Changes in semi-arid plant species associations along a livestock grazing gradient

In semi-arid ecosystems, vegetation is heterogeneously distributed, with plant species often associating in patches. These associations between species are not constant, but depend on the particular response of each species to environmental factors. Here, we investigated how plant species associations change in response to livestock grazing in a semi-arid ecosystem, Cabo de Gata-Níjar Natural Park in South East Spain. We established linear point-intercept transects at four sites with different grazing intensity, and recorded all species at each point. We investigated plant associations by comparing the number of times that each pair of species occurred at the same spatial point (co-occurrences), with the expected number of times based on species abundances. We also assessed associations for each shrub and grass species by considering all their pairs of associations and for the whole plant community by considering all pairs of associations on each site. At all sites, the plant community had a negative pattern of association, with fewer co-occurrences than expected. Negative association in the plant community increased at maximum grazing intensity. Most species associated as expected, particularly grass species, and positive associations were most important at intermediate grazing intensities. No species changed its type of association along the grazing gradient. We conclude that in the present plant community, grazing-resistant species compete among themselves and segregate in space. Some shrub species act as refuges for grazing-sensitive species that benefit from being spatially associated with shrub species, particularly at intermediate grazing intensities where positive associations were highest. At high grazing intensity, these shrubs can no longer persist and positive associations decrease due to the disappearance of refuges. Spatial associations between plant species and their response to grazing help identify the factors that organize plant communities, and may contribute to improving management of semi-arid ecosystems.     

Changes in plant interactions along a gradient of environmental stress

A combination of competition and facilitation effects operating simultaneously among plant species appears to be the rule in nature, where these effects change along productivity gradients often in a non-proportional manner. We investigated changes in competition and facilitation between a leguminous shrub, Retama sphaerocarpa , and its associate understorey species along an environmental gradient in semi-arid southeast Spain. Our results show a change in the net balance of the interaction between the shrub and several of its associated species, from clearly positive in the water-stressed, infertile environment to neutral or even negative in the more fertile habitat. There was a weakening of facilitation along the fertility gradient as a consequence of improved abiotic conditions. Competition was the most intense for below-ground resources in the less fertile environment while total competition tended to increase towards the more productive end of the gradient. Changes in the balance of the interaction between and among different plant species along the gradient of stress were caused by a decline in facilitation rather than by a change in competition. As both competition intensity and facilitation change along gradients of resource availability, plant interactions are best viewed as dynamic relationships, the outcome of which depends on abiotic conditions.     

Goose grazing selectivity along a depletion gradient

The micro-habilat food selectivity along a gradient of increasing grazing intensity of wintering pink-footed geese Anser brachyrhynchus and barnacle geese Branta leucopsis foraging on winter wheat Triticum aestivum and smooth meadow grass Poa pratensis was investigated. Protein content of ungrazed primary and secondary leaves of winter wheat did not differ, whereas the protein content of tertiary leaves was significantly lower. Secondary leaves were significantly longer than both primary and tertiary leaves, whereas tertiary leaves were significantly longer than primary leaves. Protein content of primary and secondary leaves of Poa pratensis did not differ, whereas the protein content of tertiary leaves was significantly lower. Primary leaves were significantly shorter than both secondary and tertiary leaves, which did not differ significantly.
For all leaf categories, there was an increase in the removal with increasing goose grazing intensity. The geese showed a sequential selection of winter wheat leaves according to the protein content, initially preferring primary and secondary leaves, but increasingly selecting tertiary leaves of poorer quality as high-quality leaves became depleted. Foraging on Poa pratensis. geese selected secondary leaves according to the protein content, whereas tertiary leaves were preferred to primary leaves, although these had a protein content equal to secondary leaves. This, however, may be explained by the primary leaves being too short to maintain intake rates.
It is concluded that feeding preferences cannot be established on the basis of protein contents of food plants alone and that green biomass estimates not necessarily reflect resource availability.     

Stress and disturbance determine the demographic dynamics of two grass species along a grazing gradient in Southern Mexico

Chronic anthropogenic disturbance (CAD), characterized by low-intensity but high frequency, is a major driver of environmental degradation in developing countries. CAD is a mixture of disturbance sensu stricto (DSS), that is, plant biomass removal and stress that reduces biomass production due to changes in environmental conditions. However, we still lack data on the separate effects of both components and their interaction in nature. We analyze the demographic effects of DSS and stress on two grass species in an area heavily affected by livestock raising (a widespread cause of CAD) during the last 500 year. We compared areas exposed to DSS and stress with areas without grazing but that continue experiencing a gradient of stress. Using matrix and integral projection models, we analyzed DSS and stress effects on population growth rates (λ) of two grass species and determined the relative importance of different vital rates and states for the change on λ. Disturbance and stress affected different individuals and processes. For example, changed conditions due to stress increased seedling mortality, but DSS reduced size (growth) of large plants through grazing. CAD had highly nonlinear and species-specific effects on population size structures, λ and elasticities. Such complex behavior is seemingly due to changes in the components of CAD as it intensified and synergic interactions between disturbance and stress. Given CAD's multivariate nature, these results are not surprising. Nevertheless, grouping this multitude of factors into two broad categories, namely DSS and stress, may prove a useful conceptual tool for analysis.     

Shift from facilitative to neutral interactions by the cushion plant Silene acaulis along a primary succession gradient

Background

The stress‐gradient hypothesis predicts a shift from facilitative to competitive plant interactions with decreasing abiotic stress. This has been supported by studies along elevation and temperature gradients, but also challenged by the hypothesis of a facilitation collapse at extremely harsh sites. Although facilitation is known to be important in primary succession, few studies have examined these hypotheses along primary succession gradients.

Aim

To examine whether there is a relationship between the presence of the circumpolar cushion plant Silene acaulis and other species, and if so, whether there is a shift between positive and negative interactions along a primary succession gradient in a glacier foreland.

Location

Finse, southern Norway.

Methods

We examined the performance of the common alpine forb Bistorta vivipara, species richness of vascular plants, bryophytes and lichens, and the number of seedlings and fertile vascular plants in S. acaulis cushions, and control plots without S. acaulis, along a succession gradient with increasing distance from a glacier front, and thus decreasing abiotic stress. To examine if S. acaulis cushions modify the abiotic environment, we recorded soil temperature, moisture, organic content and pH in cushions and control plots.

Results

Bistorta vivipara performed better, as shown by bigger leaves in S. acaulis cushions compared to control plots in the harshest part of the gradient close to the glacier. There were few differences in B. vivipara performance between cushion and control plots in the more benign environment further away from the glacier. This suggests a shift from facilitative to mainly neutral interactions by S. acaulis on the performance of B. vivipara with decreasing abiotic stress. A trend, although not significant, of higher vascular species richness and fertility inside S. acaulis cushions along the whole gradient, suggests that S. acaulis also facilitates community‐level species richness. The causal mechanism of this facilitation is likely that the cushions buffer extreme temperatures.

Conclusions

Our results support the stress‐gradient hypothesis for the relationship between the cushion plant S. acaulis and the performance of a single species along a primary succession gradient in a glacier foreland. S. acaulis also tended to increase vascular plant species richness and fertility regardless of stress level along the gradient, suggesting facilitation at the community level. We found no collapse of facilitation at the most stressful end of the gradient in this alpine glacier foreland.     

Phytoplankton competition along a gradient of dilution rates

Natural phytoplankton from Lake Constance was used for chemostat competition experiments performed at a variety of dilution rates. In the first series at high Si:P ratios and under uniform phosphorus limitation for all species, Synedra acus outcompeted all other species at all dilution rates up to 1.6 d^-1, only at the highest dilution rate (2.0 d^-1) Achnanthes minutissima was successful. In the second series in the absence of any Si a green algal replacement series was found, with Mougeotia thylespora dominant at the lowest dilution rates, Scenedesmus acutus at the intermediate ones, and Chlorella minutissima at the highest ones. The outcome of interspecific competition was not in contradiction with the Monod kinetics of P-limited growth of the five species, but no satisfactorily precise prediction of competitive performance can be derived from the Monod kinetics because of insufficient precision in the estimate of k _s .     

Arbuscular mycorrhizal fungi alter plant community composition along a grazing gradient in Inner Mongolia Steppe

Arbuscular mycorrhizal fungi (AMF) have a significant influence on plant productivity and diversity in non-grazing grassland. However, the interactive effects between grazing intensity and AMF on plant community composition in natural grassland communities are not well known. We conducted a field experiment that manipulated AMF colonization and grazing intensity to study the impact of AMF suppression on plant community composition and nutrient status over 2 years (2015–2016) with contrasting rainfall levels. We found that AMF root colonization was significantly reduced by the application of the fungicide benomyl as a soil drench. Grazing intensity regulated plant community composition and aboveground biomass mainly by reducing the growth of Leymus chinensis over 2 years. AMF suppression increased the growth of Chenopodium glaucum, but it did not alter other plant species across all grazing intensities. The effects of AMF suppression on plant community composition changed along a grazing gradient considerably between years: AMF suppression increased the biomass of C. glaucum across all grazing intensities in 2015, but slightly increased it in 2016. Interactions between AMF suppression and grazing intensity altered the phosphorus concentration of Stipa grandis and Cleistogenes squarrosa in 2015 but not in 2016. AMF suppression decreased the shoot phosphorus content of L. chinensis but increased that of C. glaucum across all grazing intensities. Our results indicate that grazing intensity substantially alters aboveground community biomass and affects growth of dominant species; AMF by itself have limited effects on plant communities along a grazing gradient in typical steppe.     

Impact of competition on plant performances along a flooding gradient: a multi‐species experiment

Question: 1. How do the competitive response and the importance of competition vary between species and along a flooding gradient? 2. How does the role of competition in constraining species distribution limits along the gradient vary between lower and upper limits? Location: A 1‐ha meadow within the Alzette floodplain in Luxembourg. Methods: Competitive response and importance of competition were assessed on seven meadow species differing in their tolerance to flooding. Species were cultured in monocultures and in mixtures, in three water treatments reflecting the wet, the middle and the dry end of a natural flooding gradient. We developed two models based on a multiple regression in order to express each component of competition as a function of the neighbour biomass. Results: Five species showed variations in their competitive response across water treatments; however, these species achieved either their highest or their worst competitive response in their optimal water treatment (i.e. the treatment in which the species had the highest biomass in monoculture). Competition was more important for the flood‐tolerant species in the dry treatment than for the flood‐intolerant species in the wet treatment. Conclusions: 1. Variations in species competitive responses along flooding gradients may be the result of either an amplified effect between competition and hydrological stresses, or a hierarchical effect of stress over competition. 2. The role of competition is more important in constraining the upper distribution limits of the flood‐tolerant species than the lower limits of the flood‐intolerant species along flooding gradients.     

Oak-insect herbivore interactions along a temperature and precipitation gradient

The interactions between herbivorous insects and their host plants are expected to be influenced by changing climates. Modern oaks provide an excellent system to examine this assumption because their interactions with herbivores occur over broad climatic and spatial scales, they vary in their defensive and nutritional investment in leaves by being deciduous or evergreen, and their insect herbivores range from generalists to highly specialized feeders. In this study, we surveyed leaf-litter samples of four oak species along an elevation gradient, from coastal northern California, USA, to the upper montane woodlands of the Sierra Nevada, to examine the relationship between climatic factors (mean annual temperature and precipitation) and oak herbivory levels at multiple scales; across all oak species pooled, between evergreen and deciduous species and within species.Overall, temperature and precipitation did not appear to have a significant effect on most measures of total herbivore damage (percent leaves damaged per tree, percent leaf area removed and average number of feeding damage marks per leaf) and the strongest predictor of herbivore damage overall was the identity of the host species. However, increases in precipitation were correlated with an increase in the actual leaf area removed, and specialized insects, such as those that make leaf mines and galls, were the most sensitive to differences in precipitation levels. This suggests that the effects of changing climate on some plant–insect interactions is less likely to result in broad scale increases in damage with increasing temperatures or changing precipitation levels, but is rather more likely to be dependent on the type of herbivore (specialist vs. generalist) and the scale (species vs. community) over which the effect is examined.     

Effects of grazing on plant communities and successional processes vary along an aridity gradient at a northern temperate grassland

The intermountain grasslands of North America reach their most northern geographic extent in interior British Columbia’s Cariboo-Chilcotin region. Here, this study examined the long-term effects of livestock grazing exclusion and reductions in grazing severity on plant community characteristics including plant and litter cover, species richness and abundance of leading species of 33 grassland sites across a broad aridity and soil property gradient. Across the aridity gradient, grazing reduced species richness, plant cover, and litter cover. However, the effects of grazing on dominant species varied across the gradient. In more arid grasslands, historical grazing substantially reduced cover of late-seral native bunchgrass Psuedoroegnaria spicata, and repeated measurements indicate that very long time periods are necessary for successional processes associated with recovery of native bunchgrasses. At the cool-wet end of the aridity gradient, successional processes are more rapid but dominated by exotic species Poa pratensis and Tragopogon pratensis. Recent (past 20 years) light grazing and rest-rotation have favored Poa pratensis at the expense of native needlegrasses (Achnatherum spp. and Hesperostipa spp.). We suggest that absence of a dominant large-stature native bunchgrass for mesic grasslands was a key factor in the invasion and dominance of Poa pratensis.

     

Importance and intensity of competition along a fertility gradient and across species

Questions: 1. Can the importance and the intensity of competition vary independently along a nutrient gradient? 2. Are these variations species dependent? Location: Sub‐alpine pastures of the northern French Alps. Methods: Competition intensity measures how much competition decreases the performances of an organism. Competition importance measures how much competition contributes to affect performance, among other processes (such as environmental stress or disturbance). Competition intensity and importance were measured on three co‐occurring species: Festuca rubra, a perennial grass, and two forbs of contrasting basal area, Chaerophyllum hirsutum and Alchemilla xantho‐chlora. A neighbour removal experiment was performed on Festuca rubra in three sub‐alpine grassland communities differing in fertility and on Chaerophyllum hirsutum and Alchemilla xanthochlora in the two more fertile of these communities. The importance of competition was quantified using an index proposed by Brooker et al. (2005). Results: Competition intensity and importance showed different patterns of variation along the fertility gradient for Festuca rubra: competition importance decreased with decreasing fertility whereas competition intensity did not change. The largest forb was the least affected by competition. Our results suggest that the importance of competition for all three species depended on their individual tolerance to low nutrient availability. Conclusions: 1. The distinction between the importance and the intensity of competition is helpful to explain conflicting results obtained on the variations of competition indices along productivity gradients. 2. The choice of a phytometer can affect the conclusions drawn from empirical studies.     

Plant species responses along a grazing disturbance gradient in Australian grassland

Abstract. Floristic data from paired roadside-paddock analyses from grassland in central Queensland, Australia, were ordinated. The mean direction of the vectors between these pairs was almost perfectly aligned with the indirect gradient represented by the first axis of Non-metric Multi-dimensional Scaling. It confirms anecdotal evidence of a trend from infrequently grazed roadsides to constantly grazed paddocks. The increasing abundance of annual herbs and grasses along this putative gradient is consistent with documented trends from elsewhere in the world. The response patterns of individual species along the disturbance gradient is consistent with ecological theory predicting unimodal peaks in abundance along physical environmental gradients. The ancestral perennial dominants of the grasslands, Dichanthium sericeum and D. queenslandicum, exhibited a declining response to grazing disturbance. Even the generally unpalatable perennial grass Aristida leptopoda declined considerably in the upper segments of the grazing disturbance gradient. A suite of herbaceous trailing legumes had peaks in their abundance near the middle of the grazing disturbance gradient, trends that can be readily explained by the combination of their palatability and intolerance to competition from tall perennial grasses. Several species including the noxious exotic herb Parthenium hysterophorus showed increasing abundance along the grazing disturbance gradient. The methodology may have application as a rapid method of assessing disturbance impacts elsewhere, and is most suited where a management differential between paired plots can be reliably generalized and where the physical environment is relatively monotonous.     

Grass-to-grass protection from grazing in a semi-arid steppe. Facilitation, competition, and mass effect

Plants of low palatability often serve as biotic refuges from grazing to palatable plants. Evidence for this facilitation comes from cases where the interacting species have different life form, which may minimize competition. Protected plants act as remnant seed sources that may maintain the palatable populations locally viable through mass effects. Here, we assess (1) the spatial association between a highly palatable Patagonian grass (Bromus pictus) and less preferred tussock grasses, (2) the role played by seed sources in maintaining the population in the face of heavy grazing by sheep, and (3) the facilitative and competitive components of the interaction. We quantified B. pictus density and its distance to nearest tussocks inside and outside a grazing exclosure. We also considered different distances from the exclosure, both leeward and windward, because strong westerly winds may be critical for dispersal. Additionally, we quantified several attributes of protected and unprotected B. pictus plants with and without grazing. Density of B. pictus was about 20 times greater inside the exclosure than outside. However, this difference was less pronounced in the leeward vicinity of the exclosure than in the windward one, which suggests a mass effect. B. pictus was significantly associated to less palatable tussocks, and the association became stronger under grazing and as distance from the exclosure edge increased. Protection under grazing was associated with a significant increase in plant biomass, height, tiller number, and panicle number, whereas protection in the absence of grazing, which could evidence competition, resulted in reductions of tiller number and panicle number, and an increase of height. These results suggest that in areas under grazing pressure on palatable grasses, other less palatable grasses may provide a protection from grazing that outweighs competitive effects. Such protection may generate small-scale mass effects that maintain the population at relatively high density.