Even though mammalian herbivores can exert strong indirect effects on other animals by altering the vegetation, the study of trophic cascades retains a focus on apex predators and their top-down forces. Bottom-up trophic interaction chains induced by mammalian herbivores, particularly in invertebrate food webs, remain largely unexplored. We tested whether effects of mammalian herbivores on the vegetation ricochet back up several trophic levels of the invertebrate food web. We further tested two alternative hypotheses: the strength of herbivore-induced indirect interactions either increases with plant productivity because of a concurrent higher grazing intensity, or it decreases because of a higher plant tolerance to grazing. We progressively excluded large, medium and small herbivorous mammals from replicated plots of 6 m in productive, intensively grazed short-grass vegetation and less productive, less intensively grazed tall-grass vegetation of subalpine grasslands. We measured vegetation quantity, quality, structure and composition, and determined the abundance of invertebrate herbivores, detritivores, omnivores and predators. We used structural equation modelling to test vegetation-mediated cascading effects of the different mammalian herbivores across different trophic groups of invertebrates. In the short-grass vegetation, mammals caused changes in vegetation quantity and thickness. These changes directly affected detritivorous and predatory invertebrate abundance, yet indirectly affected predatory and omnivorous invertebrates through a bottom-up trophic cascade via changes in herbivorous invertebrate abundance. In the tall-grass vegetation, mammal-induced changes in vegetation quality and composition affected detritivorous invertebrates and in turn omnivorous invertebrates, but these cascading effects were weaker than those in the short-grass vegetation. Smaller mammals were at least as important as large mammals in structuring the invertebrate food web. Our results demonstrate that differently sized mammalian herbivores can trigger trophic cascades in the local invertebrate food web. Our findings further support the hypothesis that herbivore-induced indirect interactions are stronger in more productive systems because of higher foraging intensity, as opposed to the hypothesis that a higher grazing tolerance of plants should dampen herbivore-induced indirect interactions in productive systems.
The decline of open habitats in Europe, such as semi-natural grasslands and heathlands, has caused a general decline in biodiversity, which has been well documented for butterflies. Current conservation practices often involve grazing by domestic livestock to maintain suitable butterfly habitats. The extent to which wild ungulates may play a similar role remains largely unknown. Through their rooting activity, wild boar could be effective to reduce grass encroachment and restore pioneer microhabitats that are vital to many grassland insects in temperate climates. Here, we assessed the microhabitat requirements of Pyrgus malvae, an endangered butterfly of heathland and grassland habitats in the Netherlands, with special attention for the influence of wild boar rooting. To date, oviposition site selection of this species has concentrated on calcareous grasslands, whereas we also include heathlands. Overall, larval occupancy was higher in warm, open and sparsely vegetated microhabitats, which supports earlier findings. In heathland, microhabitat occupancy was positively affected by bryophyte and litter cover. In heath-grassland mosaic, microhabitat occupancy was also influenced by bryophyte and litter cover, but in addition low grass cover increased occupancy by favouring host plants. In grassland, only low grass cover and host plant cover determined microhabitat quality. Across all habitats, occupied microhabitats were characterized by lower vegetation as well as higher average daytime temperatures than unoccupied microhabitats. We discovered that wild boar play an important role in reducing grass cover by shallow rooting in grass patches, thereby increasing host plant availability. Hence, wild boar may have an added value in maintaining and restoring P. malvae microhabitats in grassland habitats in addition to grazing by domestic livestock.
Theory predicts that mammalian herbivores affect the quantity and quality of plants on which they preferentially feed in the short term. In the longer term, they can promote either preferred or less preferred plants, depending on whether preferred plants are adapted or sensitive to grazing. Less clear are the short‐ and long‐term responses of herbivorous insects to mammalian herbivory, and how these responses depend on the specific plants or plant functional types on which the insects feed. We progressively excluded large, medium and small mammals for five growing seasons in two subalpine vegetation types with long‐term differences in mammalian grazing intensity. Short‐grass vegetation has a history of intensive grazing, while tall‐grass vegetation has been grazed less intensively. We tested whether mammals altered the abundance and body size of leafhoppers specialized on specific plant functional types (grasses, sedges, forbs, or legumes/forbs), distinguishing between short‐term (exclosures) and long‐term (vegetation types) differences in mammalian grazing pressure. Furthermore, we assessed whether leafhoppers’ responses were explained by changes in biomass or quality of the plant functional types on which they feed. In the short term, mammal exclosures increased the abundance of grass‐ and forb‐feeding leafhoppers via increases in the biomass of grasses and forbs, regardless of vegetation type. Both grasses and forbs are preferred food plants of mammals. In the long term, the biomass of sedges, which are less preferred by mammals, increased in the less intensively grazed tall‐grass vegetation. This resulted in a higher abundance of sedge‐feeding leafhoppers. The small size of these sedge feeders lowered the average leafhopper body size in the tall‐grass vegetation. Plant nutritional quality did not explain any effects of exclusions or vegetation types. Our results demonstrate that both short‐ and long‐term effects of mammalian herbivores on the biomass of specific plant functional types caused concurrent changes in the abundance of specialized herbivorous insects, which scaled up to community‐wide shifts in insect body size, a key life‐history trait. A plant‐functional‐type approach can thus help to predict how overabundance or extinction of mammalian herbivores impacts on other components of the food web at various timescales. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.12989/suppinfo is available for this article.
We study pore-scale dynamics of reactive transport in heterogeneous, dual-porosity media, wherein a reactant in the invading fluid interacts chemically with the surface of the permeable grains, leading to the irreversible reaction A aq + B s → C aq . A microfluidic porous medium was synthesized, consisting of a single layer of hydrogel pillars (grains), chemically modified to contain immobilized enzymes on the grain surfaces. Fluorescence microscopy was used to monitor the spatiotemporal evolution of the reaction product C aq at different flow rates (Pećlet values) and to characterize the impact on its transport. The experimental setup enables delineation of three key features of the temporal evolution of the reaction product within the domain: (i) the characteristic time until the rate of C aq production reaches steady state, (ii) the magnitude of the reaction rate at steady state, and (iii) the rate at which C aq is flushed from the system. These features, individually, are found to be sensitive to the value of the Pećlet number, because of the relative impact of diffusion (vs advection) on the production and spatiotemporal evolution of C aq within the system. As the Pećlet number increases, the production of C aq is reduced and the transport becomes more localized within the vicinity of the grains. The dual-porosity feature causes the residence time of the transported species to increase, by forming stagnant zones and diffusive-dominant regions within the flow field, thus enhancing the reaction potential of the system. Using complementary numerical simulations, we explore these effects for a wider range of Pećlet and Damkoḧler numbers and propose nonlinear scaling laws for the key features of the temporal evolution of C aq .
Predatory protozoa play an essential role in shaping microbial populations. Among these protozoa, Acanthamoeba are ubiquitous in the soil and aqueous environments inhabited by Listeria monocytogenes . Observations of predator–prey interactions between these two microorganisms revealed a predation strategy in which Acanthamoeba castellanii assemble L. monocytogenes in aggregates, termed backpacks, on their posterior. The rapid formation and specific location of backpacks led to the assumption that A. castellanii may recruit L. monocytogenes by releasing an attractant. However, this hypothesis has not been validated, and the mechanisms driving this process remained unknown. Here, we combined video microscopy, microfluidics, single-cell image analyses, and theoretical modeling to characterize predator–prey interactions of A. castellanii and L. monocytogenes and determined whether bacterial chemotaxis contributes to the backpack formation. Our results indicate that L. monocytogenes captures are not driven by chemotaxis. Instead, random encounters of bacteria with amoebae initialize bacterial capture and aggregation. This is supported by the strong correlation between experimentally derived capture rates and theoretical encounter models at the single-cell level. Observations of the spatial rearrangement of L. monocytogenes trapped by A. castellanii revealed that bacterial aggregation into backpacks is mainly driven by amoeboid locomotion. Overall, we show that two nonspecific, independent mechanisms, namely random encounters enhanced by bacterial motility and predator surface-bound locomotion, drive backpack formation, resulting in a bacterial aggregate on the amoeba ready for phagocytosis. Due to the prevalence of these two processes in the environment, we expect this strategy to be widespread among amoebae, contributing to their effectiveness as predators.
Drift or downstream dispersal is a fundamental process in the life cycle of many riverine organisms. In the face of rapidly declining freshwater biodiversity, there is a need to enhance our capacity to study the drift of riverine organisms, by overcoming the limitations of traditional labour‐intensive sampling methods that result in data of low temporal and spatial resolution. To address this need, we developed a new technology, the Riverine Organism Drift Imager (RODI), which combines in situ imaging with machine‐learning classification. This technique expands on the traditional methodology by replacing the collection cup of a drift net with a camera system that continuously images riverine organisms as they drift through the device. After being imaged, organisms are released into the environment unharmed. A machine‐learning classifier is used after field sampling to identify drifting organisms. Therefore, RODI provides a non‐invasive sampling method that can quantify organism drift at unprecedented temporal resolution. Multiple deployments have served to validate the performance of the technology in the field. In its current implementation, images are captured continuously for 1.5 h at 50 frames per second. We demonstrate that the quality of the resulting images enables a convolutional neural network classifier to identify organisms to the family level. The weighted F1 score, a metric for the performance of the classifier, was 94%, based on training and testing on a field‐collected dataset consisting of 4598 images of 285 organisms belonging to seven classes (one species, five families and one order). In conclusion, this work provides a proof of concept, demonstrating the viability of the deployment of RODI as an automated, in situ organism drift sampler. This novel approach offers the possibility to advance our fundamental understanding of the drift of riverine organisms and how this is affected by human impacts in natural streams while, at the same time, can serve as a cost‐effective tool for biodiversity monitoring.
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