Jan-Bernard Bouzillé

Effects of grazing by horses and/or cattle on the diversity of coastal grasslands in western France

In the coastal grasslands of western France, extensive mixed grazing by cattle and horses has been practised for many centuries. The vegetation of these old embanked grasslands varies along a topographical gradient with a hygrophilous plant community in low-lying depressions seasonally flooded, a mesophilous community on high level flats that are never flooded, and a meso-hygrophilous community on slopes where the soil remains saline. Recently, fewer horses have been grazed on these pastures and so a 6-year study was designed to investigate the effects of monospecific and mixed grazing by horses and cattle on plant community structure, composition and diversity. Mixed grazing produced the most species-rich and structurally diverse swards. Mixed grazing enhanced the development of rosette, sub-halophyte and halophyte species where the soil is saline, due to additive effects between the two herbivore species. Mixed grazing also limited the strongly competitive Elymus repens and Agrostis stolonifera on horse latrine areas, cattle grazing, thus showing a compensatory effect. The combination of additive and compensatory effects with mixed grazing could be used to manage plant diversity, heterogeneity in vegetation structure and communities of conservation value at the scale of the grassland ecosystem.

How Much does Grazing-Induced Heterogeneity Impact Plant Diversity in Wet Grasslands?

Abstract: The positive impact of grazing herbivores on plant diversity has been frequently reported in fertile grasslands. Grazing-induced heterogeneity is often evoked as an explanation for this influence. However, the relative importance of plant community heterogeneity induced by herbivores compared to other mechanisms linked to grazing remains unclear. We investigated this question by looking at 3 plant communities found in wet grasslands situated along the French Atlantic coast and traditionally grazed by horses and cattle. An experimental design set up in 1995 allowed us to compare the consequences of cattle-grazing, horse-grazing, and grazing abandonment on plant community diversity and heterogeneity. Floristic measurements made in 2007 showed that cattle and horses both had a positive impact on species richness and Shannon diversity index and that patchiness only occurred in grazed situations. The relative importance of grazing-induced patchiness within the overall positive effect of grazing on plant diversity was assessed by partitioning the diversity gain due to grazing into its additive within- and among-patch components. Grazing-induced patchiness entirely explained the increase in plant richness, whereas it accounted for only a small part of Shannon diversity. Grazing-related processes operating at the patch scale explained the main part of the increase in Shannon diversity. These processes make only a limited contribution to species recruitment, but they result in a more even species abundance distribution.

Spatial patterns of soil salinities in old embanked marshlands in western France

The spatial pattern of soil salinity was studied in commonly owned grazing meadows, embanked in the distant past (10th century), situated on the Atlantic coast of western France. The meadows were characterized by micro-relief, a relic of former salt marshes, with an amplitude that varied between 15 and 65 cm. The topographic sequence consisted of a mosaic of seasonally flooded depressions, of slopes, and of areas of flat, higher ground. The soils of the various topographic levels that were studied at 6 sites had different electrical conductivities and sodicities. The highest values occurred in slope soils (conductivity between 0.83 and 2.85 mS/cm), especially at the base of slopes, whereas the lowest values were typical of the soils of depressions (conductivity between 0.42 and 0.56 mS/cm). The deepest soil horizon (Cg) always had a higher sodicity and conductivity than that of the surface horizon at all topographic levels. The groundwater specific conductivity showed similar tendencies. A more intensive sampling of the spatial pattern of soil salinity at the Magnils-Reigniers site confirmed these results. In this sampling, only the conductivity of the soil-water extract was measured, as high correlations were shown at the 6 sites between the conductivity, sodicity, the exchangeable sodium content and soluble sodium content of the soil. The spatial pattern of salinity and sodicity can be related to the spatial structure of the vegetation, as salt-tolerant species (Juncus gerardi andHordeum marinum) occur on the slopes. This type of system does not seem to have been described elsewhere and appears to be related, via hydrologic mechanisms and the physical action of grazing animals, to the traditional management of the meadows for grazing.

Vegetation and ecological gradients in abandoned salt pans in western France

. Salt marshes along the Atlantic coast of France have been converted into solar salt pans since the 7th century. Salt production declined strongly from the 18th century onward, leading to the abandonment of many of these salt pans. High soil salinity is a residue from the original salt production and varies among salt pans according to time since abandonment, the current flooding regime and the effect of drainage measures. The relationships between the plant communities and seven environmental variables were investigated by Detrended Correspondence Analyses (DCA). Duration of flooding, electric conductivity and sodium saturation were most strongly related to variation in vegetation. The Heleo-chareto-Hippuridetum vulgaris and the Caricetum ripariae occur in lagoons with slightly saline soil that are flooded for the longest time period; the Scirpetum maritimi compacti occurs in salt pans with saline soil flooded for a long period; the Alopecuro-Juncetum gerardii occurs on saline soil where flooding is of short duration whereas the Carici-Lolietum perennis is never flooded and occurs on only slightly saline soil. Soil salinity and duration of flooding provide a satisfactory explanation of the variation in species composition in abandoned salt pans but land-use practices, especially grazing, have to be taken into account to fully understand their floristic composition.

After grazing exclusion, is there any modification of strategy for two guerrilla species: Elymus repens (L.) Gould and Agrostis stolonifera (L.)?

Elymus repens (L.) Gould and Agrostis stolonifera (L.), are competitive grasses with guerrilla strategy that invade grasslands with a low stocking rate. In this work, we tested the hypotheses that grazing exclusion facilitates vegetative development of rhizomes and stolons of these clonal grasses and that such change is a key mechanism for their abundance in set-aside grasslands. The competitive capacities of these two guerrilla species were characterised by samples in plant community (species richness and biomass) and on the level of individual species (morphometric measurements on stolons and rhizomes) during a growing season. Compared to grasslands where grazing was excluded for three years, species richness was higher in grazed site and the plant community structure differed. Indeed, with grazing exclusion, a shift from annual species with a diversified growth-form to perennial species with a tall tussock and graminoid growth-form was monitored. In ungrazed situation, Elymus repens and Agrostis stolonifera were the dominant grasses, and the standing biomass for the lowland community showed a significant increase compared to the grazed site. Vegetative development increased competitive capacities of these two guerrilla species and led by phenomenon of competitive exclusion to the disappearance of annuals species. With grazing cessation, Elymus repens was found to increase the size of aerial traits (shoot length and the number of leaves per shoot) and this may both be propitious for achieving dominance within plant communities and also maintaining its competitive local advantage. By contrast, Agrostis stolonifera showed an increase in a root trait, i.e. rhizome length, in the fenced site, which provide good ability for spatial propagation and then to explore adjacent patches. We concluded that Elymus repens presented a morphological capacity to change its colonising strategy from a guerrilla strategy to a phalanx strategy, by morphological variability of aerial organs, when it was submitted to competitive stress and environment modifications. Agrostis stolonifera showed a capacity to escape aerial competition resulting from grazing cessation, than to increase underground propagation capacity. The present study highlighted the capacities of Elymus repens to respond in an adaptative way to competitive pressure.

Responses of clonal architecture to experimental defoliation: a comparative study between ten grassland species

Clonal architecture may enable plants to effectively respond to environmental constraints but its role in plant tolerance to defoliation remains poorly documented. In several non-clonal species, modifications of plant architecture have been reported as a mechanism of plant tolerance to defoliation, yet this has been little studied in clonal plants. In a glasshouse experiment, five rhizomatous and five stoloniferous species of grazed pastures were subjected to three frequencies of defoliation in order to test two hypotheses. (1) We expected plant clonal response to defoliation to be either a more compact architecture (low clonal propagation, but high branching), or a more dispersed one (long-distance propagation and low branching). Such plastic adjustments of clonal architecture were assumed to be involved in tolerance to defoliation i.e. to promote genet performance in terms of biomass and number of ramets. (2) The response of clonal architecture to defoliation was expected to be dependent on the species and to be more plastic in stoloniferous than in rhizomatous species. Most genets of each species were tolerant to defoliation as they survived and developed in every treatment. Architectural modifications in response to defoliation did not match our predictions. Clonal growth was either maintained or reduced under defoliation. Relative growth rate (RGR) decreased in eight species, whereas defoliated genets of seven species produced as many ramets as control genets. Biomass allocation to ramet shoots remained stable for all but one species. In defoliated genets, the number and mean length of connections, and mean inter-ramet distance were equal to or lower than those in control genets. Four groups of species were distinguished according to their architectural response to defoliation and did not depend on the type of connections. We hypothesised that dense clonal architectures with low plasticity may be the most advantageous response in defoliated conditions such as in grazed pastures.

Clonal Growth Strategies Along Flooding and Grazing Gradients in Atlantic Coastal Meadows

Specific composition and species clonal traits were characterized along combined flooding and grazing gradients to answer two questions. i) To what extent does the interaction of flooding and grazing influence the clonal characteristics of the vegetation? ii) Are the effects of both environmental factors independent or interactive? This study was carried out in a wet meadow along the Atlantic coast (France). Three plant communities (hygrophilous, meso-hygrophilous and mesophilous) were distinguished along a flooding gradient and five levels of grazing pressure were controlled through an experimental design (from no grazing to heavy grazing). We monitored species composition and retrieved, for each species, the type of clonal growth organs (CGOs) and clonal traits from the CLO-PLA3 database. We identified two syndromes of clonal traits: “above-ground splitters” and “below-ground integrators”. Clonal traits played a key role in plant assembly in the studied meadows. The interaction of both environmental factors selected for particular syndromes of clonal traits; however, flooding had a stronger filtering effect than grazing. The hygrophilous community was dominated by above-ground splitters, whereas the meso-hygrophilous vegetation was dominated by below-ground integrators. In the mesophilous community, clonal composition was the most diverse and shared clonal traits with the vegetation of both the hygrophilous and meso-hygrophilous communities. Grazing impact on CGOs and clonal traits differed between plant communities, i.e., the effect of grazing was modulated by the flooding regime. This study confirmed that vegetation responses to grazing might depend on the pool of traits, primarily filtered by environmental factors such as flooding.

Growth patterns of Juncus gerardi clonal populations in a coastal habitat

Juncus gerardi populations demonstrated a logistic growth curve during the colonization stage. Shoot production by vegetative multiplication was virtually continuous from December to June. Experiments suggested that the stabilisation stage of the demographic curve reflected water deficit. Taller, fertile, winter and early spring cohorts could be distinguished from shorter, infertile end of spring and beginning of summer cohorts. Shoot emergence began in March and terminated at the end of June, when water becomes a limiting factor due to a period of water shortage, typical of the thermo-atlantic climate. Spatial extension of populations was due to rhizome growth, which ceased during flowering.Flowering in May temporarily checked growth in shoot height of all emerged cohorts. No cost of reproduction was demonstrated concerning the rate of appearance of new shoots.Although fertile shoots were taller than vegetative shoots, their growth rates were significantly lower from April onwards. The tallest fertile shoots produced the most capsules.Energy allocation to seed production is the only possible means for long-term establishment of new genotypes, and vegetative multiplication appears as the principal source of recruitment of new modules in Juncus gerardi.Resource allocation patterns in this clonal species are discussed in relation to the ecological background in the concerned marshlands and with theoretical proposals derived from models of spatial colonization strategies in clonal plants.Nomenclature: follows Flora Europaea (Tutin et al., 1964ndash;1980).

Satellite‐derived vegetation indices as surrogate of species richness and abundance of ground beetles in temperate floodplains

Remotely sensed data are frequently employed for monitoring vegetation and for estimating herbivore diversity. Their use for predicting predator arthropod species abundance and richness has also been investigated with success for ants and beetles in forests using normalised difference vegetation index (NDVI) and for beetles in mountain forests using light detection and ranging data. We investigated whether vegetation indices, derived from multispectral SPOT imagery could predict abundance and species richness of ground active spiders and ground beetles in a new ecological context, the floodplain meadows of the Loire River in Western Europe. Using pitfall traps, we collected carabids and spiders in the field. Maximum vegetation height, litter‐depth and plant species richness best explained species assemblages of both groups (multivariate analyses). NDVI and enhanced vegetation index (EVI 2) were strongly related to activity‐density and species richness for ground beetles only, EVI 2 being the best surrogate. Relationships between vegetation indices and spider assemblage patterns were either non‐significant or weak. We demonstrated that EVI 2 is a good surrogate of the abundance and richness of carabid species in a temperate floodplain, and has potential as a low cost method for mapping arthropod assemblages at large spatial scales. Our approach provides a tool which contributes to biodiversity assessment at large spatial scales. It can also contribute to the prioritisation of conservation areas and early change detection, as carabids are keystone indicators.

Impact of selective grazing on plant production and quality through floristic contrasts and current-year defoliation in a wet grassland

Grazing impacts the structure and functional properties of vegetation through floristic changes (i.e., long-term effect) and current defoliation (i.e., short-term effect). The aim of this study was to assess the relative importance of these two grazing effects on productivity (ANPP) and plant quality (C/N ratio) among plant patches submitted to a variety of grazing intensity for several years. Long-term grazing effect was measured by comparing ANPP and C/N ratio among plant patches with contrasting floristic composition. Short-term impact of grazing was measured by comparing ANPP and C/N in plant patches, with and without defoliation. Floristic contrasts led to a lower ANPP in highly grazed patches than in lightly grazed ones. This result may be related to the increasing proportion of grazing-tolerant and grazing-avoiding species with increasing grazing intensity. Vegetation C/N contrasts were recorded among grazed patches but did not linearly relate to grazing intensity. Short-term effect of current-year defoliation on ANPP was limited as vegetation compensated for biomass removal. No evidence for grazing-enhancement of ANPP was found even at moderate grazing intensity. Long-term floristic changes with grazing thus appeared to be the main driving factor of variations in ANPP. In contrast, C/N ratio showed no general and consistent variation along the grazing gradient but varied consistently depending on the community investigated, thus suggesting an effect of the species pool available.