Impact of herbivores on nitrogen cycling: contrasting effects of small and large species

Herbivores are reported to slow down as well as enhance nutrient cycling in grasslands. These conflicting results may be explained by differences in herbivore type. In this study we focus on herbivore body size as a factor that causes differences in herbivore effects on N cycling. We used an exclosure set-up in a floodplain grassland grazed by cattle, rabbits and common voles, where we subsequently excluded cattle and rabbits. Exclusion of cattle lead to an increase in vole numbers and a 1.5-fold increase in net annual N mineralization at similar herbivore densities (corrected to metabolic weight). Timing and height of the mineralization peak in spring was the same in all treatments, but mineralization in the vole-grazed treatment showed a peak in autumn, when mineralization had already declined under cattle grazing. This mineralization peak in autumn coincides with a peak in vole density and high levels of N input through vole faeces at a fine-scale distribution, whereas under cattle grazing only a few patches receive all N and most experience net nutrient removal. The other parameters that we measured, which include potential N mineralization rates measured under standardized laboratory conditions and soil parameters, plant biomass and plant nutrient content measured in the field, were the same for all three grazing treatments and could therefore not cause the observed difference. When cows were excluded, more litter accumulated in the vegetation. The formation of this litter layer may have added to the higher mineralization rates under vole grazing, through enhanced nutrient return through litter or through modification of microclimate. We conclude that different-sized herbivores have different effects on N cycling within the same habitat. Exclusion of large herbivores resulted in increased N annual mineralization under small herbivore grazing.     

Copepod feeding behaviour and egg production during a dinoflagellate bloom in the North Sea

Feeding strategies of copepods were studied during a dinoflagellate-dominated bloom in the North Sea in August 2001. The aim of this study was to evaluate the importance of mesozooplankton grazing as a biological loss factor of harmful algal blooms under natural conditions. Therefore, ingestion, egestion and egg production experiments were performed with the most abundant copepod species Calanus helgolandicus, Temora longicornis and Acartia sp. feeding on the natural phytoplankton community. Dinophysis norvegica and Ceratium furca were the most abundant dinoflagellate species at the time of the experiments. Grazing experiments as well as examination of fecal pellet content revealed C. helgolandicus fed efficiently on D. norvegica. Ingestion rates up to 47 cells female⁻¹ h⁻¹ were measured and a large proportion of the C. helgolandicus fecal pellets contained intact D. norvegica cells. Dinophysis cells were rarely seen in fecal pellets produced by T. longicornis, and never observed in pellets produced by Acartia sp. The ingestion rate of C. furca, which was the dominating Ceratium species, mimicked that of D. norvegica. C. helgolandicus grazed significantly on C. furca (16 cells female⁻¹ h⁻¹), while the ingestion rate of T. longicornis was low and Acartia sp. was not able to graze on C. furca. Egg production experiments revealed that 92% of the C. helgolandicus females produced eggs. The specific egg production rate and the proportion of females producing eggs among T. longicornis were low. This field experiment clearly shows that some copepod species feed efficiently on D. norvegica and C. furca under natural conditions, which may affect the bloom development of these dinoflagellates.     

In situ feeding rates and grazing impact of Mesopodopsis africana O. Tattersall in the St Lucia Estuary, South Africa

Mesopodopsis africana is an important mysid in southern African coastal zooplankton and a key species in the St Lucia estuarine lake, which is currently undergoing severe desiccation owing to freshwater deprivation. M. africana populations through much of the system are consequently under severe environmental stress. This study investigates the grazing dynamics of this mysid species, in relation to autotrophic food availability and other environmental constraints in two contrasting areas of the St Lucia Estuary, Charters Creek, heavily affected by the desiccation process and the Mouth, virtually under unchanged conditions. Gut evacuation experiments were conducted once each during the day and the night. Evacuation rates were consistently higher during the night, ranging from 0.27 to 0.33 h^-1 at Charters Creek and from 1.13 to 1.24 h^-1 at the Mouth. Ingestion rates were, therefore, higher at the Mouth resulting in population grazing impacts of 2.5% of the total microalgal biomass, while the grazing impact at Charters Creek was only 0.5%. The spatial variation in ingestion rates could be attributed to seasonal differences in gut evacuation rates, differences in the mean size of mysids used, or the physicochemical conditions present at the two stations. It is suggested that mysid populations at Charters Creek are predominantly driven by bottom-up forces, initiated by the harsh environmental conditions. Despite the lower ingestion rates exhibited at Charters Creek, results indicate that these mysids are capable of meeting all their energetic requirements from a microalgal diet alone, although they may also utilise a heterotrophic diet.     

Broad‐scale vegetation‐environment relationships in Eurasian high‐latitude areas

Question: How is tundra vegetation related to climatic, soil chemical, geological variables and grazing across a very large section of the Eurasian arctic area? We were particularly interested in broad‐scale vegetation‐environment relationships and how well do the patterns conform to climate‐vegetation schemes. Material and Methods: We sampled vegetation in 1132 plots from 16 sites from different parts of the Eurasian tundra. Clustering and ordination techniques were used for analysing compositional patterns. Vegetation‐environment relationships were analysed by fitting of environmental vectors and smooth surfaces onto non‐metric multidimensional scaling scattergrams. Results: Dominant vegetation differentiation was associated with a complex set of environmental variables. A general trend differentiated cold and continental areas from relatively warm and weakly continental areas, and several soil chemical and physical variables were associated with this broad‐scaled differentiation. Especially soil chemical variables related to soil acidity (pH, Ca) showed linear relationships with the dominant vegetation gradient. This was closely related to increasing cryoperturbation, decreasing precipitation and cooler conditions. Remarkable differences among relatively adjacent sites suggest that local factors such as geological properties and lemming grazing may strongly drive vegetation differentiation. Conclusions: Vegetation differentiation in tundra areas conforms to a major ecocline underlain by a complex set of environmental gradients, where precipitation, thermal conditions and soil chemical and physical processes are coupled. However, local factors such as bedrock conditions and lemming grazing may cause marked deviations from the general climate‐vegetation models. Overall, soil chemical factors (pH, Ca) turned out to have linear relationship with the broad‐scale differentiation of arctic vegetation.     

Infertile times: response to damage in genets of the clonal sedge Carex bigelowii

Tolerance to grazing is a plant trait that can be adaptive in systems where plants are subjected to a diversity of herbivore attack types. To test the tolerance ability of the clonal sedge Carex bigelowii, which is food plant to several herbivores in alpine and arctic areas, and the potential fitness costs of this tolerance, replicated units of genets were subjected to three levels of damage throughout three consecutive seasons. The three levels of treatment were no damage, light damage and heavy damage, and the damage was conducted by tearing off all plant material at 3 and 0 cm above-ground respectively. The genets had no tolerance under damage in terms of sexual reproduction. In terms of clonal reproduction the genets showed tolerance under light damage but not under heavy damage. However, no fitness cost was found for this tolerance ability, i.e. genets had higher reproduction and growth under no damage. The average ramet weight had a similar decrease under both a low and high damage treatment. Changed partitioning of biomass between plant parts and reduced concentration of total non-structural carbohydrates (TNC) in storage organs are possible mechanisms for the ability to uphold clonal reproduction in response to damage. There were no significant indications that tolerance ability or its fitness cost differed between genets. Our results suggest that when subjected to heavy damage genets will only reproduce vegetatively. Consequently, it seems C. bigelowii has evolved to allocate resources to the survival of an already successful genet rather than to a potential new genet of unknown success.     

Patterns of ash (Fraxinus excelsior L.) colonization in mountain grasslands: the importance of management practices

Woody colonization of grasslands is often associated with changes in abiotic or biotic conditions or a combination of both. Widely used as fodder and litter in the past traditional agro-pastoral system, ash (Fraxinus excelsior L.) has now become a colonizing species of mountain grasslands in the French Pyrenees. Its present distribution is dependent on past human activities and it is locally controlled by propagule pressure and abiotic conditions. However, even when all favourable conditions are met, all the potentially colonizable grasslands are not invaded. We hypothesize that management practices should play a crucial role in the control of ash colonization. From empirical field surveys we have compared the botanical composition of a set of grasslands (present and former) differing in management practices and level of ash colonization. We have displayed a kind of successional gradient positively linked to both ash cover and height but not to the age of trees. We have tested the relationships between ash presence in grassland and management types i.e. cutting and/or grazing, management intensity and some grassland communities’ features i.e. total and local specific richness and species heterogeneity. Mixed use (cutting and grazing) is negatively linked to ash presence in grassland whereas grazing alone positively. Mixed use and high grazing intensity are directly preventing ash seedlings establishment, when low grazing intensity is allowing ash seedlings establishment indirectly through herbaceous vegetation neglected by livestock. Our results show the existence of a limit between grasslands with and without established ashes corresponding to a threshold in the intensity of use. Under this threshold, when ash is established, the colonization process seems to become irreversible. Ash possesses the ability of compensatory growth and therefore under a high grazing intensity develops a subterranean vegetative reproduction. However the question remains at which stage of seedling development and grazing intensity these strategies could occur.     

Soil Properties and their Spatial Pattern in a Degraded Sandy Grassland under Post-grazing Restoration, Inner Mongolia, Northern China

In this study, we use classical and geostatistical methods to identify characteristics of some selected soil properties including soil particle size distribution, soil organic carbon, total nitrogen, pH and electrical conductivity and their spatial variation in a 5-year recovery degraded sandy grassland after two different grazing intensity disturbance: post-heavy-grazing restoration grassland (HGR) and post-moderately grazing restoration grassland (MGR), respectively, in Horqin steppe, Inner Mongolia, northern China. The objective was to examine effect of grazing intensity on spatial heterogeneity of soil properties. One hundred soil samples were taken from the soil layer 0–15 cm in depth of a grid of 10 m×10 m under each treatment. The results showed that soil fine fractions (very fine sand, 0.1–0.05 mm and silt + clay, <0.05 mm), soil organic carbon and total nitrogen concentrations were significant lower and their coefficients of variation significant higher under the HGR than under the MGR. Geostatistical analysis of soil heterogeneity revealed that soil particle size fractions, organic carbon and total nitrogen showed different degree of spatial dependence with exponential or spherical semivariograms on the scale measured under HGR and MGR. The spatial structured variance account for a large proportion of the sample variance in HGR plot ranging from 88% to 97% for soil particle fractions, organic C and total N, however, except for organic C (88.8%), the structured variance only account for 50% of the sample variance for soil particle fractions and total N in the MGR plot. The ranges of spatial autocorrelation for coarse-fine sand, very fine sand, silt + clay, organic C and total N were 13.7 m, 15.8 m, 15.2 m, 22.2 m and 21.9 m in HGR plot, respectively, and was smaller than in MGR plot with the corresponding distance of 350 m, 144.6 m, 45.7 m, 27.3 m and 30.3 m, respectively. This suggested that overgrazing resulted in an increase in soil heterogeneity. Soil organic C and total N were associated closely with soil particle fractions, and the kriging-interpolated maps showed that the spatial distribution of soil organic C and total N corresponded to the distribution patterns of soil particle fractions, indicating that high degree of spatial heterogeneity in soil properties was linked to the distribution of vegetative and bare sand patches. The results suggested that the degree of soil heterogeneity at field scale can be used as an index for indicating the extent of grassland desertification. Also, the changes in soil heterogeneity may in turn influence vegetative succession and restoration process of degraded sandy grassland ecosystem.     

Eutrophication-induced changes in benthic algae affect the behaviour and fitness of the marine amphipod Gammarus locusta

This study, conducted in mesocosms, natural field sites, and in laboratory aquaria, showed that eutrophication altered the nutrient status and dominance patterns among marine macroalgae, which in turn, stimulated gammaridean density. Gammaridean abundance correlated positively with both nutrient addition and the amount of green algae (also stimulated by nutrient enrichment). Path analysis indicated that the direct effect of nutrients on gammaridean density was of less importance than the indirect effect through increased production of green algae. In cage colonisation experiments, either in the field or in a control mesocosm kept under ambient nutrient conditions, more gammarids colonised nutrient enriched algae (E-algae) than algae with ambient nutrient levels (A-algae). Gammarus locusta generally grew faster on nutrient enriched algal specimens and when reared on green rather than on brown algae (fucoids). The nutrient status of periphytic algae did not affect gammaridean growth significantly, but the number of egg-carrying females (and thus egg production) was significantly higher among gammarids reared on E-periphyton. The gammaridean habitat preference order (red > green > brown > periphyton) was almost the reverse of their growth rate in feeding assays (periphyton > green > brown). This implies that macroalgae may be more important as a habitat than as a food source for these animals, which then have to become mobile in search of optimal food items. In this process, algal nutrient content was important as the gammarids in our study actively chose high quality nutrient-rich food, which, in addition, increased their fitness. Stimulated growth rates and egg production may ultimately lead to population increase, which, combined with the preference for high nutrient food items may dampen the initial effect of nutrient enrichment (i.e. blooms of green macroalgae) in shallow coastal waters.     

Population and clonal level responses of a perennial grass following fire in the northern Chihuahuan Desert

Relationships involving fire and perennial grasses are controversial in Chihuahuan Desert grasslands of southern New Mexico, USA. Research suggests that fire delays the resprouting of perennial grasses well after two growing seasons. However, such results are confounded by livestock grazing, soil erosion, and drought. Additionally, post-fire grass responses may depend on initial clone size. We evaluated the effects of fire, grazing, and clone size on Bouteloua eriopoda (black grama) in southern New Mexico grasslands. Four 2-ha plots were established in each of four sites. Fire and grazing were applied or not applied in 1999 such that four treatment combinations were assigned randomly to plots within each site. Within each plot, small (0–10 cm² basal area), medium (10–30 cm²), and large ( > 30 cm²) clones were initially mapped in five 0.91-m² quadrats where grass attributes and litter cover were evaluated before and at the end of two growing seasons following fire. Maximum fire temperature was also measured. At a population level, canopy and litter cover were each approximately 50% less in burned than unburned areas. However, compared to initial levels, canopy height had increased by 10% at the end of the study, regardless of fire. At a clonal level, basal cover reductions were attributed mostly to large clones that survived fire. Smaller clone densities had decreased by as much as 19% in burned compared to unburned areas, and fire reduced the basal cover of medium clones. Basal and canopy cover, recruitment, and clone basal area decreased with increased fire temperatures. Almost all responses were independent of grazing, and interactive effects of grazing and fire were not detected. Fire did not kill all perennial grass clones, regardless of size. However, rapid responses were likely influenced by above-average precipitation after fire. Future studies in desert grasslands should examine how perennial grass dynamics are affected by fire, precipitation patterns, and interactions with grazing.     

High Bird Species Diversity in Structurally Heterogeneous Farmland in Western Kenya

Tropical ecosystems are globally important for bird diversity. In many tropical regions, land‐use intensification has caused conversion of natural forests into human‐modified habitats, such as secondary forests and heterogeneous agricultural landscapes. Despite previous research, the distribution of bird communities in these forest‐farmland mosaics is not well understood. To achieve a comprehensive understanding of bird diversity and community turnover in a human‐modified Kenyan landscape, we recorded bird communities at 20 sites covering the complete habitat gradient from forest (near natural forest, secondary forest) to farmland (subsistence farmland, sugarcane plantation) using point counts and distance sampling. Bird density and species richness were on average higher in farmland than in forest habitats. Within forest and farmland, bird density and species richness increased with vegetation structural diversity, i.e., were higher in near natural than in secondary forest and in subsistence farmland than in sugarcane plantations. Bird communities in forest and farmland habitats were very distinct and very few forest specialists occurred in farmland habitats. Moreover, insectivorous bird species declined in farmland habitats whereas carnivores and herbivores increased. Our study confirms that tropical farmlands can hardly accommodate forest specialist species. Contrary to most previous studies, our findings show that structurally rich tropical farmlands hold a surprisingly rich and distinct bird community that is threatened by conversion of subsistence farmland into sugarcane plantations. We conclude that conservation strategies in the tropics must go beyond rain forest protection and should integrate structurally heterogeneous agroecosystems into conservation plans that aim at maintaining the diverse bird communities of tropical forest‐farmland mosaics.