Intensity-dependent effects of cattle and sheep grazing in sand grasslands — Does livestock type really matter?

Aims

By analysing cattle- and sheep-grazed sand grasslands, we tested the following hypotheses: (i) livestock type has a stronger effect on the vegetation characteristics than grazing intensity; (ii) sheep grazing results in lower biomass and species and functional diversity than cattle grazing, regardless of intensity; and (iii) increased grazing intensity causes a shift of the trait composition in grasslands.

Location

Sand grasslands in the Nyírség region, East Hungary.

Methods

We selected 26 sand grassland sites grazed by cattle or sheep and classified them into four intensity levels. Vegetation composition was surveyed in 2 m × 2 m plots. We harvested the above-ground biomass from 20 cm × 20 cm plots; then dried and sorted it to live biomass, litter, moss, and lichen. We compared Rao dissimilarity index, species richness, Shannon diversity, evenness, and the community-weighted means of nine vegetative and generative traits along a grazing intensity gradient. We calculated functional richness, evenness, and divergence for comparison.

Results

We found that some diversity metrics and community-weighted means of most studied traits were significantly affected by grazing intensity. Several characteristics were also affected by the interaction of grazing intensity and livestock type, but none of the studied characteristics was affected by livestock type in itself. Increasing Rao dissimilarity index peaking at the fourth grazing intensity level was detected, but for other multitrait indices, no such changes were proven, except for functional divergence, which was the lowest at the first intensity level. Graminoid, forb, and litter biomass were significantly affected by intensity, but none of the biomass fractions was affected by livestock type.

Conclusions

We suggest that for the management of sand grasslands, grazing intensity should be carefully adjusted, considering not only livestock units per hectare. For practical recommendations, well-defined, long-term experiments studying different livestock and habitat types along an intensity gradient would be essential.     

Evaluating the benefits of CAP reforms: Can afforestations restore bird diversity in Mediterranean Spain?

Current Common Agricultural Policy (CAP) subsidies in the Mediterranean region tend to prioritize afforestation on former arable land with oaks rather than pines because pine plantations would maintain lower biological diversities than native forests. Nevertheless, no thorough evaluations of the conservation values of pine plantations as compared to oak remnants have been carried out to date. We analyze the diversity and conservation value of bird assemblages breeding in 200 remnants of Holm oak Quercus ilex woodlands and 82 mature (>50-year-old) pine plantations in central Spain, a Mediterranean region mostly devoted to arable farming. Species–area relationships were compared between forest types. The conservation value of bird assemblages was assessed using the “Species of European Conservation Concern” (SPEC) classification of Burfield and van Bommel [(2004). Birds in Europe: Populations estimates, trends and conservation status. Cambridge: BirdLife International]. Overall numbers of bird species maintained by oak and pine archipelagoes were rather similar, but species–area relationships differed between forest types. Intercepts were higher in oak fragments, whereas slopes were steeper in pine plantations. Small oak fragments held more species (mainly Mediterranean Sylvia warblers) than plantations, whereas large plantations held more species than large oak remnants. Differences in species–area relationships seemed to be due to differences in vegetation structure, especially understorey shrub cover and tree height and cover. We recorded nine SPECs, all exclusive (6) or near-exclusive (3) to oak woodlands, although such woodlands do not appear to be critical for their conservation. Hence, we conclude that pine afforestations have played a role for maintaining and restoring forest bird communities in the farming landscapes of central Spain. Promoting large and shrubby plantations would enhance their conservation value for breeding birds, together with promoting growth, regeneration and expansion of Holm oak remnants by means of set-aside measures previous or alternative to oak reafforestation. The increasing importance of non-commercial as compared to commercial values of Mediterranean forests would justify subsidizing the proposed policy.     

Is vascular plant species diversity a predictor of bryophyte species diversity in Mediterranean forests?

This study aimed to (i) investigate the congruence among the species composition and diversity of bryophytes and vascular plants in forests; (ii) test if site prioritization for conservation aims by the maximization of the pooled number of vascular plant species is effective to maximize the pooled number of bryophyte species. The study was performed in six forests in Tuscany, Italy. Four-hundred and twenty vascular plant species (61 of which were woody) and 128 bryophyte species were recorded in 109 plots. Despite the good predictive value of the compositional patterns of both woody plants and total vascular with respect to the compositional pattern of bryophytes, the species richness of the latter was only marginally related to the species richness of the former two. Bryophyte rare species were not spatially related to rare plant species and neither coincided with the sites of highest plant species richness. The species accumulation curves of bryophytes behaved differently with respect to those of woody plants or total vascular plants. Reserve selection analysis based on the maximization of the pooled species richness of either woody plants or total vascular plants were not effective in maximizing the pooled species richness of bryophytes. This study indicates that species diversity of vascular plants is not likely to be a good indicator of the bryophyte species diversity in Mediterranean forests.     

Does wild boar rooting affect livestock grazing areas in alpine grasslands?

Interactions between traditional livestock management practices and wildlife activities are important in the conservation of many mountain ecosystems including the summer rangelands in the Spanish Central Pyrenees, where rooting by wild boar (Sus scrofa) is a large disturbance that can reduce the amount of area available to grazing livestock. This study explored the likely impact of wild boar rooting on Pyrenean grasslands. It quantified the extent of wild boar rooting in livestock grazing areas and determined whether wild boars selected or avoided areas depending on the type of livestock and stocking rates. Wild boar rooting affected 16% of livestock grazing area and occurred in sites that were grazed by cattle, rather than by sheep. In addition, a preference for areas that had intermediate stocking rates was found. The relationship between the increase in the number of wild boars and trends in livestock management suggests that the extent of wild boar rooting will increase especially in cattle grazing areas, and therefore, the area available for cattle grazing in Pyrenean mountain rangelands would decrease significantly.     

Does plant phylogenetic diversity increase invertebrate herbivory in managed grasslands?

Plant diversity and land-use intensity have been shown to affect invertebrate herbivory. Several hypotheses predict positive (e.g. associational susceptibility) or negative (e.g. associational resistance) relationships of herbivory with plant species richness. Also, the strength and direction of reported relationships vary greatly between studies leading to the conclusion that relationships either depend on the specific system studied or that other unconsidered factors are more important. Here, we hypothesized that plant phylogenetic diversity is a stronger predictor of invertebrate herbivory than plant species richness because it integrates additional information about the phenotypical and functional composition of communities. We assessed the community-wide invertebrate herbivory, plant species richness and phylogenetic diversity across a range of land-use intensities including a total of 145 managed grasslands in three regions in Germany. Increasing land-use intensity decreased plant species richness and phylogenetic diversity. Plant species richness did not predict invertebrate herbivory. By contrast herbivory moderately increased with increasing plant phylogenetic diversity even after accounting for the effects of region and land use. The strength of direct effects of land-use intensity and indirect effects via altered phylogenetic diversity on herbivory, however, varied among regions. Our results suggest that increasing phylogenetic diversity of plant communities increases invertebrate herbivory probably by providing higher resource diversity. Differences between regions underline the need to account for regional peculiarities when attempting to generalize land-use effects on invertebrate herbivory.     

Which functional processes control the short-term effect of grazing on net primary production in grasslands?

For unicellular organisms, a lack of effects of local species richness on ecosystem function has been proposed due to their locally high species richness and their ubiquitous distribution. High dispersal ability and high individual numbers may enable unicellular taxa to occur everywhere. Using our own and published data sets on uni- and multicellular organisms, we conducted thorough statistical analyses to test whether (1) unicellular taxa show higher relative local species richness compared to multicellular taxa, (2) unicellular taxa show lower slopes of the species:area relationships and species:individuals relationships, and (3) the species composition of unicellular taxa is less influenced by geographic distance compared to multicellular taxa. We found higher local species richness compared to the global species pool for unicellular organisms than for metazoan taxa. The difference was significant if global species richness was conservatively estimated but not if extrapolated, and therefore higher richness estimates were used. Both microalgae and protozoans showed lower slopes between species richness and sample size (area or individuals) compared to macrozoobenthos, also indicating higher local species richness for unicellular taxa. The similarity of species composition of both benthic diatoms and ciliates decreased with increasing geographic distance. This indicated restricted dispersal ability of protists and the absence of ubiquity. However, a steeper slope between similarity and distance was found for polychaetes and corals, suggesting a stronger effect of distance on the dispersal of metazoans compared to unicellular taxa. In conclusion, we found partly different species richness patterns among uni- and multicellular eukaryotes, but no strict ubiquity of unicellular taxa. Therefore, the effect of local unicellular species richness on ecosystem function has to be reanalyzed. Macroecological patterns suggested for multicellular organisms may differ in unicellular communities.     

Can livestock grazing maintain landscape diversity and stability in an ecosystem that evolved with wild herbivores?

The livestock industry is converting mountain ecosystems in central Argentina into rocky deserts. However, it helps to conserve plant biodiversity, presumably because the ecosystem has evolved under wild herbivores now locally extinct. We hypothesized that low or moderate livestock stocking rates, instead of the high stocking rates currently used for commercial production, might mimic pre-hispanic herbivore pressures. Thus, the mosaic of physiognomies necessary to maintain landscape diversity and soil integrity could be preserved. To test this hypothesis we tracked physiognomic changes in 200 plots (16 m² each) under different stocking rates, including livestock exclusion, for five years. Contrary to our expectations, we found that both low and moderate stocking rates failed to maintain landscape diversity. As observed for livestock exclusion, low to moderate stocking rates promote retraction and, finally, elimination of short grazing lawns and their replacement by tall tussock grasslands, or possibly by woody vegetation. In turn, heavy grazing pressure maintains the desired short lawn patches in the landscape, but also promotes a concomitant loss and eventual elimination of woodlands, together with an expansion of bare rock as a consequence of soil erosion. These results indicate that the present mosaic of physiognomic types is difficult to maintain and raise questions about the past stability of the ecosystem. We suggest that during the last 400 years of livestock production in the ecosystem, short lawns have been maintained by means of heavy stocking rates and anthropogenic fires ignited to eliminate tussocks. However, this relative stability of lawns has been attained concomitantly with a progressive loss of soils and woodland area. We also discuss some possible explanations for the maintenance of short lawns before the introduction of livestock, based on main climatic and floristic shifts during the late Holocene.     

Stimulation of soil nitrification and denitrification by grazing in grasslands: do changes in plant species composition matter?

Stimulation of nitrification and denitrification by long term (from years to decades) grazing has commonly been reported in different grassland ecosystems. However, grazing generally induces important changes in plant species composition, and whether changes in nitrification and denitrification are primarily due to changes in vegetation composition has never been tested. We compared soil nitrification- and denitrification-enzyme activities (NEA and DEA, respectively) between semi-natural grassland sites experiencing intensive (IG) and light (LG) grazing/mowing regimes for 13 years. Mean NEA and DEA (i.e. observed from random soil sampling) were higher in IG than LG sites. The NEA/DEA ratio was higher in IG than LG sites, indicating a higher stimulation of nitrification. Marked changes in plant species composition were observed in response to the grazing/mowing regime. In particular, the specific phytomass volume of Elymus repens was lower in IG than LG sites, whereas the specific volume of Lolium perenne was higher in IG than LG sites. In contrast, the specific volume of Holcus lanatus, Poa trivialis and Arrhenatherum elatius were not significantly different between treatments. Soils sampled beneath grass tussocks of the last three species exhibited higher DEA, NEA and NEA/DEA ratio in IG than LG sites. For a given grazing regime, plant species did not affect significantly soil DEA, NEA and NEA/DEA ratio. The modification of plant species composition is thus not the primary factor driving changes in nitrification and denitrification in semi-natural grassland ecosystems experiencing long term intensive grazing. Factors such as trampling, N returned in animal excreta, and/or modification of N uptake and C exudation by frequently defoliated plants could be responsible for the enhanced microbial activities.     

Can switches in disturbance type improve habitat for grassland birds in semi-arid grasslands of South-Eastern Australia?

Grasslands are one of the most endangered and degraded ecosystems globally. Switches in disturbance type can restore grassland function and improve conservation outcomes for fauna, but land-use legacies can limit the capacity of biota to respond positively to shifts in disturbance type, making it difficult to predict ecological outcomes. In semi-arid grasslands of South-Eastern Australia, habitat for grassland birds is managed using livestock grazing, a practice that has continued for >150 years. It is unknown if outcomes for birds can be improved by a switch in disturbance type as alternatives to livestock grazing have not been explored. We compare the effects of status-quo livestock grazing with alternative biomass management tools (no management, ‘crash’ grazing, planned fire) on birds, vegetation structure, and food resources using a BACI design across 3 years (2017–2019). We found crash grazing and fire produced more open, shorter grassy swards, with less exotic grass cover, compared to status-quo grazing or disturbance exclusion. Preferred habitat structure for the critically endangered plains-wanderer (Pedionomus torquatus) was maximized by status-quo grazing. Grassland birds responded in opposing ways to a disturbance switch. Brown songlarks (Megalurus cruralis) and Horsfield's bushlarks (Mirafra javanica) responded positively to disturbance exclusion, while stubble quail (Coturnix pectoralis) responded negatively to crash grazing. Australasian pipits (Anthus novaeseelandiae) were more frequent in response to status-quo grazing. Our findings suggest that multiple disturbance types should be used if the aim is to promote the spectrum of vegetation structures and food sources required to support a diverse grassland bird community in semi-arid grasslands of Australia.     

Species introduction in restoration projects – Evaluation of different techniques for the establishment of semi-natural grasslands in Central and Northwestern Europe

During recent decades, many studies have shown that the successful restoration of species-rich grasslands is often seed-limited because of depleted seed banks and limited seed dispersal in modern fragmented landscapes. In Europe, commercial seed mixtures, which are widely used for restoration measures, mostly consist of species and varieties of non-local provenance. The regional biodiversity of a given landscape, however, can be preserved only when seeds or plants of local provenance are used in restoration projects. Furthermore, the transfer of suitable target species of local provenance can strongly enhance restoration success.We review and evaluate the success of currently used near-natural methods for the introduction of target plant species (e.g. seeding of site-specific seed mixtures, transfer of fresh seed-containing hay, vacuum harvesting, transfer of turves or seed-containing soil) on restoration sites, ranging from dry and mesic meadows to floodplain grasslands and fens. Own data combined with literature findings show species establishment rates during the initial phase as well as the persistence of target species during long-term vegetation development on restoration sites.In conclusion, our review indicates that seed limitation can be overcome successfully by most of the reviewed measures for species introduction. The establishment of species-rich grasslands is most successful when seeds, seed-containing plant material or soil are spread on bare soil of ex-arable fields after tilling or topsoil removal, or on raw soils, e.g. in mined areas. In species-poor grasslands without soil disturbance and on older ex-arable fields with dense weed vegetation, final transfer rates were the lowest. For future restoration projects, suitable measures have to be chosen carefully from case to case as they differ considerably in costs and logistic effort. Long-term prospects for restored grassland are especially good when management can be incorporated in agricultural systems.     

Can tolerant allogeneic cells restore nude mice?

BALB/c-nunu mice have been injected with allogeneic or tolerant allogeneic spleen or thymus cells and sheep red blood cells (SRBC). From day 3 to day 10 the mice were bled daily and the antiSRBC antibody was assayed by hemagglutination. No difference was found between recipients of normal allogeneic cells or tolerant allogeneic cells. Both showed a transient response but the response was maintained only in recipients of congenic cells.     

Use of PMSG antiserum in superovulated cattle?

Two Pregnant Mare Serum Gonadotrophin (PMSG) antisera were tested in 174 dairy cows that were superovulated with PMSG and were then given prostaglandin at 60 hours after PMSG. At 48 hours after injection of prostaglandin, the cows were given either PMSG antiserum (monoclonal (n=56) or polyclonal (n=57)), or saline as control (n=61). Ova (n=1,206) were recovered either nonsurgically or after slaughter. Of these, 757 were evaluated morphologically to be transferable embryos. A proportion of these embryos (n=295 from 52 flushed donors) were transferred to synchronized recipients and the pregnancy results were recorded. The reproductive function of 37 flushed donors was followed for 6 months after superovulation. No significant effect of the PMSG antisera could be demonstrated in any of the parameters studied (i.e., ovulation rate, number of follicles at collection, total yield of ova, fertilization rate, number of transferable embryos, pregnancy results after transfer of embryos, or period required by the donor cows for restitution of reproductive function after superovulation and recovery). It is concluded that use of PMSG antiserum did not improve the embryo yield in terms of the number and quality of transferable embryos or enhance normalization of reproductive function of the donor in the 6-month period after superovulation. Therefore, in an embryo transfer operation, the routine use of PMSG antiserum in a PMSG superovulation regimen in cattle is not recommended.     

How do subordinate and dominant species in semi-natural mountain grasslands relate to productivity and land-use change?

Changes in agricultural practices of semi-natural mountain grasslands are expected to modify plant community structure and shift dominance patterns. Using vegetation surveys of 11 sites in semi-natural grasslands of the Swiss Jura and Swiss and French Alps, we determined the relative contribution of dominant, subordinate and transient plant species in grazed and abandoned communities and observed their changes along a gradient of productivity and in response to abandonment of pasturing.The results confirm the humpbacked diversity–productivity relationship in semi-natural grassland, which is due to the increase of subordinate species number at intermediate productivity levels. Grazed communities, at the lower or higher end of the species diversity gradient, suffered higher species loss after grazing abandonment. Species loss after abandonment of pasturing was mainly due to a higher reduction in the number of subordinate species, as a consequence of the increasing proportion of dominant species.When plant biodiversity maintenance is the aim, our results have direct implications for the way grasslands should be managed. Indeed, while intensification and abandonment have been accelerated since few decades, our findings in this multi-site analysis confirm the importance of maintaining intermediate levels of pasturing to preserve biodiversity.     

Does restoration of a habitat‐forming seaweed restore associated faunal diversity?

Declines of habitat‐forming organisms in terrestrial and marine systems can lead to changes in community‐wide biodiversity. The dominant habitat‐forming macroalga Phyllospora comosa (Fucales) went locally extinct along the metropolitan coastline of Sydney in the 1980s. However, the consequences of that disappearance to the associated faunal diversity in these habitats, and whether Phyllospora is ecologically redundant with respect to the biodiversity it supports, are not known. Efforts are underway to restore Phyllospora, and the capacity to enhance local biodiversity is an important component of the rationale for restoration. We compared epifaunal diversity (abundances and composition) between Phyllospora and two other co‐occurring habitat‐forming algae, the kelp Ecklonia radiata and the fucoid Sargassum vestitum, and determined whether Phyllospora transplanted to Sydney developed different epifaunal communities than undisturbed thalli and controls. Where the 3 species naturally co‐occurred, Phyllospora supported different abundances of taxa than Ecklonia and Sargassum, as well as different composition at finer scales, which suggests that this species is not completely redundant and that its disappearance may have affected local biodiversity. Similarly, assemblages on transplanted Phyllospora differed from those on Ecklonia and Sargassum at restored sites, but did not always resemble assemblages from extant natural Phyllospora populations, even 18 months after transplantation. These experiments indicate that restoration of key habitat‐forming seaweeds not only recovers the algal species but also reduces risks of losing habitat diversity for epifauna and their consumers. However, restoration of all the original biodiversity associated with these seaweeds can be a difficult, complex, and long‐term process.     

Is livestock grazing compatible with amphibian diversity in the High Mountains of Córdoba, Argentina?

The High Mountains of Córdoba, Argentina have a long evolutionary history of grazing by large herbivores. However, about 400?years ago, European livestock were introduced and gradually replaced native herbivores. Since the 1920s, domestic herbivores have been the only large herbivores present in the area, causing severe soil erosion and a threat to the system diversity. The endemic fauna of the region includes four amphibian species. We evaluated the effect of livestock rearing on amphibian diversity and water bodies in woodlands and grasslands of the High Mountains of Córdoba. The work was conducted on stream stretches and ponds in two contrasting grazing situations: an area with livestock presence and another area where livestock was excluded 14?years ago. The following variables were recorded at each sampling site: amphibian richness and abundance, percentage of emergent, submerged and peripheral vegetation in areas surrounding the water bodies, water pH, and water dissolved O₂. No significant differences were detected in amphibian diversity between streams of both grazing situations, whereas a greater diversity (p?<?0.01) was observed in ponds in grazed grasslands. Our results suggest that livestock rearing, qualitatively measured as grazing and 14?years of livestock exclusion, in the study area would not have negative effects on amphibian diversity. This finding might be due to the long evolutionary grazing history of the area, large-scale livestock exclusion exhibiting a novel scenario. The ongoing reintroduction of native grazers may provide the benefits of grazing without the consequent soil erosion and habitat degradation associated with domestic livestock.     

Can a solitary avian species use collective detection? An assay in semi-natural conditions

Collective detection (e.g., enhanced predator detection through the vigilance of conspecifics) is expected to have evolved particularly in social species. However, we assessed the degree to which an avian territorial species (California towhee Pipilo crissalis) would use social cues about predation in a semi-natural assay. We also exposed a social species (house finch Carpodacus mexicanus) to similar conditions. California towhees increased scanning rates when foraging with conspecifics, whereas house finches increased scanning rates when foraging solitarily, suggesting that vigilance in these species is regulated mostly through interference competition and through predation risk, respectively. California towhees did not show early detection, and actually the last detector in the group delayed detection in relation to solitary individuals. House finches benefited from early detection, but the second and last detectors maintained detection at the level of solitary individuals. California towhees increased the chances of fleeing when in groups in relation to solitary conditions, but this effect was less pronounced in the last detector. House finches always fled across conditions. Overall, an asocial avian species may use collective detection, but limited to certain types of cues: responses were more pronounced to overt (conspecifics walking or fleeing) rather than subtle (conspecifics becoming alert or crouching) social cues.     

Can edaphic factors demonstrate landscape-scale differences in vegetation responses to grazing?

We focused on land units as landscape characteristics and selected seven typical land units on a land catena comprising two areas of southern Mongolia. Hierarchical analysis was used to test the hypothesis that a land unit’s edaphic factors could explain the differences in vegetation responses to grazing. We established the survey sites at increasing distances from a livestock camp or water point within each land unit, then analysed patterns of change in floristic and functional compositions, vegetation volume and soil properties within each land unit to reveal differences in vegetation responses to grazing. We also examined the variations in floristic and functional compositions across land units to identify the edaphic factors that may underlie these differences. Changes in vegetation and soil properties at increasing distances from a camp or water point within each land unit were into three different patterns. Ordination techniques consistently indicated that land unit groups categorised using edaphic factors corresponded to those categorised using response patterns. Our study revealed that edaphic factors were responsible for the observed landscape-scale differences in vegetation responses to grazing in the study areas. In addition, the mechanisms underlying vegetation responses to grazing may have been primarily determined by edaphic factors.