How extreme summer weather may limit control of Festuca paniculata by mowing in subalpine grasslands

Background: The tussock grass Festuca paniculata can become strongly dominant in subalpine grasslands after cessation of mowing. The depletion of water-soluble carbohydrate (WSC) reserves has been suggested as a mechanism by which mowing can contain this species. By affecting plant physiology and especially by favouring WSC accumulation, extreme summer weather (i.e. exceptionally hot and dry) could however counterbalance the effects of mowing on WSC reserves in F. paniculata. The relevance of this hypothesis needs to be tested in the current context of climate and land-use changes.

Aims: We investigated (1) the physiological mechanisms that control the growth of F. paniculata, (2) how they are affected by mowing and (3) whether extreme summer heat and drought could influence physiological mechanisms and thereby the ecological response of F. paniculata to mowing.

Methods: In a field experiment we manipulated weather and mowing during two summers. For current summer weather (W0), ambient temperature was unchanged and precipitation was adjusted on the past 30-year average. Extreme summer weather (W+) corresponded to a seasonal change (+1 °C, –80% in precipitation compared to W0) and a three-week heatwave (+4.3 °C) in the first year. In addition, vegetation was either mown at 5 cm in late summer (M) or left unmown (U). Concentrations and absolute contents of WSC contained in tiller bases, leaf nitrogen concentration (LNC), vegetative multiplication, plant growth and leaf senescence were measured from one to four times, depending on the variable considered, throughout the summer of the second year of the experiment.

Results: As compared to the unmown treatment, late-summer mowing decreased tillering, tussock size and LNC, regardless of the summer weather treatment. However, it depleted WSC pools, including fructans, only under current summer weather (W0).

Conclusions: These results suggest that extreme summer heat and drought could alleviate the sensitivity of F. paniculata to mowing. They raise the question of the consequences of recurrent summer extremes for conservation management in subalpine grasslands.     

A leachate a day keeps the seedlings away: mowing and the inhibitory effects of Festuca paniculata in subalpine grasslands

Background and Aims

Is the release of allelochemicals by the dominant tussock grass Festuca paniculata responsible for its dominance by inhibiting growth of neighbour grasses in subalpine grasslands? As such a community is also structured by mowing practices, what could be the impact of mowing on allelopathy?

Methods

A design was used that isolated allelopathy from resource competition by separating donor plants (Festuca paniculata) from target plants (F. paniculata, Dactylis glomerata and Bromus erectus). Leachates from donor pots containing bare soil, unmown F. paniculata or mown F. paniculata continuously irrigated target pots containing seedlings. Activated carbon was added in half of the target pots to adsorb potential allelochemicals. C and N analyses of target potting soil were used to test for any effect of treatments on resources. Total phenol concentration was measured in the solutions flowing from donor to target pots.

Results

Festuca paniculata leachates inhibited seedling growth of D. glomerata and B. erectus. Inhibition was correlated with polyphenol concentration, and was not due to resource competition for nitrogen. Mowing the leaves of the donor plants did not significantly increase this inhibition. The activated carbon treatment was not conclusive as it inhibited the seedling growing under control pots with only bare soil.

Conclusions

The results suggest that allelopathy may be at least partly responsible for F. paniculata dominance in subalpine meadows by inhibition of colonization by neighbouring species.Key words: Allelopathy, chemical interference, mowing, activated carbon, polyphenols, Festuca paniculata, Bromus erectus, Dactylis glomerata, subalpine, competition     

Short-time effects of land-use changes on O-horizon in subalpine grasslands

Due to economic needs, a process of abandonment of unfavourably situated areas and intensified cultivation on favoured areas has taken place in the alpine region. This abandonment not only means the loss of a cultural landscape based on centuries of care but also interference with an established ecological balance. We wanted to investigate the impact of land-use, such as mowing, fertilisation and irrigation and the effects of abandonment on the O-horizon type, quantity and quality. Therefore, we selected areas with different types of land use but with equivalent site conditions in the Passeier Valley (Bolzano-South Tyrol province) for investigation. The areas studied were as follows: an organically fertilised and irrigated, intensively managed hay meadow mown once a year; an organically fertilised, intensively managed hay meadow mown once a year; a lightly managed hay meadow mown every second year; a lightly managed hay meadow mown every 3–4 years; an area densely covered with dwarf shrubs abandoned 10 years ago; and a reforested area abandoned 30 years ago. These selected areas represent different intensities of land use or stages of succession. On these areas, vegetation, phytomass, litter quantity, humus type, O-horizon quantity, litter decomposition and C and N storage, were analysed in detail. Abandonment causes the percentage of coverage with grasses and herbs to decrease from 100 to 10%, while lignified species increase from 0 to 90%. This leads to growth of the phytomass pools, which in turn causes an increase in the annual litter quantity (from 117 to 525 g m⁻²). Significant changes in litter quality in the case of reduced use or abandonment increases the litter’s resistance to decomposition, decreasing overall decomposition. Litter quantity and quality then have a significant impact on the O-horizon quantity. This increases strongly with decreasing intensity of land use or longer times since abandonment (115–1,180 g m⁻²). This leads to a significant increase of C and N storage and a change in O-horizon type ranging from no definite O-horizon or mull O-horizons on regularly used areas to moder on abandoned areas.     

Plant response traits mediate the effects of subalpine grasslands on soil moisture

* In subalpine grasslands, changes in abiotic conditions with decreased management intensity alter the functional composition of plant communities, leading to modifications of ecosystem properties. Here, it is hypothesized that the nature of plant feedbacks on soil moisture is determined by the values of key traits at the community level. * As community functional parameters of grasslands change along a gradient of land uses, those traits that respond most to differences in abiotic conditions produced by land use changes were identified. A vegetation removal experiment was then conducted to determine how each plant community affected soil moisture. * Soil moisture was negatively correlated with community root length and positively correlated with canopy height, whereas average leaf area was associated with productivity. These traits were successfully used to predict the effects on soil moisture of each plant community in the removal experiment. This result was validated using data from an additional set of fields. * These findings demonstrate that the modification of soil moisture following land use change in subalpine grasslands can be mediated through those plant functional traits that respond to water availability.     

Spatial patterns of species distributions in grazed subalpine grasslands

Spatial patterns of species diversity have important influences on the functioning of ecosystems, and the effect of livestock grazing on spatial heterogeneity can differ depending on the scale of the analysis. This study examined the effects of grazing on the spatial patterns of species distributions and whether the effects of grazing on the spatial distributions of a species vary with the scale of the analysis. Data were collected at three locations in the subalpine grasslands of Ordesa-Monte Perdido National Park and Aísa Valley, Central Pyrenees, Spain, which differed in mean stocking rates. Aspect explained about one-third of the environmental variation in species distributions. In flat areas, spatial variation in species composition varied with grazing intensity at two scales. At a coarse scale (among vegetation patches), grazing promoted patchiness, and among-transect variation in species diversity and grazing intensity were positively correlated. At a fine scale (within vegetation patches), the disruption of the self-organizing processes of the species spatial distributions resulted in a reduction in the long-range spatial autocorrelations of some of the characteristic species and in the homogenization of species spatial distributions. The presence of encroaching Echinospartum horridum had a significant influence on the effect of grazing on south-facing grassland slopes.     

The dynamics of two-species allelopathic competition with optimal harvesting

This paper analyses a bionomic model of two competitive species in the presence of toxicity with different harvesting efforts. An interesting dynamics in the first quadrant is analysed and two saddle-node bifurcations are detected for different bifurcation parameters. It is noted that under certain parametric restrictions, the model has a unique positive equilibrium point that is globally asymptotically stable whenever it is locally stable. It is also noted that the model can have zero, one or two feasible equilibria appearing through saddle-node bifurcations. The non-existence of a limit cycle in the interior of the first quadrant is also discussed using the Poincare-Dulac criteria. The saddle-node bifurcations are studied using Sotomayor's theorem. Numerical simulations are carried out to validate the analytical findings. The conditions for the existence of bionomic equilibria are discussed and an optimal harvesting policy is derived using Pontryagin's maximum principle.     

Warming effects on consumption and intraspecific interference competition depend on predator metabolism

1.?Model analyses show that the stability of population dynamics and food web persistence increase with the strength of interference competition. Despite this critical importance for community stability, little is known about how external factors such as the environmental temperature affect intraspecific interference competition. 2.?We aimed to fill this void by studying the functional responses of two ground beetle species of different body size, Pterostichus melanarius and Poecilus versicolor. These functional response experiments were replicated across four predator densities and two temperatures to address the impact of temperature on intraspecific interference competition. 3.?We generally expected that warming should increase the speed of movement, encounter rates and in consequence interference among predator individuals. In our experiment, this expectation was supported by the results obtained for the larger predator, P.?melanarius, whereas the opposite pattern characterized the interference behaviour of the smaller predator P.?versicolor. 4.?These results suggest potentially nontrivial implications for the effects of environmental temperature on intraspecific interference competition, for which we propose an explanation based on the different sensitivity to warming of metabolic rates of both species. As expected, increasing temperature led to stronger interference competition of the larger species, P.?melanarius, which exhibited a weaker increase in metabolic rate with increasing temperature. The stronger increase in the metabolic rate of the smaller predator, P.?versicolor, had to be compensated by increasing searching activity for prey, which did not leave time for increasing interference. 5.?Together, these results suggest that any generalization how interference competition responds to warming should also take the species' metabolic response to temperature increases into account.     

The dynamics of two-species allelopathic competition with optimal harvesting

This paper analyses a bionomic model of two competitive species in the presence of toxicity with different harvesting efforts. An interesting dynamics in the first quadrant is analysed and two saddle-node bifurcations are detected for different bifurcation parameters. It is noted that under certain parametric restrictions, the model has a unique positive equilibrium point that is globally asymptotically stable whenever it is locally stable. It is also noted that the model can have zero, one or two feasible equilibria appearing through saddle-node bifurcations. The non-existence of a limit cycle in the interior of the first quadrant is also discussed using the Poincare–Dulac criteria. The saddle-node bifurcations are studied using Sotomayor's theorem. Numerical simulations are carried out to validate the analytical findings. The conditions for the existence of bionomic equilibria are discussed and an optimal harvesting policy is derived using Pontryagin's maximum principle.     

Low impact of climate change on subalpine grasslands in the Swiss Northern Alps

While phenological shifts and migration of isolated species under climate change have already been observed on alpine summits, very few studies have focused on community composition changes in subalpine grasslands. Here we use permanent plots monitored since 1954 and precisely located phytosociological censuses from 1970 to study compositional changes of subalpine grasslands in two distinct regions of the Swiss Northern Alps. In both areas, warming trends during the monitoring period were associated with changes in land management (abandonment of goat and sheep pasturing or grazing replaced by mowing). Old and recent inventories were compared with correspondence analyses (CA). Ecological indicator values, community‐affinities and biological traits of the species were used to infer the factors responsible for triggering the observed changes. In both regions, subalpine grasslands were stable with smaller changes than have previously been observed in alpine environments. Only a few species appeared or disappeared and changes were generally limited to increasing or decreasing frequency and cover of certain taxa. At one site, grazing abandonment favored fallow species. Some of these species were located at their upper altitudinal distribution limits and may have spread because of rising temperatures. In both areas, declining species were predominantly alpine and low‐growing species; their decline was probably due to increased competition (e.g., shadow) with more vigorous subalpine taxa no longer limited by grazing. We conclude that vegetation communities can respond rapidly to warming as long as colonization is facilitated by available space or structural change. In the subalpine grasslands studies, changes were mainly driven by land management. These communities have a dense vegetation cover and newly arriving herbaceous species preferring warmer conditions may take some time to establish themselves. However, climate disturbances, such as exceptional drought, may accelerate community changes by opening gaps for new species.     

Effects of mowing on fungal endophytes and arbuscular mycorrhizal fungi in subalpine grasslands

In French subalpine grasslands, cessation of mowing promotes dominance of Festuca paniculata, which alters plant diversity and ecosystem functioning. One of the mechanisms underpinning such effects may be linked to simultaneous changes in the abundance of fungal symbionts such as endophytes and arbuscular mycorrhizal fungi. In field conditions, mowing reduced the abundance of the endophyte Neotyphodium sp. in leaves of F. paniculata by a factor of 6, and increased mycorrhizal densities by a factor of 15 in the soil. In greenhouse experiments, the mycorrhizal colonization of Trifolium pratense and Allium porrum increased 3- fold and 3.8- fold respectively in mown vs unmown grassland soil. Significantly reduced growth of the two host plants was also observed on soil from the unmown grassland. Such opposite effects of mowing on the two functional groups of fungal symbionts could suggest interactions between these two groups, which in turn could contribute to structuring plant communities in subalpine grasslands.     

Effects of Festuca paniculata on the compensatory growth response of Centaurea uniflora in the French Alps

The independent effects of herbivores and neighbors on plants are generally negative, and therefore the combined effects of these interactions are generally assumed to have additive or multiplicative negative effects on plant growth. However, because herbivores can stimulate the growth of plants (compensation). and neighbors can facilitate each other, the combined effects of herbivory and plant-plant interactions can be highly variable and poorly predicted by current competition and plant-herbivore theory. In some cases in North America, Festuca species appear to facilitate invasive Centaurea species and enhance their compensatory responses in controlled greenhouse conditions. We explored the interactions between herbivory and neighbor effects in the French Alps by testing the effect of the neighbor, Festuca paniculata L., on the compensatory growth response of defoliated Centaurea uniflora L. over two growing seasons. Seventy percent of aboveground C. uniflora biomass was clipped at each of seven times throughout two growing seasons in the presence or absence of F. paniculata. Centaurea uniflora compensated for severe damage in the first year, but was negatively affected by defoliation in the second year. Defoliating C. uniflora reduced final aboveground biomass by 44% and flower number by 64%, but did not affect survival. Unlike observations for other Centaurea and Festuca species, F. paniculata had significant competitive effects on C. uniflora. Festuca paniculata neither enhanced compensatory responses of C. uniflora nor increased the negative effects of defoliation. Our results show that compensatory responses can weaken over time, but that neighboring plants do not necessarily increase the negative effects of defoliation.     

Encroachment of shrubs into subalpine grasslands in the Pyrenees changes the plant-soil stoichiometry spectrum

Plant and Soil - Shrub encroachment has been reported over a large proportion of the subalpine grasslands across Europe and is expected to have an important impact on the biogeochemical cycle of...     

Linkages between grazing history and herbivore exclusion on decomposition rates in mineral soils of subalpine grasslands

Background & aims

Herbivore-driven changes to soil properties can influence the decomposition rate of organic material and therefore soil carbon cycling within grassland ecosystems. We investigated how aboveground foraging mammalian and invertebrate herbivores affect mineral soil decomposition rates and associated soil properties in two subalpine vegetation types (short-grass and tall-grass) with different grazing histories.

Methods

Using exclosures with differing mesh sizes, we progressively excluded large, medium and small mammals and invertebrates from the two vegetation types in the Swiss National Park (SNP). Mineral soil decomposition rates were assessed using the cotton cloth (standard substrate) method between May and September 2010.

Results

Decomposition displayed strong spatio-temporal variability, best explained by soil temperature. Exclusion of large mammals increased decomposition rates, but further exclusion reduced decomposition rates again in the lightly grazed (tall-grass) vegetation. No difference among treatments was found in the heavily grazed (short-grass) vegetation. Heavily grazed areas had higher decomposition rates than the lightly grazed areas because of higher soil temperatures. Microbial biomass carbon and soil C:N ratio were also linked to spatio-temporal decomposition patterns, but not to grazing history.

Conclusions

Despite altering some of the environmental controls of decomposition, cellulose decomposition rates in the SNP’s subalpine grasslands appear to be mostly resistant to short-term herbivore exclusion.     

The early effects of afforestation on biodiversity of grasslands in Ireland

The target rate of afforestation in Ireland over the next 30 years is 20,000 ha per year, which would result in an increase of the forest cover from the current 10% to 17%. In order to promote sustainable forest management practices, it is essential to know the composition and conservation value of habitats where afforestation is planned and the effects of subsequent planting upon biodiversity. The objectives of this study were to investigate changes in vegetation composition and diversity of grasslands 5 years after afforestation with Sitka spruce (Picea sitchensis) and determine the primary ecological and management factors responsible for these changes. Species cover, environmental and management data were collected from 16 afforested and unplanted improved and wet grassland site pairs in Ireland. Our results indicate that 5 years after tree planting, there were significant changes in richness, composition, and abundance of species. Competitive and vigorous grasses were more abundant in planted than in unplanted sites, as were generalist species found in both open and wooded habitats, while small-stature shade-sensitive species were less abundant. Vascular plant species richness and Shannon’s diversity index were higher in unplanted wet grassland, than in the planted sites. Bryophyte species richness was higher in planted improved grassland than in unplanted sites. The differences were primarily the result of the exclusion of grazing, ground preparation, changes in nutrient management and drainage for afforestation. Drainage ditches provided a temporary habitat for less competitive species, but the overall effect of drainage was to reduce the diversity of species dependent on wet conditions. Variance partitioning showed differences in the relative influences of environmental and management variables on biodiversity in the two habitats, probably due to the greater pre-afforestation grazing pressure and fertilisation levels in improved grasslands. The differences in biodiversity between planted and unplanted grasslands indicate that afforestation represents a threat to semi-natural habitats where distinctive and highly localised plant communities could potentially occur.     

The effects of cue competition on timing in pigeons

We studied the effects of cue competition on timing in both overshadowing and blocking operant procedures with pigeons. A white center key delivered reward when pecked 30 s after a red or green sidekey was presented and 10 s after presentation of the alternate color on the other sidekey. In Experiment 1, key presentations were concurrent during training trials for overshadow-condition pigeons, while side key presentations were separated across training trials for control birds. In Experiment 2, half of the birds (Blocking group) were given pre-exposure trials to either the 10-s or 30-s sidekey condition. Both blocking-condition and control birds were then given trials of concurrent side key presentations. Peak time curves were compared between experimental and control conditions. The results showed blocking of timing accuracy of a long (30-s) stimulus by a short (10-s) stimulus, but no evidence for overshadowing of timing accuracy.     

Density dynamics of bumblebees in subalpine meadows: competition and resource limitation

Densities of Bombus flavifrons and B. rufocinctus were measured over one summer in 23 and 11 discrete subalpine meadows, respectively. I assessed the relation between observed bee densities and meadow size, elevation, floristics, temperature, and time of day by multiple regression. Multivariate regression solutions for each week of observation accounted for 38 to 61% of the variation in density for B. flavifrons and 47 to 87% for B. rufocinctus . Temperature or time of day consistently accounted for the most variation in density of the two species. In early summer meadow size or elevation also accounted for a large proportion of explained variation in B. flavifrons density; as the summer progressed meadow floristics increasingly contributed to the explanation of variation in both species.
In mid- to late summer a statistically significant portion of the variation in densities not explained by habitat characteristics was accounted for by densities of congeners: i.e., densities of B. flavifrons and B. rufocinctus were negatively related in eight meadows where they co-occurred. In experimental meadows where all Bombus but one of the two target species were removed. B. rufocinctus underwent significant positive density compensation but B. flavifrons did not. To the extent that B. flavifrons appeared to limit the density of B. rufocinctus these statistical and experimental results are consistent with the hypothesis that competition limits local bumblebee densities.