Organic farming and landscape structure: effects on insect-pollinated plant diversity in intensively managed grasslands

Parallel declines in insect-pollinated plants and their pollinators have been reported as a result of agricultural intensification. Intensive arable plant communities have previously been shown to contain higher proportions of self-pollinated plants compared to natural or semi-natural plant communities. Though intensive grasslands are widespread, it is not known whether they show similar patterns to arable systems nor whether local and/or landscape factors are influential. We investigated plant community composition in 10 pairs of organic and conventional dairy farms across Ireland in relation to the local and landscape context. Relationships between plant groups and local factors (farming system, position in field and soil parameters) and landscape factors (e.g. landscape complexity) were investigated. The percentage cover of unimproved grassland was used as an inverse predictor of landscape complexity, as it was negatively correlated with habitat-type diversity. Intensive grasslands (organic and conventional) contained more insect-pollinated forbs than non-insect pollinated forbs. Organic field centres contained more insect-pollinated forbs than conventional field centres. Insect-pollinated forb richness in field edges (but not field centres) increased with increasing landscape complexity (% unimproved grassland) within 1, 3, 4 and 5km radii around sites, whereas non-insect pollinated forb richness was unrelated to landscape complexity. Pollination systems within intensive grassland communities may be different from those in arable systems. Our results indicate that organic management increases plant richness in field centres, but that landscape complexity exerts strong influences in both organic and conventional field edges. Insect-pollinated forb richness, unlike that for non-insect pollinated forbs, showed positive relationships to landscape complexity reflecting what has been documented for bees and other pollinators. The insect-pollinated forbs, their pollinators and landscape context are clearly linked. This needs to be taken into account when managing and conserving insect-pollinated plant and pollinator communities.     

Shrinking by numbers: landscape context affects the species composition but not the quantitative structure of local food webs

1. With habitat fragmentation spreading around the world, there is a pressing need to understand its impacts on local food webs. To date, few studies have examined the effects of landscape context on multiple local communities in a quantitative, spatially realistic setting. 2. To examine how the isolation of a food web affects its structure, we construct local food webs of specialist herbivores and their natural enemies on 82 individual oaks (Quercus robur) growing in different landscape contexts. 3. Across this set of webs, we find that communities in isolated habitat patches not only contained fewer species than did well-connected ones, but also differed in species composition. 4. Surprisingly, the effects observed in terms of species composition were not reflected in the quantitative interaction structure of local food webs: landscape context had no detectable effect on either the interaction evenness, linkage density, connectance, generality or vulnerability of local webs. 5. We conclude that the quantitative structure of food webs may be stable in the face of habitat fragmentation, despite clear-cut impacts on individual species. This finding offers hope-inspiring news for conservation, but should clearly be verified by empirical studies across both naturally and more recently fragmented systems.     

Decreases in Fire Spread Probability with Forest Age Promotes Alternative Community States, Reduced Resilience to Climate Variability and Large Fire Regime Shifts

The generalization that plant communities increase in flammability as they age and invariably lead to resilient self-organized landscape mosaics is being increasingly challenged. Plant communities often exhibit rapidly saturating or even hump-shaped age-flammability trajectories and landscapes often display strong non-linear behaviors, abrupt shifts, and self-reinforcing alternative community states. This plethora of fire-landscape interactions calls for a more general model that considers alternative age-flammability rules. We simulated landscape dynamics assuming communities that (1) increase in flammability with age and (2) gain flammability up to a certain age followed by a slight and moderate loss to a constant value. Simulations were run under combinations of ignition frequency and interannual climatic variability. Age-increasing fire probability promoted high resilience to changes in ignition frequency and climatic variability whereas humpbacked-shaped age-flammability led to strong non-linear behaviors. Moderate (20%) reductions in mature compared to peak flammability produced the least resilient behaviors. The relatively non-flammable mature forest matrix intersected by young flammable patches is prone to break up and disintegrate with slight increases in ignition/climate variability causing large-scale shifts in the fire regime because large fires were able to sweep through the more continuous young/flammable landscape. Contrary to the dominant perception, fire suppression in landscapes with positive feedbacks may effectively reduce fire occurrence by allowing less flammable later stage communities composed of longer lived, obligate seeders to replace earlier stages of light demanding, often more flammable resprouters. Conversely, increases in anthropogenic ignitions, a common global trend of many forested regions may, in synergism with increased climate variability, induce abrupt shifts, and large-scale forest degradation.     

Cascading effects of climate change on plankton community structure

Plankton communities account for at least half of global primary production and play a key role in the global carbon cycle. Warming and acidification may alter the interaction chains in these communities from the bottom and top of the food web. Yet, the relative importance of these potentially complex interactions has not yet been quantified. Here, we examine the isolated and combined effects of warming, acidification, and reductions in phytoplankton and predator abundances in a series of factorial experiments. We find that warming directly impacts the top of the food web, but that the intermediate trophic groups are more strongly influenced by indirect effects mediated by altered top‐down interactions. Direct manipulations of predator and phytoplankton abundance reveal similar strong top‐down interactions following top predator decline. A meta‐analysis of published experiments further supports the conclusion that warming has stronger direct impacts on the top and bottom of the food web rather than the intermediate trophic groups, with important differences between freshwater and marine plankton communities. Our results reveal that the trophic effect of warming cascading down from the top of the plankton food web is a powerful agent of global change.     

Effects of Multi-chain Omnivory on the Strength of Trophic Control in Lakes

Omnivory has been implicated in both diffusing and intensifying the effects of consumer control in food chains. Some have postulated that the strong, community level, top-down control apparent in lakes is not expressed in terrestrial systems because terrestrial food webs are reticulate, with high degrees of omnivory and diverse plant communities. In contrast, lake food webs are depicted as simple linear chains based on phytoplankton-derived energy. Here, we explore the dynamic implications of recent evidence showing that attached algal (periphyton) carbon contributes substantially to lake primary and secondary productivity, including fish production. Periphyton production represents a cryptic energy source in oligotrophic and mesotrophic lakes that is overlooked by previous theoretical treatment of trophic control in lakes. Literature data demonstrate that many fish are multi-chain omnivores, exploiting food chains based on both littoral and pelagic primary producers. Using consumer-resource models, we examine how multiple food chains affect fourth-level trophic control across nutrient gradients in lakes. The models predict that the stabilizing effects of linked food chains are strongest in lakes where both phytoplankton and periphyton contribute substantially to production of higher trophic levels. This stabilization enables a strong and persistent top down control on the pelagic food chain in mesotrophic lakes. The extension of classical trophic cascade theory to incorporate more complex food web structures driven by multi-chain predators provides a conceptual framework for analysis of reticulate food webs in ecosystems.     

Benthic suspension feeders: their paramount role in littoral marine food webs

In recent years, particular attention has been paid to coupling and energy transfer between benthos and plankton. Because of their abundance, certain benthic suspension feeders have been shown to have a major impact in marine ecosystems. They capture large quantities of particles and might directly regulate primary production and indirectly regulate secondary production in littoral food chains. Suspension feeders develop dense, three-dimensional communities whose structural complexity depends on flow speed. It has been postulated that these communities can self-organize to enhance food capture and thus establish boundary systems capable of successfully exploiting a less structured system, namely, the plankton.     

An abstract cell model that describes the self-organization of cell function in living systems

The principal aim of systems biology is to search for general principles that govern living systems. We develop an abstract dynamic model of a cell, rooted in Mesarovi? and Takahara's general systems theory. In this conceptual framework the function of the cell is delineated by the dynamic processes it can realize. We abstract basic cellular processes, i.e., metabolism, signalling, gene expression, into a mapping and consider cell functions, i.e., cell differentiation, proliferation, etc. as processes that determine the basic cellular processes that realize a particular cell function. We then postulate the existence of a 'coordination principle' that determines cell function. These ideas are condensed into a theorem: If basic cellular processes for the control and regulation of cell functions are present, then the coordination of cell functions is realized autonomously from within the system. Inspired by Robert Rosen's notion of closure to efficient causation, introduced as a necessary condition for a natural system to be an organism, we show that for a mathematical model of a self-organizing cell the associated category must be cartesian closed. Although the semantics of our cell model differ from Rosen's (M,R)-systems, the proof of our theorem supports (in parts) Rosen's argument that living cells have non-simulable properties. Whereas models that form cartesian closed categories can capture self-organization (which is a, if not the, fundamental property of living systems), conventional computer simulations of these models (such as virtual cells) cannot. Simulations can mimic living systems, but they are not like living systems.     

Refuge increases food chain length: modeled impacts of littoral structure in lake food webs

Food chain length (FCL) represents a fundamental metric within ecology because it has implications for ecosystem function and responses to environmental change. Omnivory between linked food chains situated within large ecosystems can increase FCL, whereas overlap of food chains within small or spatially compressed ecosystems is generally thought to decrease FCL. Yet FCL varies widely in small ecosystems and the mechanisms underlying determinants of FCL in these systems is unclear. In small shallow lakes, littoral structure is a predictor of FCL but it is unclear whether this is due to productivity or refuge mechanisms. Here we provide evidence, using consumer resource food web modules parameterized with empirical data, that refuge in spatially compressed ecosystems has the ability on its own to increase the trophic position of top predators by increasing the biomass of top and intermediate predators across a range of common food web module structures. Our results suggest that refuge is an important driver of FCL in small ecosystems, which has implications for determining responses of these systems to environmental change.     

Are Antarctic suspension-feeding communities different from those elsewhere in the world?

This paper reviews the trophic ecology of benthic suspension feeders in Antarctic shelf communities, studied within SCAR's EASIZ Programme, in comparison with published information from other seas. Dense benthic suspension-feeder communities capture large quantities of particles and may directly regulate primary, and indirectly, secondary production in littoral food chains. Most work has been performed in temperate and tropical seas; however, little is known about suspension feeders in cold environments. Recent studies on Antarctic littoral benthic suspension feeders suggest the period of winter inactivity may last only a few weeks. This contrasts with the hypothesis that in Antarctic communities there is a prolonged period of minimal activity lasting at least 6 months during the austral winter. Results from other oceans may explain how dense benthic communities could develop under such conditions. Alternative food sources, i.e. the "fine fraction", sediment resuspension, lateral advection and efficient food assimilation may play a significant role in the development of suspension-feeder dominated, very diversified, high biomass and three-dimensionally structured communities on the Antarctic shelf.     

Landscape-level effects on cynipid component communities of “orphaned” native shrubs

Native plants that persist in agricultural landscapes can be important for conserving and re-connecting fragmented biological communities, particularly to arthropods that live in and on them. However, their value as habitat may depend upon landscape context, which can differently impact species among trophic levels. We examined the communities of gall-inducing cynipid wasps and their parasitoids associated with a native wild rose, Rosa woodsii, in the Palouse region of southeastern Washington State and adjacent Idaho, to determine how this arthropod community varies with the landscape context of the host plant. Nine species of gall wasps (Diplolepis spp.), 11 species of parasitoids, and the inquiline cynipid, Periclistus sp. were sampled from R. woodsii shrubs throughout the Palouse region. We examined whether any gall-inducing cynipids were absent from R. woodsii in particular landscape contexts. We tested the relationship between community structure and landscape variables including landscape diversity and the proportion of prairie, agriculture, introduced grasslands, and built environments. All gall-inducer species occurred in every landscape context, although some species were more common on roses within prairie remnants. Gall-inducer and parasitoid species richness was positively correlated with the proportion of prairie in the landscape at radii from 100 to 1,500 m, with the scale of this effect differing between the two sample years. Landscape diversity had little or no effect on species richness. These results suggest that although R. woodsii supports diverse arthropod communities throughout a fragmented landscape, its greatest conservation potential will be realized if prairie remnants are conserved as well.     

Inter‐patch movement in an experimental system: the effects of life history and the environment

An important process for the persistence of populations subjected to habitat loss and fragmentation is the dispersal of individuals between habitat patches. Dispersal involves emigration from a habitat patch, movement between patches through the surrounding landscape, and immigration into a new suitable habitat patch. Both landscape and physical condition of the disperser are known to influence dispersal ability, although disentangling these effects can often be difficult in the wild. In one of the first studies of its kind, we used an invertebrate model system to investigate how dispersal success is affected by the interaction between the habitat condition, as determined by food availability, and life history characteristics (which are also influenced by food availability). Dispersal of juvenile and adult mites (male and female) from either high food or low food natal patches were tested separately in connected three patch systems where the intervening habitat patches were suitable (food supplied) or unsuitable (no food supplied). We found that dispersal success was reduced when low food habitat patches were coupled to colonising patches via unsuitable intervening patches. Larger body size was shown to be a good predictor of dispersal success, particularly when the intervening landscape is unsuitable. Our results suggest that there is an interaction between habitat fragmentation and habitat suitability in determining dispersal success: if patches degrade in suitability and this affects the ability to disperse successfully then the effective connectance across landscapes may be lowered. Understanding these consequences will be important in informing our understanding of how species, and the communities in which they are embedded, may potentially respond to habitat fragmentation.     

Negative effects of pesticides on wild bee communities can be buffered by landscape context

Wild bee communities provide underappreciated but critical agricultural pollination services. Given predicted global shortages in pollination services, managing agroecosystems to support thriving wild bee communities is, therefore, central to ensuring sustainable food production. Benefits of natural (including semi-natural) habitat for wild bee abundance and diversity on farms are well documented. By contrast, few studies have examined toxicity of pesticides on wild bees, let alone effects of farm-level pesticide exposure on entire bee communities. Whether beneficial natural areas could mediate effects of harmful pesticides on wild bees is also unknown. Here, we assess the effect of conventional pesticide use on the wild bee community visiting apple (Malus domestica) within a gradient of percentage natural area in the landscape. Wild bee community abundance and species richness decreased linearly with increasing pesticide use in orchards one year after application; however, pesticide effects on wild bees were buffered by increasing proportion of natural habitat in the surrounding landscape. A significant contribution of fungicides to observed pesticide effects suggests deleterious properties of a class of pesticides that was, until recently, considered benign to bees. Our results demonstrate extended benefits of natural areas for wild pollinators and highlight the importance of considering the landscape context when weighing up the costs of pest management on crop pollination services.     

Trophic assimilation efficiency markedly increases at higher trophic levels in four-level host–parasitoid food chain

Trophic assimilation efficiency (conversion of resource biomass into consumer biomass) is thought to be a limiting factor for food chain length in natural communities. In host–parasitoid systems, which account for the majority of terrestrial consumer interactions, a high trophic assimilation efficiency may be expected at higher trophic levels because of the close match of resource composition of host tissue and the consumer''s resource requirements, which would allow for longer food chains. We measured efficiency of biomass transfer along an aphid-primary–secondary–tertiary parasitoid food chain and used stable isotope analysis to confirm trophic levels. We show high efficiency in biomass transfer along the food chain. From the third to the fourth trophic level, the proportion of host biomass transferred was 45%, 65% and 73%, respectively, for three secondary parasitoid species. For two parasitoid species that can act at the fourth and fifth trophic levels, we show markedly increased trophic assimilation efficiencies at the higher trophic level, which increased from 45 to 63% and 73 to 93%, respectively. In common with other food chains, δ¹⁵N increased with trophic level, with trophic discrimination factors (Δ¹⁵N) 1.34 and 1.49‰ from primary parasitoids to endoparasitic and ectoparasitic secondary parasitoids, respectively, and 0.78‰ from secondary to tertiary parasitoids. Owing to the extraordinarily high efficiency of hyperparasitoids, cryptic higher trophic levels may exist in host–parasitoid communities, which could alter our understanding of the dynamics and drivers of community structure of these important systems.     

Developing sustainable food supply chains

This paper reviews the opportunities available for food businesses to encourage consumers to eat healthier and more nutritious diets, to invest in more sustainable manufacturing and distribution systems and to develop procurement systems based on more sustainable forms of agriculture. The important factors in developing more sustainable supply chains are identified as the type of supply chain involved and the individual business attitude to extending responsibility for product quality into social and environmental performance within their own supply chains. Interpersonal trust and working to standards are both important to build more sustainable local and many conserved food supply chains, but inadequate to transform mainstream agriculture and raw material supplies to the manufactured and commodity food markets. Cooperation among food manufacturers, retailers, NGOs, governmental and farmers' organizations is vital in order to raise standards for some supply chains and to enable farmers to adopt more sustainable agricultural practices.     

Relative contribution of agroforestry,rainforest and openland to local and regional bee diversity

Due to increasing human modification of tropical landscapes, the relative importance of natural habitats and agricultural systems has become a major conservation topic to counteract global species loss. We investigated the contribution of tropical primary forest, cacao agroforestry systems of varying management practices and openland to the temporal and spatial variation of diversity of native bee communities in the herb layer (Apidae, Hymenoptera) in Sulawesi (Indonesia). Local bee density and diversity were highest in openland, followed by agroforestry systems and were lowest in primary forests, revealing the importance of herbaceous food resources in the understorey. In contrast, highest regional bee richness was found in agroforestry systems, because of high community dissimilarity. Multidimensional scaling supported these findings with openland habitats showing more compactly clustered bee species communities than agroforestry habitats. In conclusion, the herb associated bee community profited from the opening of the landscape as a result of agricultural activities, while agroforestry systems increased bee species richness especially on a regional scale due to high management diversity.     

Yield and dynamics of tritrophic food chains

Strong relationships between yield and dynamic behavior of tritrophic food chains are pointed out by analyzing the classical Rosenzweig-MacArthur model. On the one hand, food chains are subdivided into undersupplied and oversupplied categories, the first being those in which a marginal increase of nutrient supply to the bottom produces a marginal increase of mean yield at the top. On the other hand, a detailed bifurcation analysis proves that dynamic complexity first increases with nutrient supply (from stationary to a low-frequency cyclic regime and, finally, to chaos) and then decreases (from chaos to a high-frequency cyclic regime). A careful comparison of the two analyses supports the conclusion that food chains cycling at high frequency are oversupplied, while all others are undersupplied. A straightforward consequence of this result is that maximization of food yield requires a chaotic regime. This regime turns out to be very often on the edge of a potential catastrophic collapse of the top component of the food chain. In other words, optimality implies very complex and dangerous dynamics, as intuitively understood long ago for ditrophic food chains by Rosenzweig in his famous article on the paradox of enrichment.     

Climate change enhances the negative effects of predation risk on an intermediate consumer

Predators are a major source of stress in natural systems because their prey must balance the benefits of feeding with the risk of being eaten. Although this ‘fear’ of being eaten often drives the organization and dynamics of many natural systems, we know little about how such risk effects will be altered by climate change. Here, we examined the interactive consequences of predator avoidance and projected climate warming in a three‐level rocky intertidal food chain. We found that both predation risk and increased air and sea temperatures suppressed the foraging of prey in the middle trophic level, suggesting that warming may further enhance the top‐down control of predators on communities. Prey growth efficiency, which measures the efficiency of energy transfer between trophic levels, became negative when prey were subjected to predation risk and warming. Thus, the combined effects of these stressors may represent an important tipping point for individual fitness and the efficiency of energy transfer in natural food chains. In contrast, we detected no adverse effects of warming on the top predator and the basal resources. Hence, the consequences of projected warming may be particularly challenging for intermediate consumers residing in food chains where risk dominates predator‐prey interactions.     

Connecting landscape structure and patterns in body size distributions

Understanding the interaction between community structure and landscape structure represents a pressing theoretical challenge of great applied importance considering the increasing structural modification of ecosystems through habitat loss and fragmentation. Dispersal ability and energetic demands coupled to body size determine the landscape structure experienced by an organism, which could essentially be fragmented for small individuals but continuous for large ones. Although discontinuities in species assemblages have been predicted and detected, no explicit association between habitat structure and body size distributions has been demonstrated. In this contribution, we propose that body size structure in local communities should reflect such different perceptions of landscape structure. To this end, we explore this association in a simple metacommunity located in the Atacama Desert, in northern Chile. Using graph theory we found that species of different size and trophic position (carnivores and herbivores) perceive the landscape at contrasting spatial scales. In each community (n = 31) we determined the observed and the expected body size distributions – in a random sample from the metacommunity of 18 727 individuals –, which allowed us to identify the body sizes at which an overrepresentation or underrepresentation of individuals occur. Such aggregations and discontinuities in body sizes were related, for carnivores, to patch location within the landscape, and to the internal banded vegetation pattern within patches for herbivores. Our study shows, for the first time, an empirical connection between the spatial distribution of communities, their local attributes, and the existence and locations of discontinuities and aggregations in body size distributions.     

Modularity along organism dispersal gradients challenges a prevailing view of abrupt transitions in animal landscape perception

A common property of landscapes and metacommunities is the occurrence of abrupt shifts in connectivity along gradients of individual dispersal abilities. Animals with short‐range dispersal capability perceive fragmented landscapes, but organisms moving across critical thresholds perceive continuous landscapes. This qualitative shift in landscape perception may determine several attributes of local communities and the dynamics of whole metacommunities. Modularity describes the existence in some communities of relatively high numbers of mutual connections favoring the movement of neighboring individuals (even when each individual is able to reach any patch in the landscape). Local patch linkages and metacommunity connectivity along gradients of dispersal ability have been reported frequently. However, the intermediate level of structure captured by modularity has not been considered. We evaluated landscape connectivity and modularity along gradients of individual dispersal abilities. Random landscapes with different degrees of cell aggregation and occupancy were simulated; we also analyzed ten real ecosystems. An expected, a shift in landscape connectivity was always detected; modularity consistently decreased gradually along dispersal gradients in both simulated networks and empirical landscapes. Neutral metacommunities within simulated landscapes demonstrated that modularity and connectivity may reflect landscape traits in the shaping of metacommunity diversity. Average beta‐diversity was strongly associated with modularity, particularly with low migration rates, while connectivity trends tracked changes in beta‐diversity at intermediate to high migrations rates. Consequently, while some species are able to perceive abrupt transitions in the landscape, many others probably experience a gradual continuum in landscape perception, contrary to predictions from previous analyses. Furthermore, the gradual behavior of modularity indicates that it may represent an exceptional early‐warning tool that measures system distance to tipping points. Our study highlights the multiple perceptions that different species may have of a single landscape and shows, for the first time, a theoretical and empirical relationship between landscape modularity, and metacommunity diversity.     

Fumarole-Supported Islands of Biodiversity within a Hyperarid, High-Elevation Landscape on Socompa Volcano, Puna de Atacama, Andes

Fumarolic activity supports the growth of mat-like photoautotrophic communities near the summit (at 6,051 m) of Socompa Volcano in the arid core of the Andes mountains. These communities are isolated within a barren, high-elevation landscape where sparse vascular plants extend to only 4,600 m. Here, we combine biogeochemical and molecular-phylogenetic approaches to characterize the bacterial and eucaryotic assemblages associated with fumarolic and nonfumarolic grounds on Socompa. Small-subunit rRNA genes were PCR amplified, cloned, and sequenced from two fumarolic soil samples and two reference soil samples, including the volcanic debris that covers most of the mountain. The nonfumarolic, dry, volcanic soil was similar in nutrient status to the most extreme Antarctic Dry Valley or Atacama Desert soils, hosted relatively limited microbial communities dominated by Actinobacteria and Fungi, and contained no photoautotrophs. In contrast, modest fumarolic inputs were associated with elevated soil moisture and nutrient levels, the presence of chlorophyll a, and ¹³C-rich soil organic carbon. Moreover, this soil hosted diverse photoautotroph-dominated assemblages that contained novel lineages and exhibited structure and composition comparable to those of a wetland near the base of Socompa (3,661-m elevation). Fumarole-associated eucaryotes were particularly diverse, with an abundance of green algal lineages and a novel clade of microarthropods. Our data suggest that volcanic degassing of water and ¹³C-rich CO₂ sustains fumarole-associated primary producers, leading to a complex microbial ecosystem within this otherwise barren landscape. Finally, we found that human activities have likely impacted the fumarolic soils and that fumarole-supported photoautotrophic communities may be exceptionally sensitive to anthropogenic disturbance.