20 cm)亚系统中,有机碳储量为3.162×105t;每年约有1.4105×104t碳通过土壤呼吸释放到大气圈,长白山高山冻原植被-土壤系统现已成为3.6471×105t碳储库。