Trait-mediated effects on flowers: artificial spiders deceive pollinators and decrease plant fitness

Although predators can affect foraging behaviors of floral visitors, rarely is it known if these top-down effects of predators may cascade to plant fitness through trait-mediated interactions. In this study we manipulated artificial crab spiders on flowers of Rubus rosifolius to test the effects of predation risk on flower-visiting insects and strength of trait-mediated indirect effects to plant fitness. In addition, we tested which predator traits (e.g., forelimbs, abdomen) are recognized and avoided by pollinators. Total visitation rate was higher for control flowers than for flowers with an artificial crab spider. In addition, flowers with a sphere (simulating a spider abdomen) were more frequently visited than those with forelimbs or the entire spider model. Furthermore, the presence of artificial spiders decreased individual seed set by 42% and fruit biomass by 50%. Our findings indicate that pollinators, mostly bees, recognize and avoid flowers with predation risk; forelimbs seem to be the predator trait recognized and avoided by hymenopterans. Additionally, predator avoidance by pollinators resulted in pollen limitation, thereby affecting some components of plant fitness (fruit biomass and seed number). Because most pollinator species that recognized predation risk visited many other plant species, trait-mediated indirect effects of spiders cascading down to plant fitness may be a common phenomenon in the Atlantic rainforest ecosystem.     

Habitat patch shape, not corridors, determines herbivory and fruit production of an annual plant

Habitat corridors confer many conservation benefits by increasing movement of organisms between habitat patches, but the benefits for some species may exact costs for others. For example, corridors may increase the abundance of consumers in a habitat to the detriment of the species they consume. In this study we assessed the impact of corridors on insect herbivory of a native plant, Solanum americanum, in large-scale, experimentally fragmented landscapes. We quantified leaf herbivory and assessed fruit production as a proxy for plant fitness. We also conducted field surveys of grasshoppers (Orthoptera), a group of abundant, generalist herbivores that feed on S. americanum, and we used exclosure cages to explicitly link grasshopper herbivory to fruit production of individual S. americanum. The presence of corridors did not increase herbivory or decrease plant fruit production. Likewise, corridors did not increase grasshopper abundance. Instead, patches in our landscapes with the least amount of edge habitat and the greatest amount of warmer "core" area had the highest levels of herbivory, the largest cost to plant fruit production as a result of herbivory, and the most grasshoppers. Thus habitat quality, governed by patch shape, can be more important than connectivity for determining levels of herbivory and the impact of herbivory on plant fitness in fragmented landscapes.     

Direct and indirect effects of ants on a forest-floor food web

Interactions among predators that prey on each other and are potential competitors for shared prey (intraguild [IG] predators) are widespread in terrestrial ecosystems and have the potential to strongly influence the dynamics of terrestrial food webs. Ants and spiders are abundant and ubiquitous terrestrial IG predators, yet the strength and consequences of interactions between them are largely unknown. In the leaf-litter food web of a deciduous forest in Kentucky (USA), we tested the direct and indirect effects of ants on spiders and a category of shared prey (Collembola) by experimentally subsidizing ants in open plots in two field experiments. In the first experiment, ant activity was increased, and the density of ants in the litter was doubled, by placing carbohydrate and protein baits in the center of each plot. Gnaphosa spiders were almost twice as abundant and Schizocosa spiders were half as abundant in baited plots relative to controls. There were more tomocerid Collembola in baited plots, suggesting possible indirect effects on Collembola caused by ant-spider interactions. The second experiment, in which screening of two mesh sizes selectively excluded large and small worker ants from a sugar bait, revealed that the large ants, primarily Camponotus, could alone induce similar effects on spiders. Gnaphosa biomass density was almost twice as high in the plots where large ants were more active, whereas Schizocosa biomass density was reduced by half in these plots. Although tomocerid densities did not differ between treatments, tomocerid numbers were negatively correlated with the activity of Formica, another large ant species. Path analysis failed to support the hypothesis that the ant Camponotus indirectly affected tomocerid Collembola through effects on densities of spiders. However, path analysis also revealed other indirect effects of Camponotus affecting tomocerids. These results illustrate the complexity of interactions between and within two major IG predator groups with disparate predatory behaviors, complexities that will have consequences for functioning of the forest-floor food web.     

Negative effects of vertebrate herbivores on invertebrates in a coastal dune community

Although competition has been a major focus in ecology for the past century, most empirical and theoretical studies in this area have emphasized interactions between closely related species. However, there is growing evidence that negative interactions among distantly related taxa also occur and may be far more important than previously thought. In this study, we took advantage of an 11-year-old replicated vertebrate-exclosure experiment in a coastal dune community in northern California, USA, to examine the effects of the two most common vertebrate herbivores (jackrabbits and black-tailed deer) on the abundance of the three most visible invertebrate herbivores (two snail, a moth, and a grasshopper species). Our results indicate that four of the six possible pairwise interactions were significantly negative for the invertebrates. Jackrabbits reduced the abundances of snails by 44-75%, tiger moth caterpillars by 36%, and grasshoppers by 62%. Deer reduced the abundances of snails by 32%, increased the abundances of caterpillars by 31%, and had no measurable effect on grasshopper abundance. Our data also revealed that jackrabbits significantly decreased the volume of forbs and common shrubs and the flowering by grasses in our study plots. We were unable to detect an effect of deer on these measures of vegetation. These results suggest that by changing vegetation, jackrabbits may reduce invertebrate populations that are limited by food, protective structures, or microclimate provided by plants. Of these three mechanisms, only shade was strongly supported as limiting snail numbers in smaller-scale manipulations. In most systems, as in this one, the number of pairs of distantly related herbivores far exceeds the number of pairs of congeners. Since interactions among distantly related herbivores may be common in many cases, these interactions are likely to be important and should receive far more attention from ecologists.     

Climate heterogeneity modulates impact of warming on tropical insects

Evolutionary history and physiology mediate species responses to climate change. Tropical species that do not naturally experience high temperature variability have a narrow thermal tolerance compared to similar taxa at temperate latitudes and could therefore be most vulnerable to warming. However, the thermal adaptation of a species may also be influenced by spatial temperature variations over its geographical range. Spatial climate gradients, especially from topography, may also broaden thermal tolerance and therefore act to buffer warming impacts. Here we show that for low-seasonality environments, high spatial heterogeneity in temperature correlates significantly with greater warming tolerance in insects globally. Based on this relationship, we find that climate change projections of direct physiological impacts on insect fitness highlight the vulnerability of tropical lowland areas to future warming. Thus, in addition to seasonality, spatial heterogeneity may play a critical role in thermal adaptation and climate change impacts particularly in the tropics.     

Effects of Widespread Drought‐Induced Aspen Mortality on Understory Plants

Forest die‐off around the world is expected to increase in coming decades as temperature increases due to climate change. Forest die‐off will likely affect understory plant communities, which have substantial influence on regional biological diversity, ecosystem function, and land–atmosphere interactions, but how die‐off alters these plant communities is largely unknown. We examined changes in understory plant communities following a widespread, drought‐induced die‐off of trembling aspen (Populus tremuloides) in the western United States. We assessed shrub and herbaceous cover and volume in quadrats in 55 plots located across a wide range of levels of aspen mortality. We measured species richness and composition of herbaceous plant communities by recording species presence and absence in 12 sets of paired (1 healthy, 1 dying) aspen plots. Although understory composition in healthy and dying stands was heterogeneous across the landscape, shrub abundance, cover, and volume were higher and abundance of herbaceous species, cover, and volume were lower in dying aspen stands. Shrub cover and volume increased from 2009 to 2011 in dying stands, which suggests that shrub growth and expansion is ongoing. Species richness of herbs declined by 23% in dying stands. Composition of herbs differed significantly between dying and healthy stands. Richness of non‐native species did not differ between stand types. The understory community in dying aspen stands was not similar to other shrub‐dominated plant communities in the region and may constitute a novel community. Our results suggest that changes in understory plant communities as forests die off could be a significant indirect effect of climate change on biological diversity and forest communities. Efectos de la Mortalidad Extensiva de Álamos Inducida por Sequía sobre Plantas del Sotobosque     

Biotic habitat complexity controls species diversity and nutrient effects on net biomass production

Canopy-forming plants and algae commonly contribute to spatial variation in habitat complexity for associated organisms and thereby create a biotic patchiness of communities. In this study, we tested for interaction effects between biotic habitat complexity and resource availability on net biomass production and species diversity of understory macroalgae by factorial field manipulations of light, nutrients, and algal canopy cover in a subtidal rocky-shore community. Presence of algal canopy cover and/or artificial shadings limited net biomass production and facilitated species diversity. Artificial shadings reduced light to levels similar to those under canopy cover, and net biomass production was significantly and positively correlated to light availability. Considering the comparable and dependent experimental effects from shadings and canopy cover, the results strongly suggest that canopy cover controlled net biomass production and species diversity by limiting light and thereby limiting resource availability for community production. Canopy cover also controlled experimental nutrient effects by preventing a significant increase in net biomass production from nutrient enrichment recorded in ambient light (no shading). Changes in species diversity were mediated by changes in species dominance patterns and species evenness, where canopy cover and shadings facilitated slow-growing crust-forming species and suppressed spatial dominance by Fucus vesiculosus, which was the main contributor to net production of algal biomass. The demonstrated impacts of biotic habitat complexity on biomass production and local diversity contribute significantly to understanding the importance of functionally important species and biodiversity for ecosystem processes. In particular, this study demonstrates how loss of a dominant species and decreased habitat complexity change the response of the remaining assembly to resource loading. This is of potential significance for marine conservation since resource loading often promotes low habitat complexity and canopy species are among the first groups lost in degraded aquatic systems.     

草原不同沙化地段蝗虫与植物群落多样性的变化及相互关系的数值分析

通过对植物和蝗虫种群生物量的调查，研究了宁夏盐池县干草原沙化过程中植物和蝗虫群落结构、多样性与丰富度的变化，以及其相互关系．结果表明，植物和蝗虫群落随着草地的沙化程度表现出序列性演替过程，蝗虫群落多样性直接受植物群密结构变化的影响，但二者的变化趋势并不完全平行．草地沙化对蝗虫群落多样性和丰富度是一种灾变性影响：沙化导致植被结构与土壤硬度的变化，造成蝗虫产卵地条件的改变．     

Trade-offs in foraging success and predation risk with spatial position in colonial spiders

Summary Colonial web-building spiders respond to trade-offs between selective forces relative to spatial position within colonies and thus provide support for the selfish herd theory. The size distribution of spiders within colonies of Metepeira incrassata, a colonial orb-weaver (Araneae: Araneidae) from tropical Mexico is nonrandom; larger (mature) spiders and females guarding eggsacs are more prevalent in the center, whereas more small (immature) spiders are found on the periphery. Experimental field studies with spiders of selected size classes show that larger spiders actively and aggressively seek protected positions in the center of the colony webbing, even though prey availability and capture rates are significantly higher on the periphery. Attacks by predatory wasps, other spiders, and hummingbirds are more frequent on the periphery than in the core of the colony. Reproductive females on the periphery are at greater risk because they are captured more often than smaller spiders, and if their egg sacs consequently remain unguarded, chances of cocoon parasitism are increased. As a result, spiders in the core of the colony have greater reproductive success, producing more egg sacs with greater hatching frequency. Colonial spiders thus appear to be making a trade-off between foraging and protection from predation and show a spatial organization predicted by the selfish herd theory. The influence of such trade-offs on individual fitness and the structure of colonies is discussed. Offprint requests to: G.W. Uetz     

Indirect effects of prey swamping: differential seed predation during a bamboo masting event

Resource pulses often involve extraordinary increases in prey availability that "swamp" consumers and reverberate through indirect interactions affecting other community members. We developed a model that predicts predator-mediated indirect effects induced by an epidemic prey on co-occurring prey types differing in relative profitability/preference and validated our model by examining current-season and delayed effects of a bamboo mass seeding event on seed survival of canopy tree species in mixed Patagonian forests. The model shows that predator foraging behavior, prey profitability, and the scale of prey swamping influence the character and strength of short-term indirect effects on various alternative prey. When in large prey-swamped patches, nonselective predators decrease predation on all prey types. Selective predators, instead, only benefit prey of similar quality to the swamping species, while very low or high preference prey remain unaffected. Negative indirect effects (apparent competition) may override such positive effects (apparent mutualism), especially for highly preferred prey, when prey-swamped patches are small enough to allow predator aggregation and/or predators show a reproductive numerical response to elevated food supply. Seed predation patterns during bamboo (Chusquea culeou) masting were consistent with predicted short-term indirect effects mediated by a selective predator foraging in large prey-swamped patches. Bamboo seeds and similarly-sized Austrocedrus chilensis (ciprés) and Nothofagus obliqua (roble) seeds suffered lower predation in bamboo flowered than nonflowered patches. Predation rates on the small-seeded Nothofagus dombeyi (coihue) and the large-seeded Nothofagus alpina (rauli) were independent of bamboo flowering. Indirect positive effects were transient; three months after bamboo seeding, granivores preyed heavily upon all seed types, irrespective of patch flowering condition. Moreover, one year after bamboo seeding, predation rates on the most preferred seed (rauli) was higher in flowered than in nonflowered patches. Despite rapid predator numerical responses, short-term positive effects can still influence community recruitment dynamics because surviving seeds may find refuge beneath the litter produced by bamboo dieback. Together, our theoretical analysis and experiments indicate that indirect effects experienced by alternative prey during and after prey-swamping episodes need not be universal but can change across a prey quality spectrum, and they critically depend on predator-foraging rules and the spatial scale of swamping.     

Social spiders catch larger prey: a study of Anelosimus eximius (Araneae: Theridiidae)

Summary During a 1-year-study in tropical Panama, prey of the social theridiid Anelosimus eximius was analysed at two locations and compared with the potential prey spectrum according to sweepnet catches, pitfall traps and bowl traps. Compared with other web-building spiders, A. eximius catch an unusually high number of large insects: about 90% are flying ants, beetles,lepidopterans hemipterans, cockroaches and grasshoppers. This is the result of a communal strategy to overwhelm prey. Webs are maintained commonly, and several spiders attack an entangled insect simultaneously. More spiders participate on insects that are larger and struggle more. The ability to catch large prey insects is discussed as a major driving factor for sociality in spiders.     

Land Forms and Winter Habitat Refugia in the Conservation of Montane Grasshoppers in Southern Africa

Abstract: The importance of topography in evaluation of insect conservation is discussed with reference to grasshoppers in the Natal Drakensberg Mountains, South Africa For these insects, the hilltops act as thermal refugia from winter cold air drainage. Additionally, the increased insolation on the eastern and northern sides of the hilltops, compared with the western and southern sides, attracts the grasshoppers. Crevices in the hill summits provide further microrefugia Burning reduces grasshopper numbers by half and although many species are adapted to burn conditions, the reduced grass cover accentuates the thermal influences. Other animals, and plants, also selectively inhabit these hilltops, making them highly significant conservation refugia in a matrix of thermally inhospitable land. Evaluations for biological conservation should take note of the strong influence that land forms may have on plant and animal local distributions.     

温带落叶林的植物物候特征及其对气候变化的响应

植被物候是气候变化对生物圈产生长期或短期影响的重要指示因子。气候变化已经明显改变了许多物种的营养生长和繁殖物候,尤其是在温带地区。研究温带森林物候变化及其对全球变暖的响应,对认识森林物种共存,协同进化以及森林保护和经营等有重要意义。通过概述温带森林下物候研究的进展发现,光照和积温是影响木本植物展叶及繁殖物候的关键因素,林下层树木通过更早展叶,以尽量减少生长季林冠层遮阴对下层树木生长的影响,更早时期开花的树木具有从顶部向四周次第开花的时空格局,林冠层树种开花具有较好的同步性。而草本植物的物候通常受融雪时间和冠层动态的影响更大,并且,温带森林下不同生活史对策的草本植物的物候特征对气候变化的响应也不尽相同,存在明显的季节动态。繁殖物候、光照的季节变化、光合特征、授粉成功之间的联系决定了林下不同繁殖特性的草本植物的繁殖成功率。量化的、多指标、多对象的定位监测是精准物候研究的基础,物候变化的机理和建立可预测的物候模型将是未来研究的重点。     

Parasite facilitates plant species coexistence in a coastal wetland

Outbreaks of infectious agents in natural ecosystems are on the rise. Understanding host-pathogen interactions and their impact on community composition may be central to the conservation of biological diversity. Infectious agents can convey both exploitive and facilitative effects that regulate host populations and community structure. Parasitic angiosperms are highly conspicuous in many plant communities, and they provide a tractable model for understanding parasite effects in multispecies communities. I examined host identity and variation in host infectivity of a holoparasitic vine (Cuscuta salina) within a California salt marsh. In a two-year parasite removal experiment, I measured the effect of C. salina on its most frequent host, a rare hemiparasite, and the plant community. C. salina clearly suppressed the dominant host, but rare plant fitness and plant species diversity were enhanced through indirect effects. Priority effects played a role in the strength of the outcome due to the timing of life history characteristics. The differential influence of parasites on the fecundity of multiple hosts can change population dynamics, benefit rare species, and alter community structure. The continuum of negative to positive consequences of parasitic interactions deserves more attention if we are to understand community dynamics and successfully restore tidal wetlands.     

Switching to Plan B: changes in the escape tactics of two grasshopper species (Acrididae: Orthoptera) in response to repeated predatory approaches

Most studies examining escape behaviour have considered single approaches and single fleeing responses; few have considered how organisms’ response is influenced by persistent pursuit. We explored fleeing behaviour of two grasshopper species to test whether they modified escape behaviour when approached repeatedly. Schistocerca alutacea did not increase flight initiation distance (FID) upon repeated approach but fled farther. Psinidia fenestralis increased its FID on the second approach but decreased its flight distance over successive escapes. Both species showed a bimodal pattern of flight direction, either flying directly away or flying perpendicular to the direction of the observer’s approach. Neither species showed a significant pattern of flight direction or change in flight direction with successive escapes. Most (88 %) P. fenestralis initially landed on sand, but after repeated approaches an increasing proportion landed in grass and hid. Both species therefore changed escape behaviour with persistent pursuit but used different tactics, suiting their flight ability or camouflage, and optimised habitat use. Three grasshopper species have now been examined for responses to repeated approach by predators and all show different tactics supporting escape decision theory. Our results emphasise the variety of escape responses across species and how the dynamic nature of escape responses vary according to an animal’s situation. Rather than single optimum escape options, each grasshopper species shows a range of responses, which vary with risk from persistent predators. Although grasshoppers provide an excellent model, it would be profitable to examine responses of a range of species according to levels of predation risk.     

Nymphal density, behavioral development, and life history in a field cricket

Population density regulates the strength of intraspecific competition and may thereby be reflected in life-history variables, such as development time, growth rate, or investment in immune defense. However, population density may also affect the fitness payoffs of different behaviors and consequently shape the development of personality. Here we studied if population density during nymphal development (one, four, or ten individuals raised together) affects the level of boldness, measured as the latency time to recover from freezing and emerge from a shelter, aggressiveness towards conspecifics or their correlation at the adult stage in the field crickets, Gryllus integer. In addition, we tested if individuals invest more resources in immune function or speed up their development in response to a high conspecifics density during ontogeny. Nymphal density did not affect adult boldness or aggressiveness towards conspecific males per se, but these variables showed a negative association, i.e., indicated an unconventional behavioral syndrome in the highest density treatment. Supporting the effectiveness of density treatments in inducing plastic responses, individuals reached maturity sooner and invested more resources in immune function in the highest nymphal density group compared to groups consisting of one or four individuals. Our results suggest that population density may play an important role in shaping both the realized life history and development of behavioral syndromes.     

Direct and indirect effects of climate and habitat factors on butterfly diversity

Many factors, including climate, resource availability, and habitat diversity, have been proposed as determinants of global diversity, but the links among them have rarely been studied. Using structural equation modeling (SEM), we investigated direct and indirect effects of climate variables, host-plant richness, and habitat diversity on butterfly species richness across Britain, at 20-km grid resolution. These factors were all important determinants of butterfly diversity, but their relative contributions differed between habitat generalists and specialists, and whether the effects were direct or indirect. Climate variables had strong effects on habitat generalists, whereas host-plant richness and habitat diversity contributed relatively more for habitat specialists. Considering total effects (direct and indirect together), climate variables had the strongest link to butterfly species richness for all groups of species. The results suggest that different mechanistic hypotheses to explain species richness may be more appropriate for habitat generalists and specialists, with generalists hypothesized to show direct physiological limitations and specialists additionally being constrained by trophic interactions (climate affecting host-plant richness).     

Stereochemistry affects sesquiterpene lactone bioactivity against an herbivorous grasshopper

Sesquiterpene lactones are defensive compounds which protect plants against a variety of herbivores and other natural enemies. Sesquiterpene lactones from higher plants can be divided into two groups based on the stereochemistry of their lactone ring junction, either cis-fused or trans-fused. It is unclear whether and how this variation affects potentially important ecological interactions. To investigate whether stereochemical variation in sesquiterpene lactone ring junctions can influence resistance to herbivorous insects, we performed controlled feeding trials with two pairs of diastereomeric sesquiterpene lactones and examined the deterrent effect of each compound on feeding by the polyphagous grasshopper Schistocerca americana (Drury). Sesquiterpene lactone stereochemistry and concentration significantly influenced feeding behavior with grasshoppers consuming less of the trans-fused compounds than the cis-fused compounds. To our knowledge, this is the first demonstration that sesquiterpene lactone ring junction stereochemistry influences the feeding behavior of herbivores. Because this stereochemical trait polymorphism is widely distributed in nature, it could have substantial consequences for the ecology and evolution of large groups of plants, particularly the Asteraceae.     

Comparison of spatial overlap between the polychaetes Nereis virens and Nephtys caeca in two intertidal estuarine environments

An intensive study of spatial overlap between the polychaetes Nereis virens (Sars) and Nephtys caeca (Fabricius) was conducted in 1992 on two tidal sand flats, which differ by their exposure to dominant winds and residual currents in the Lower St. Lawrence Estuary (Québec, Canada). Results showed that spatial overlap (Lloyd's index and spatial distribution) was higher among adults of both species at the lower tidal elevation of l'Anse-à-l'Orignal (north-east oriented). Results also suggest weaker interspecific interactions among juveniles of both species because of limited spatial overlap. In Baie-du-Ha!Ha! (south-west oriented), spatial overlap was greater than that observed in l'Anse-à-l'Orignal and appeared important in adults as well as in juveniles. Levins' directional measure of competition indicated an asymmetric spatial overlap between N. virens and N. caeca in Baie-du-Ha!Ha! and a symmetric overlap in l'Anse-à-l'Orignal. The intraspecific encounter values, estimated from Lloyd's mathematical expression, were significantly higher than values of interspecific interactions only in l'Anse-à-l'Orignal. Moreover, no vertical stratification was found inside the sediment, with no effect of the densities and individual body weights of the polychaetes. Juveniles of both species mainly inhabited the organic-rich upper portion of the sediments (0 to 12 cm), while adults colonised greater depths (>25 cm) where organic matter content was lower. A complementary field experiment was conducted in 1993 to investigate interspecific interactions (predation and competition) existing between N. virens and N. caeca. Results from this experiment depend on which species was first-introduced and showed an important variation in mortality rates between allopatric and sympatric conditions. The influence of competition and predation on the structure of these populations is also discussed in relation to their spatial overlap.