Additive and non‐additive effects of birch genotypic diversity on arthropod herbivory in a long‐term field experiment

Herbivores are important drivers of plant population dynamics and community composition in natural and managed systems. Intraspecific genetic diversity of long‐lived plants like trees might shape patterns of herbivory by different guilds of herbivores that trees experience through time. However, previous studies on plant genetic diversity effects on herbivores have been largely short‐term. We investigated how tree genotypic variation and diversity influence herbivory of silver birch Betula pendula in a long‐term field experiment. Using clones of eight genotypes, we constructed experimental plots consisting of one, two, four or eight genotypes, and measured damage by five guilds of arthropod herbivores twice a year over three different years (four, six and nine years after the experiment was established). Genotypes varied significantly for most types of herbivore damage, but genotype resistance rankings often shifted over time, and none of the clones was more resistant than all others to all types of herbivores. At the plot level, birch genotypic diversity had significant positive additive effect on leaf rollers and negative non‐additive effects on chewing herbivores and gall makers. In contrast, leaf‐mining and leaf‐tying damage was not influenced by birch genotypic diversity. Within diverse plots, the direction of genotypic diversity effects varied depending on birch genotype, some having lower and some having higher herbivory in mixed stands. This research highlights the importance of long‐term studies including different feeding guilds of herbivores to understand the effects of plant genetic diversity on arthropod communities. Different responses of various feeding guilds to genotypic diversity and shifts in resistance of individual genotypes over time indicate that genotypic mixtures are unlikely to result in overall reduction in herbivory over time.     

Community‐wide effects of below‐ground rhizobia on above‐ground arthropods

1. Plants take nutrients for their growth and reproduction from not only soil but also symbiotic microbes in the rhizosphere, and therefore below‐ground microbes may indirectly influence the above‐ground arthropod community through changes in the quality and quantity of plants. 2. Rhizobia are root‐nodulating bacteria that provide NH₄⁺ to legume plants. We examined bottom‐up effects of rhizobia on the community properties of the arthropods on host plants, using a root‐nodulating soybean strain (R+) and a non‐nodulating strain (R?) in a common garden. 3. R+ plants grew larger and produced a greater number of leaves than R? plants. We observed 28 species of herbivores and three taxonomic groups of predators on R+ and R? plants. The herbivorous species were classified into sap feeders (12 species) and chewers (16 species). 4. The species richness of overall herbivores, sap feeders, and chewers on R+ plants was greater than that on R? plants. Rhizobia positively affected the abundance of chewers. 5. The community composition of herbivores was significantly different between R? and R+ plants, although species diversity and evenness did not differ. 6. Rhizobia‐induced bottom‐up effects were transmitted to the third trophic level. The abundance, taxonomic richness, and diversity of the predators on R+ plants were greater but evenness was lower than those on R? plants. The community composition of predators was not affected by rhizobia. 7. These results indicate that the below‐ground microbes initiated bottom‐up effects on above‐ground herbivores and predators through trophic levels.     

Reversed effects of grazing on plant diversity: the role of below‐ground competition and size symmetry

Grazing is known as one of the key factors for diversity and community composition in grassland ecosystems, but the response of plant communities towards grazing varies remarkably between sites with different environmental conditions. It is generally accepted that grazing increases plant diversity in productive environments, while it tends to reduce diversity in unproductive habitats (grazing reversal hypothesis). Despite empirical evidence for this pattern the mechanistic link between modes of plant–plant competition and grazing response at the community level still remains poorly understood. Root‐competition in particular has rarely been included in theoretical studies, although it has been hypothesized that variations in productivity and grazing regime can alter the relative importance of shoot‐ and root‐competition. We therefore developed an individual‐based model based on plant functional traits to investigate the response of a grassland community towards grazing. Models of different complexity, either incorporating only shoot competition or with distinct shoot‐ and root‐competition, were used to study the interactive effects of grazing, resource availability, and the mode of competition (size‐symmetric or asymmetric). The pattern predicted by the grazing reversal hypothesis (GRH) can only be explained by our model if shoot‐ and root‐competition are explicitly considered and if size asymmetry of above‐ and symmetry of below‐ground competition is assumed. For this scenario, the model additionally reproduced empirically observed plant trait responses: erect and large plant functional types (PFTs) dominated without grazing, while frequent grazing favoured small PFTs with a rosette growth form. We conclude that interactions between shoot‐ and root‐competition and size symmetry/asymmetry of plant–plant interactions are crucial in order to understand grazing response under different habitat productivities. Our results suggest that future empirical trait surveys in grassland communities should include root traits, which have been largely ignored in previous studies, in order to improve predictions of plants’ responses to grazing.     

Crop identity and memory effects on aboveground arthropods in a long‐term crop rotation experiment

Agricultural landscapes are globally dominated by monocultures under intensive management. This is one of the main reasons for biodiversity loss and insect population decline in many regions all over the world. Agroecosystem biodiversity in these areas can be enhanced by cropping system diversification, such as crop rotations. Yet, long‐term studies on effects of crop rotations on aboveground agrobiodiversity are lacking. We set up a 10‐year long‐term crop rotation experiment in Central Germany and monitored the temporal dynamics of aboveground arthropods over a full cultivation period to investigate influence of current and preceding crop identity and cropping system diversification on activity density, species richness, and community structure. We found that species composition was strongly influenced by currently grown crop although effect on arthropods varied between species groups. Especially, winter oilseed rape strongly affects arthropod community structure. Interestingly, we were also able to show an influence of the preceding crops, indicating an ecological memory effect in the aboveground arthropod community. Our results show that crop identity of both currently and previously grown crops in crop rotations may lead to an increase in arthropod activity density and changes in species composition. Diversified crop rotations including appropriate crops can be an easily implemented tool to increase arthropod biodiversity and biomass at large spatial and temporal scales, particularly in areas dominated by a single crop (e.g., wheat, maize). Our results may help to design optimized crop rotations for large‐scale enhancement of insect biodiversity in agroecosystems.     

The effect of long‐term repeated burning and fire exclusion on above‐ and below‐ground Blackbutt (Eucalyptus pilularis) forest vegetation assemblages

We used a long‐term fire experiment in south‐east Queensland, Australia, to determine the effects of frequent prescribed burning and fire exclusion on understorey vegetation (<7.5 m) richness and density in Eucalyptus pilularis forest. Our study provided a point in time assessment of the standing vegetation and soil‐stored vegetation at two experimental sites with treatments of biennial burning, quadrennial burning since 1971–1972 and no burning since 1969. Vegetation composition, density and richness of certain plant groups in the standing and soil‐stored vegetation were influenced by fire treatments. The density of resprouting plants <3 m in height was higher in the biennially burnt treatment than in the unburnt treatment, but resprouters 3–7.5 m in height were absent from the biennial burning treatment. Obligate seeder richness and density in the standing vegetation was not significantly influenced by the fire treatments, but richness of this plant group in the seed bank was higher in the quadrennial treatment at one site and in the long unburnt treatment at the other site. Long unburnt treatments had an understorey of rainforest species, while biennial burning at one site and quadrennial burning at the other site were associated with greater standing grass density relative to the unburnt treatment. This difference in vegetation composition due to fire regime potentially influences the flammability of the standing understorey vegetation. Significant interactions between fire regime and site, apparent in the standing and soil‐stored vegetation, demonstrate the high degree of natural variability in vegetation community responses to fire regimes.     

Effect of burning and mowing on grass and forb diversity in a long‐term grassland experiment

Abstract. Disturbance may be an important determinant of plant community composition and diversity owing to its effects on competitive interactions, resources, dominance and vigour. The effect of type, timing and frequency of disturbance on grass and forb species richness was examined using data from a long‐term (> 50 yr) grassland burning and mowing experiment in KwaZulu‐Natal, South Africa. Grass species richness declined considerably (> 50%) in the absence of disturbance, whereas forb species richness was unaffected. Annual burning in sites not mown in summer tended to increase grass species richness relative to triennial burning (22% increase) with the reverse being true in sites mown twice in summer (37% decline). Forb species richness declined by 25% in sites mown twice in summer relative to sites mown in early summer only. Disturbance was necessary to achieve maximum grass species richness presumably by removing litter and increasing the availability of light. The interaction of time of mowing in summer (early versus late) and time of burning during the dormant period (spring versus winter) had the most dramatic effect on species richness. Time of burning had no effect on richness in sites mown in early summer, but winter burning resulted in a dramatic decline (27–42%) in richness in sites mown in late summer. This effect may be related to possible greater soil desiccation with this combination of disturbances.     

The long‐term adaptation of bacterial communities in metal‐contaminated sediments: a metaproteogenomic study

The aim of the study was to understand the effect of a long‐term metal exposure (110 years) on sediment microbial communities. Two freshwater sites, Férin and MetalEurop, differing by one order of magnitude in metal levels (MetalEurop: 3218 mg Zn kg^?1; 913 mg Pb kg^?1) were compared by shotgun metaproteogenomics. A total of 69–118 Mpb of DNA and 943–1241 proteins were obtained. Phymm BL analysis of the DNA sequences indicated that the phylogenetic profile was similar in both stations and that β‐Proteobacteria were dominant. However, subtle but significant changes were observed for some bacteria: e.g. Pseudomonas (+0.4%), Leptothrix (–0.4%), Thiobacillus (+0.36%) and Acidovorax (+0.48%). Using the stamp software, the two communities were found to be functionally very similar. However, significant genetic differences (10^?6 < P < 10^?3) were observed for three SEED categories: synthesis of exopolymeric substances, virulence and defence mechanisms (including czcA metal efflux genes), and elements involved in horizontal gene transfer. The CzcA protein was found by metaproteomics in MetalEurop, but the levels were too low to allow comparisons. It is concluded that bacterial communities in freshwater sediments may adapt to high metal levels without broad changes in the structure of the population.     

Plant‐mediated interactions among above‐ground and below‐ground life stages of a root‐feeding weevil

1. Interactions among herbivores mediated by plant responses to herbivore injury may have large impacts on herbivore population densities. Responses may persist for weeks after injury and may affect not only the initial (inducing) herbivore, but also herbivores that are spatially or temporally separated from the initial attacker. 2. In many plant–insect interactions, multiple life stages of the insect may be associated with the same plant, and these various stages may interact indirectly with one another via induced responses. The rice water weevil (RWW), Lissorhoptrus oryzophilus, a serious global pest of rice, is one such insect. A series of experiments were performed with root‐feeding larvae and leaf‐feeding adults of the RWW using three conventional rice varieties. 3. The first objective of this study was to test whether RWW adult feeding on rice leaves resulted in altered oviposition by subsequent adults. The hypothesis for the first objective was that RWW adult feeding would decrease plant suitability, resulting in reduced oviposition by subsequent adults. 4. The second objective was to test whether injury by RWW larvae to rice roots resulted in altered oviposition by subsequent adults. The hypothesis for the second objective was that below‐ground RWW larval feeding would decrease plant suitability of rice to above‐ground RWW adults, resulting in decreased oviposition. 5. Results provided inconsistent support for the first hypothesis, indicating that responses differed among combinations of variety and injury level. Conversely, consistent support for the second hypothesis was found, indicating that larval feeding on roots decreased suitability of rice plants for oviposition.     

Elevated temperature prolongs long‐term effects of a pesticide on Daphnia spp. due to altered competition in zooplankton communities

Considerable research efforts have been made to predict the influences of climate change on species composition in biological communities. However, little is known about how changing environmental conditions and anthropogenic pollution can affect aquatic communities in combination. We investigated the influence of short warming periods on the response of a zooplankton community to the insecticide esfenvalerate at a range of environmentally realistic concentrations (0.03, 0.3 and 3 μg L^?1) in 55 outdoor pond microcosms. Warming periods increased the cumulative water temperature, but did not exceed the maximum temperature measured under ambient conditions. Under warming conditions alone the abundance of some zooplankton taxa increased selectively compared to ambient conditions. This resulted in a shift in the community composition that had not recovered by the end of the experiment, 8 weeks after the last warming period. Regarding the pesticide exposure, short‐term effects of esfenvalerate on the community structure and the sensitive taxa Daphnia spp. did not differ between the two temperature regimes. In contrast, long‐term effects of esfenvalerate on Daphnia spp., a taxon that did not benefit from elevated temperatures, were observed twice as long under warming than under ambient conditions. This resulted in long‐term effects on Daphnia spp. until 4 months after contamination at 3 μg L^?1 esfenvalerate. Under both temperature regimes, we identified strength of interspecific competition as the mechanism determining the time until recovery. However, enhanced interspecific competition under warming conditions was prolonged and explained the delayed recovery of Daphnia spp. from esfenvalerate. These results show that, for realistic prediction of the combined effects of changing environmental factors and toxicants on sensitive taxa, the impacts of stressors on the biotic interactions within the community need to be considered.     

Responses of macroinvertebrate communities to long‐term flow variability in a Sonoran Desert stream

Current global models predict a hotter and drier climate in the southwestern United States with anticipated increases in drought frequency and severity coupled with changes in flash flood regimes. Such changes would likely have important ecological consequences, particularly for stream and riparian ecosystems already subject to frequent hydrologic disturbance. This study assessed the potential response of aquatic macroinvertebrates to interannual variation in hydrology in a spatially intermittent desert stream (Sycamore Creek, AZ). We compiled data on the recovery of macroinvertebrate communities following spring floods, with successional sequences captured 11 times over a 16‐year period (1983–1999). This period encompassed a transition from perennial to intermittent flow in this system, and included a record drought in 1989–1990. Results show that while the size of floods initiating sequences had little explanatory power, changes in macroinvertebrate community structure during postflood succession were closely associated with antecedent flooding and drought. Year‐to‐year differences in benthic communities integrated taxon‐specific responses to antecedent disturbance, including differential resistance to channel drying, use of hyporheic refugia, and variable rates of recovery once stream flow resumed. The long‐term consequences of drying on community structure were only evident during later stages of postflood succession, illustrating an interaction between flood and drought recovery processes in this system. Our observations highlight the potential for predicted climate changes in this region to have marked and long‐lasting consequences for benthic communities in desert streams.     

Functional composition of plant communities determines the spatial and temporal stability of soil microbial properties in a long‐term plant diversity experiment

Stable provisioning of ecosystem functions and services is crucial for human well‐being in a changing world. Two essential ecological components driving vital ecosystem functions in terrestrial ecosystems are plant diversity and soil microorganisms. In this study, we tracked soil microbial basal respiration and biomass over a time period of 12 years in a grassland biodiversity experiment (the Jena Experiment) and examined the role of plant diversity and plant functional group composition for the spatial and temporal stability of soil microbial properties (basal respiration and biomass) in bulk‐soil. Spatial and temporal stability were calculated as the inverse coefficient of variation (CV^?1) of soil microbial respiration and biomass measured from soil samples taken over space and time, respectively. We found that 1) plant species richness consistently increased soil microbial properties after a time lag of four years since the establishment of the experimental plots, 2) plant species richness had minor effects on the spatial stability of soil microbial properties, whereas 3) the functional composition of plant communities significantly affected spatial stability of soil microbial properties, with legumes and tall herbs reducing both the spatial stability of microbial respiration and biomass, while grasses increased the latter, and 4) the effect of plant diversity on temporal stability of soil microbial properties turned from being negative to neutral, suggesting that the recovery of soil microbial communities from former arable land‐use takes more than a decade. Our results highlight the importance of plant functional group composition for the spatial and temporal stability of soil microbial properties, and hence for microbially‐driven ecosystem processes, such as decomposition and element cycling, in temperate semi‐natural grassland.     

Effects of long‐term grazing management on sand dune vegetation of high conservation interest

Question: Can long‐term grazing management maintain and restore species‐rich sand dune plant communities within a sand dune site of high conservation interest? Location: Newborough Warren, North Wales, UK. Methods: Vegetation changes that occurred between 1987 and 2003, subsequent to grazing by domestic livestock being introduced to the site after decades with little or no stock grazing, were analysed using data collected from permanent monitoring quadrats over a 16‐year period. Results: At the plant community level, grazing brought about a shift from a tall‐grass dominated, species‐poor community to a more species‐rich community in the dry dunes, but did not change community type in dune slacks. However, at the species level, grazing enhanced the abundance of some desired perennial, annual and biennial species, graminoids and bryophytes in both habitat types. The increased frequency of positive indicator species for habitat condition suggests that grazing was beneficial for species of conservation interest. Ellenberg nitrogen (N) values decreased after grazing in dry habitats but showed no long‐term change independent of grazing, suggesting no increase in site fertility over the study period. Surprisingly, light (L) values also decreased in the dry dunes after grazing. Conclusions: Long‐term grazing management can play an important role for the conservation of dune communities and associated species. Because of its positive effects on species diversity, plant communities and habitat condition in sand dunes, livestock grazing is recommended for conservation management.     

Long‐term effects of exclusion of grazing stock on degraded herbaceous plant communities in a riparian Eucalyptus camaldulensis forest in south‐eastern Australia

Abstract Stock grazing has degraded many riparian ecosystems around the world. However, the potential for ecosystem recovery following the removal of grazing stock is poorly known. We developed a conceptual model to predict the responses of native and exotic herbaceous plants to grazing exclusion, based on site productivity and the degree of initial vegetation degradation. The effects of excluding grazing stock on richness, cover and composition of herbaceous plants were examined over 12 years in the degraded understorey of a riparian forest in Gulpa Island State Forest in south‐eastern Australia. We predicted that grazing exclusion would lead to limited changes in vegetation cover, richness and composition, owing to presumed low site productivity and the high degree of understorey degradation. Results showed that the cover, richness and composition of native and exotic species varied significantly among years. Over all plots, regions and years, total cover was slightly but significantly lower in grazed than in ungrazed plots (43.4% vs. 50.8%). While the cover of native plants increased over time in both treatments, the rate of increase was slightly greater in ungrazed plots. Grazing exclusion had no effect on the richness of native or exotic species, but had a significant but minor impact on plant composition, with different common species (mostly exotics) being promoted or diminished in ungrazed plots. The composition of grazed and ungrazed areas did not become more different over time. Overall, the effects that could be attributed to grazing exclusion were relatively minor and transient. Results are consistent with predictions based on site productivity and initial degradation, and should not be extrapolated to other more productive, or less degraded, riparian systems.     

Temperature mean and variance alter phytoplankton biomass and biodiversity in a long‐term microcosm experiment

Global predictions of increasing mean temperature and its variance result in concerns about the fate of biodiversity under future climatic conditions. On the local scale of interacting species, the response of diversity to warming will depend much on the change in interspecific interactions such as competition or predation. Using a phytoplankton model community, we conducted a 2 × 2 × 2 factorial long‐term microcosm experiment (16 months) with an underlying seasonal temperature profile. We manipulated the mean temperature (T_mean), the temperature variance (T_var), and the presence of consumers (ciliates) and monitored treatment effects on algal biomass, species composition and species richness. Temperature effects on algal biomass depended on the seasonal development and consumer presence. Algal biomass decreased with increasing T_mean at consumer absence, but increased at consumer presence. Overall, algal biomass decreased with increasing T_var. Consumer presence reduced algal biomass from summer to winter, but then ciliates and the consumer effect disappeared. Almost all treatment combinations collapsed to monocultures after 16 months, but extinction occurred faster at higher T_mean (especially at consumer absence) and slower at consumer presence (especially at higher T_mean). Contrasting our predictions, increasing T_var reduced richness and increased extinction rate. The treatment effects on biomass and richness were not independent, as algal species richness and biomass were positively correlated. Moreover, accelerated loss of species was consistently correlated to higher temporal variability in biomass. In conclusion, altered temperature regimes strongly affected algal biomass and diversity by interdependently altering competitive and consumer interactions.