Interaction between spiders and ticks-ancient arthropod predatory behavior?

Ticks are ectoparasite vectors of pathogens affecting human and animal health worldwide. Rational integration of different control interventions including plant-derived repellents and acaricides, management of natural predators, and vaccines is required for innovative approaches to reduce the risks associated with ticks and tick-borne diseases. How tick populations are naturally controlled is always a question. Tick interactions with other arthropods including predators evolved from ancient times. In this study, Cretaceous (ca. 100 Mya) Burmese amber inclusions were identified as probably related to Compluriscutula vetulum (Acari: Ixodida: Ixodidae) tick larvae and spider silk. As illustrated in this study, ancient interactions between ticks and spiders may support arthropod predatory behavior as a natural control intervention. Rational integrative management of different tick control interventions including natural predators under a One Health perspective will contribute to effectively and sustainably reducing the risks associated with ticks and tick-borne diseases.

Infrequent Long-Range Dispersal and Evolution of a Top Terrestrial Arthropod Predator in the Sub-Antarctic.

AbstractThe sub-Antarctic terrestrial ecosystems survive on isolated oceanic islands in the path of circumpolar currents and winds that have raged for more than 30 million years and are shaped by climatic cycles that surpass the tolerance limits of many species. Surprisingly little is known about how these ecosystems assembled their native terrestrial fauna and how such processes have changed over time. Here, we demonstrate the patterns and timing of colonization and speciation in the largest and dominant arthropod predators in the eastern sub-Antarctic: spiders of the genus Myro. Our results indicate that this lineage originated from Australia before the Plio-Pleistocenic glacial cycles and underwent an adaptive radiation on the Crozet archipelago, from where one native species colonized multiple remote archipelagos via the Antarctic circumpolar current across thousands of kilometers. The results indicate limited natural connectivity between terrestrial macroinvertebrate faunas in the eastern sub-Antarctic and partial survival of repeated glaciations in the Plio-Pleistocene. Furthermore, our findings highlight that by integrating arthropod taxa from multiple continents, the climatically more stable volcanic Crozet archipelago played a critical role in the evolution and distribution of arthropod life in the sub-Antarctic.

Broken Ring Speciation in California Mygalomorph Spiders (Nemesiidae, Calisoga).

AbstractIdealized ring species, with approximately continuous gene flow around a geographic barrier but singular reproductive isolation at a ring terminus, are rare in nature. A broken ring species model preserves the geographic setting and fundamental features of an idealized model but accommodates varying degrees of gene flow restriction over complex landscapes through evolutionary time. Here we examine broken ring species dynamics in Calisoga spiders, which, like the classic ring species Ensatina salamanders, are distributed around the Central Valley of California. Using nuclear and mitogenomic data, we test key predictions of common ancestry, ringlike biogeography, biogeographic timing, population connectivity, and terminal overlap. We show that a ring complex of populations shares a single common ancestor, and from an ancestral area in the Sierra Nevada mountains, two distributional and phylogenomic arms encircle the Central Valley. Isolation by distance occurs along these distributional arms, although gene flow restriction is also evident. Where divergent lineages meet in the South Coast Ranges, we find rare lineage sympatry, without evidence for nuclear gene flow and with clear evidence for morphological and ecological divergence. We discuss general insights provided by broken ring species and how such a model could be explored and extended in other systems and future studies.

Does female control and male mating system predict courtship investment and mating outcomes? A comparative study in five widow spider species (genus Latrodectus) tested under similar laboratory conditions.

BACKGROUND: Male courtship investment may evolve in response to the male's expectation of future mating opportunities or the degree of female control during mating interactions. We used a comparative approach to test this hypotheses by assessing the courtship and mating behaviors of five widow spider species (genus Latrodectus) under common laboratory conditions. We predicted male investment in courtship would be higher in species where males mate only once because of high cannibalism rates (monogyny, L. geometricus, L. hasselti, L. mirabilis), compared to species with rare cannibalism (L. mactans, L. hesperus) in which males should reserve energy for future mating opportunities. Increased male investment, measured as courtship duration, might also evolve with increased female control over mating outcomes if females prefer longer courtships. We tested this by assessing the frequency of copulations, timing of sexual cannibalism, and the degree of female-biased size dimorphism, which is expected to be negatively correlated with the energetic cost of rebuffing male mating attempts. RESULTS: Copulation frequency was consistently lower in species with extreme female-skewed size dimorphism, and where sexual cannibalism was more prevalent, suggesting the importance of female control for mating outcomes. We confirmed significant interspecific variation in average courtship duration, but contrary to predictions, it was not predicted by male mating system, and there was no consistent link between courtship duration and sexual size dimorphism. CONCLUSION: We show that the degree of sexual dimorphism is not only correlated with sexual cannibalism, but also with mating success since restriction of male copulation frequency by female Latrodectus affects paternity. However, predictions about male mating system or female control affecting courtship duration were not supported. We propose that the form of female control over mating and cannibalism, and male responses, might be more informative for understanding the evolution of courtship duration. For example, male tactics to avoid female aggression may drive lower courtship duration in species like L. mirabilis. Nonetheless, our results differ from inferences based on published studies of each species in isolation, illuminating the need for standardized data collection for behavioural comparative studies.

Changes in the material properties of spider glue droplet proteins accompanied shifts in prey capture biomechanics as cobweb spiders diverged from their orb weaving ancestors.

Although descended from orb weavers, spiders in the family Theridiidae spin cobwebs whose sticky prey capture gumfoot lines extend from a silk tangle to a surface below. When a crawling insect contacts glue droplets at the bottom of a gumfoot line, the line's weak pyriform anchor releases, causing the taut line to contract, pulling the insect from the surface and making its struggles to escape ineffective. To determine if this change in prey capture biomechanics was accompanied by a change in the material properties of theridiid glue, we characterized the elastic modulus and toughness of the glue droplet proteins of four theridiid species at 20-90 % relative humidity and compared their properties with those of 13 orb weaving species in the families Tetragnathidae and Araneidae. Compared to orb weavers, theridiid glue proteins had low extensions per protein volume and low elastic modulus and toughness values. These differences are likely explained by the loss of tension on a gumfoot line when its anchor fails, which may prioritize glue droplet adhesion rather than extension. Similarities in theridiid glue droplet properties did not reflect these species' evolutionary relationships. Instead, they appear associated with differences in web architecture. Two species that had stiffer gumfoot support lines and longer and more closely spaced gumfoot lines also had stiffer glue proteins. These lines may store more energy, and, when their anchors release, require stiffer glue to resist the more forceful upward thrust of a prey. STATEMENT OF SIGNIFICANCE: When a crawling insect contacts glue droplets on a theridiid cobweb's gumfoot line, this taut line's anchor fails and the insect is hoisted upward, rendering its struggles to escape ineffective. This strategy contrasts with that of orb weaving ancestors, which rely on more closely spaced prey capture threads to intercept and retain flying insects. A comparison of the elastic modulus and toughness of gumfoot and orb web glue proteins shows that this change in prey capture biomechanics is associated with reductions in the stiffness and toughness of cobweb glue. Unlike orb web capture threads, whose droplets extend in a coordinated fashion to sum adhesive forces, gumfoot lines become untethered, which prioritizes glue droplet adhesive contact over glue droplet extension.

Taxon-dependent diversity response along a temperate elevation gradient covered by grassland.

Elevational gradients constitute excellent systems for understanding the mechanisms that generate and maintain global biodiversity patterns. Climatic gradients associated with elevation show strong influence on species distribution in mountains. The study of mountains covered by the same habitat type is an ideal scenario to compare alternatives to the energy hypotheses. Our aim was to investigate how changes in climatic conditions along the elevational gradient drive α- and ß-diversity of four taxa in a mountain system located within a grassland biome. We sampled ants, spiders, birds and plants, and measured climatic variables at six elevational bands (with 10 sampling sites each) established between 470 and 1,000 masl on a mountain from the Ventania Mountain System, Argentina. Species richness per site and ß-diversity (turnover and nestedness) between the lowest band and upper sites were estimated. For most taxa, species richness declined at high elevations and energy, through temperature, was the major driver of species richness for ants, plants and birds, prevailing over productivity and water availability. The major ß-diversity component was turnover for plants, spiders and birds, and nestedness for ants. The unique environmental conditions of the upper bands could favour the occurrence of specialist and endemic species.

Warming alters cascading effects of a dominant arthropod predator on fungal community composition in the Arctic.

Rapid climate change in the Arctic is altering microbial structure and function, with important consequences for the global ecosystem. Emerging evidence suggests organisms in higher trophic levels may also influence microbial communities, but whether warming alters these effects is unclear. Wolf spiders are dominant Arctic predators whose densities are expected to increase with warming. These predators have temperature-dependent effects on decomposition via their consumption of fungal-feeding detritivores, suggesting they may indirectly affect the microbial structure as well. To address this, we used a fully factorial mesocosm experiment to test the effects of wolf spider density and warming on litter microbial structure in Arctic tundra. We deployed replicate litter bags at the surface and belowground in the organic soil profile and analyzed the litter for bacterial and fungal community structure, mass loss, and nutrient characteristics after 2 and 14 months. We found there were significant interactive effects of wolf spider density and warming on fungal but not bacterial communities. Specifically, higher wolf spider densities caused greater fungal diversity under ambient temperature but lower fungal diversity under warming at the soil surface. We also observed interactive treatment effects on fungal composition belowground. Wolf spider density influenced surface bacterial composition, but the effects did not change with warming. These findings suggest a widespread predator can have indirect, cascading effects on litter microbes and that effects on fungi specifically shift under future expected levels of warming. Overall, our study highlights that trophic interactions may play important, albeit overlooked, roles in driving microbial responses to warming in Arctic terrestrial ecosystems. IMPORTANCE: The Arctic contains nearly half of the global pool of soil organic carbon and is one of the fastest warming regions on the planet. Accelerated decomposition of soil organic carbon due to warming could cause positive feedbacks to climate change through increased greenhouse gas emissions; thus, changes in ecological dynamics in this region are of global relevance. Microbial structure is an important driver of decomposition and is affected by both abiotic and biotic conditions. Yet how activities of soil-dwelling organisms in higher trophic levels influence microbial structure and function is unclear. In this study, we demonstrate that predicted changes in abundances of a dominant predator and warming interactively affect the structure of litter-dwelling fungal communities in the Arctic. These findings suggest predators may have widespread, indirect cascading effects on microbial communities, which could influence ecosystem responses to future climate change.

Impact of pre-copulatory sexual cannibalism on genetic diversity and efficacy of selection.

Factors that increase reproductive variance among individuals act to reduce effective population size (Ne), which accelerates the loss of genetic diversity and decreases the efficacy of purifying selection. These factors include sexual cannibalism, offspring investment and mating system. Pre-copulatory sexual cannibalism, where the female consumes the male prior to mating, exacerbates this effect. We performed comparative transcriptomics in two spider species, the cannibalistic Trechaleoides biocellata and the non-cannibalistic T. keyserlingi, to generate genomic evidence to support these predictions. First, we estimated heterozygosity and found that genetic diversity is relatively lower in the cannibalistic species. Second, we calculated dN/dS ratios as a measure of purifying selection; a higher dN/dS ratio indicated relaxed purifying selection in the cannibalistic species. These results are consistent with the hypothesis that sexual cannibalism impacts operational sex ratio and demographic processes, which interact with evolutionary forces to shape the genetic structure of populations. However, other factors such as the mating system and life-history traits contribute to shaping Ne. Comparative analyses across multiple contrasting species pairs would be required to disentangle these effects. Our study highlights that extreme behaviours such as pre-copulatory cannibalism may have profound eco-evolutionary effects.

Heuristic test reveals little effect of learning and maturation on early prey capture experiences in a web-building spider.

Behaviors can vary throughout an animal's life and this variation can often be explained by changes associated with learning and/or maturing. Currently, there is little consensus regarding how these processes interact to affect behaviors. Here we proposed a heuristic approach to disentangle the effects of learning and maturation on behavior and applied it to the predatory behaviors of Physocyclus globosus spiderlings. We varied the degree of prey difficulty and familiarity spiderlings received along the first instar and across the molt to the second instar and quantified the time spiderlings spent wrapping prey, as a proxy for prey capture efficiency. We found no overall evidence for learning or maturation. Changes in efficiency were mainly due to the switch from difficult to easy prey, or vice versa. However, there was one treatment where spiderlings improved in efficiency before and after the molt, without a switch in prey type. This provides some indication that difficult prey may offer more opportunity for learning or maturation to impact behavior. Although we found little effect of learning or maturation on prey capture efficiency, we suggest that our heuristic approach is effective and could be useful in investigating these processes in other behaviors and other animals.

All-you-can-eat buffet: A spider-specialized bat species (Myotis emarginatus) turns into a pest fly eater around cattle.

Determining the dietary spectrum of European insectivorous bats over time is the cornerstone of their conservation, as it will aid our understanding of foraging behavior plasticity in response to plummeting insect populations. Despite the global decline in insects, a restricted number of arthropod pest species thrive. Yet past research has overlooked the potential of European bats to suppress pests harmful to woodlands or livestock, in spite of their economic relevance. Here we investigated the diet composition, its breeding season variations and pest consumption of an insectivorous bat species (Myotis emarginatus), at the northern edge of its range (Wallonia, Belgium). We also explored the prey ecology to gain insight into the hunting strategies and foraging habitats of this bat species. We used DNA metabarcoding to amplify two COI markers within 195 bat droppings collected in June, July and August, thereby identifying 512 prey taxa predominated by Diptera, Araneae and Lepidoptera. Overall, in 97% of the samples we detected at least one of the 58 potential pest taxa, 41 of which targeting trees. The June samples were marked by a diet rich in orb-weaver spiders, in accordance with the archetypal diet of M. emarginatus bats. However, during the highly energy demanding July-August parturition and lactation period, roughly 55% of the dropping samples contained two cattle fly pests (Stomoxys calcitrans and Musca domestica). Moreover, among the 88 Diptera species preyed upon by M. emarginatus in July and August, these flies accounted for around 50% of the taxa occurrences. This plasticity-the switch from a spider-rich to a fly-rich diet-seems providential considering the dramatic ongoing drop in insect populations but this involves ensuring bat-friendly cattle farming. Our results revealed that bats widely consume pest entomofauna, thereby highlighting their potential role as allies of forest managers and farmers.

Note on hydrostatic skeletons: muscles operating within a pressurized environment.

Muscles and muscle fibers are volume-constant constructs that deform when contracted and develop internal pressures. However, muscles embedded in hydrostatic skeletons are also exposed to external pressures generated by their activity. For two examples, the pressure generation in spiders and in annelids, we used simplified biomechanical models to demonstrate that high intracellular pressures diminishing the resulting tensile stress of the muscle fibers are avoided in the hydrostatic skeleton. The findings are relevant for a better understanding of the design and functionality of biological hydrostatic skeletons.

Eye-specific detection and a multi-eye integration model of biological motion perception.

'Biological motion' refers to the distinctive kinematics observed in many living organisms, where visually perceivable points on the animal move at fixed distances from each other. Across the animal kingdom, many species have developed specialized visual circuitry to recognize such biological motion and to discriminate it from other patterns. Recently, this ability has been observed in the distributed visual system of jumping spiders. These eight-eyed animals use six eyes to perceive motion, while the remaining two (the principal anterior medial eyes) are shifted across the visual scene to further inspect detected objects. When presented with a biologically moving stimulus and a random one, jumping spiders turn to face the latter, clearly demonstrating the ability to discriminate between them. However, it remains unclear whether the principal eyes are necessary for this behavior, whether all secondary eyes can perform this discrimination, or whether a single eye-pair is specialized for this task. Here, we systematically tested the ability of jumping spiders to discriminate between biological and random visual stimuli by testing each eye-pair alone. Spiders were able to discriminate stimuli only when the anterior lateral eyes were unblocked, and performed at chance levels in other configurations. Interestingly, spiders showed a preference for biological motion over random stimuli - unlike in past work. We therefore propose a new model describing how specialization of the anterior lateral eyes for detecting biological motion contributes to multi-eye integration in this system. This integration generates more complex behavior through the combination of simple, single-eye responses. We posit that this in-built modularity may be a solution to the limited resources of these invertebrates' brains, constituting a novel approach to visual processing.

Hot and scared: how do heatwaves and predation risk impact resource acquisition and allocation?

Heatwaves are increasingly prevalent and can constrain investment into important life-history traits. In addition to heatwaves, animals regularly encounter threats from other organisms in their environments, such as predators. The combination of these two environmental factors introduces a decision-making conflict-heat exposure requires more food intake to fuel investment into fitness-related traits, but foraging in the presence of predators increases the threat of mortality. Thus, we used female variable field crickets (Gryllus lineaticeps) to investigate the effects of heatwaves in conjunction with predation risk (exposed food and water sources, and exposure to scent from black widow spiders, Latrodectus hesperus) on resource acquisition (food intake) and allocation (investment into ovarian and somatic tissues). A simulated heatwave increased food intake and the allocation of resources to reproductive investment. Crickets exposed to high predation risk reduced food intake, but they were able to maintain reproductive investment at an expense to investment into somatic tissue. Thus, heatwaves and predation risk deprioritized investment into self-maintenance, which may impair key physiological processes. This study is an important step towards understanding the ecology of fear in a warming world.

It is hot and cold here: the role of thermotolerance in the ability of spiders to colonize tree plantations in the southern Atlantic Forest.

Worldwide, with the decline of natural habitats, species with reduced niche breadth (specialists) are at greater risk of extinction as they cannot colonise or persist in disturbed habitat types. However, the role of thermal tolerance as a critical trait in understanding changes in species diversity in disturbed habitats, e.g., due to forest replacement by tree plantations, is still understudied. To examine the role of thermal tolerance on the responses of specialist and generalist species to habitat disturbances, we measured and compared local temperature throughout the year and thermotolerance traits [upper (CTmax) and lower (CTmin) thermal limits] of the most abundant species of spiders from different guilds inhabiting pine tree plantations and native Atlantic Forests in South America. Following the thermal adaptation hypothesis, we predicted that generalist species would show a wider thermal tolerance range (i.e., lower CTmin and higher CTmax) than forest specialist species. As expected, generalist species showed significantly higher CTmax and lower CTmin values than specialist species with wider thermal tolerance ranges than forest specialist species. These differences are more marked in orb weavers than in aerial hunter spiders. Our study supports the specialisation disturbance and thermal hypotheses. It highlights that habitat-specialist species are more vulnerable to environmental changes associated with vegetation structure and microclimatic conditions. Moreover, thermal tolerance is a key response trait to explain the Atlantic Forest spider's ability (or inability) to colonise and persist in human-productive land uses.

Distinct communities under the snow: describing characteristics of subnivium arthropod communities.

Arthropods are active during the winter in temperate regions. Many use the seasonal snowpack as a buffer against harsh ambient conditions and are active in a refugium known as the subnivium. While the use of the subnivium by arthropods is well established, far less is known about subnivium community composition, abundance, biomass, and diversity and how these characteristics compare with the community in the summer. Understanding subnivium communities is especially important given the observed and anticipated changes in snowpack depth and duration due to the changing climate. We compared subnivium arthropod communities with those active during the summer using pitfall trapping in northern New Hampshire. We found that compositions of ground-active arthropod communities in the subnivium differed from those in the summer. The subnivium arthropod community featured moderate levels of richness and other measures of diversity that tended to be lower than the summer community. More strikingly, the subnivium community was much lower in overall abundance and biomass. Interestingly, some arthropods were dominant in the subnivium but either rare or absent in summer collections. These putative "subnivium specialists" included the spider Cicurina brevis (Emerton 1890) (Araneae: Hahniidae) and 3 rove beetles (Coleoptera: Staphylinidae): Arpedium cribratum Fauvel, 1878, Lesteva pallipes LeConte, 1863, and Porrhodites inflatus (Hatch, 1957). This study provides a detailed account of the subnivium arthropod community, establishes baseline information on arthropod communities in temperate forests of northeastern North America, and explores the idea of subnivium specialist taxa that are highly active in winter and might be especially vulnerable to climate change.

Land-use change in the past 40 years explains shifts in arthropod community traits.

Understanding how anthropogenic activities induce changes in the functional traits of arthropod communities is critical to assessing their ecological consequences. However, we largely lack comprehensive assessments of the long-term impact of global-change drivers on the trait composition of arthropod communities across a large number of species and sites. This knowledge gap critically hampers our ability to predict human-driven impacts on communities and ecosystems. Here, we use a dataset of 1.73 million individuals from 877 species to study how four functionally important traits of carabid beetles and spiders (i.e. body size, duration of activity period, tolerance to drought, and dispersal capacity) have changed at the community level across ~40 years in different types of land use and as a consequence of land use changes (that is, urbanisation and loss of woody vegetation) at the landscape scale in Switzerland. The results show that the mean body size in carabid communities declined in all types of land use, with particularly stronger declines in croplands compared to forests. Furthermore, the length of the activity period and the tolerance to drought of spider communities decreased in most land use types. The average body size of carabid communities in landscapes with increased urbanisation in the last ~40 years tended to decrease. However, the length of the activity period, the tolerance to drought, and the dispersal capacity did not change significantly. Furthermore, urbanisation promoted increases in the average dispersal capacities of spider communities. Additionally, urbanisation favoured spider communities with larger body sizes and longer activity periods. The loss of woody areas at the landscape level was associated with trait shifts to carabid communities with larger body sizes, shorter activity periods, higher drought tolerances and strongly decreased dispersal capacities. Decreases in activity periods and dispersal capacities were also found in spider communities. Our study demonstrates that human-induced changes in land use alter key functional traits of carabid and spider communities in the long term. The detected trait shifts in arthropod communities likely have important consequences for their functional roles in ecosystems.

Foraging rates from metabarcoding: Predators have reduced functional responses in wild, diverse prey communities.

Functional responses describe foraging rates across prey densities and underlie many fundamental ecological processes. Most functional response knowledge comes from simplified lab experiments, but we do not know whether these experiments accurately represent foraging in nature. In addition, the difficulty of conducting multispecies functional response experiments means that it is unclear whether interaction strengths are weakened in the presence of multiple prey types. We developed a novel method to estimate wild predators' foraging rates from metabarcoding data and use this method to present functional responses for wild wolf spiders foraging on 27 prey families. These field functional responses were considerably reduced compared to lab functional responses. We further find that foraging is sometimes increased in the presence of other prey types, contrary to expectations. Our novel method for estimating field foraging rates will allow researchers to determine functional responses for wild predators and address long-standing questions about foraging in nature.

Brushing aside doubts: an evaluation of the beat-sheet brushing technique for detecting the Nearctic tree trunk sheetweaver (Araneae: Linyphiidae).

It is important to have reliable information on the presence/absence, population structure, and density of animals across their natural range. Detecting small organisms, however, such as the Nearctic tree trunk sheetweaver spider Drapetisca alteranda Chamberlin 1909 (Araneae: Linyphiidae), presents challenges due to its diminutive size and cryptic nature. We used a capture/recapture study to determine the detection and recapture probabilities of this spider using a standard beat sheet technique adopted for surveying tree trunks. Spiders were released on 3 different tree species that provided a range of microhabitats, including variable bark surface area and furrow depth/width. Microhabitat features played a small role in the timing of spider recapture (i.e., slower rate of recapture as furrowing increased). However, our results demonstrated 100% detection across replicate experiments and individual recapture probabilities exceeding 90% in most situations, with no significant differences in recapture observed among tree species and with respect to tree circumference. Furthermore, we show that most spiders could be recaptured within 2 sampling revolutions around the tree trunk, and there was no difference in the probability of collecting male and female spiders (although they differ markedly in size). Finally, we found no difference among brushers, supporting the idea that this method is replicable across collectors and studies. Collectively, we establish confidence in the ecological knowledge obtained with this technique and encourage its application with similar species and systems.

Expectancy and attention bias to spiders: Dissecting anticipation and allocation processes using ERPs.

The current registered report focused on the temporal dynamics of the relationship between expectancy and attention toward threat, to better understand the mechanisms underlying the prioritization of threat detection over expectancy. In the current event-related potentials experiment, a-priori expectancy was manipulated, and attention bias was measured, using a well-validated paradigm. A visual search array was presented, with one of two targets: spiders (threatening) or birds (neutral). A verbal cue stating the likelihood of encountering a target preceded the array, creating congruent and incongruent trials. Following cue presentation, preparatory processes were examined using the contingent negative variation (CNV) component. Following target presentation, two components were measured: early posterior negativity (EPN) and late positive potential (LPP), reflecting early and late stages of natural selective attention toward emotional stimuli, respectively. Behaviorally, spiders were found faster than birds, and congruency effects emerged for both targets. For the CNV, a non-significant trend of more negative amplitudes following spider cues emerged. As expected, EPN and LPP amplitudes were larger for spider targets compared to bird targets. Data-driven, exploratory, topographical analyses revealed different patterns of activation for bird cues compared to spider cues. Furthermore, 400-500 ms post-target, a congruency effect was revealed only for bird targets. Together, these results demonstrate that while expectancy for spider appearance is evident in differential neural preparation, the actual appearance of spider target overrides this expectancy effect and only in later stages of processing does the cueing effect come again into play.

Trophic cascades within and across ecosystems: The role of anti-predatory defences, predator type and detritus quality.

Species in one ecosystem can indirectly affect multiple biodiversity components and ecosystem functions of adjacent ecosystems. The magnitude of these cross-ecosystem effects depends on the attributes of the organisms involved in the interactions, including traits of the predator, prey and basal resource. However, it is unclear how predators with cross-ecosystem habitat interact with predators with single-ecosystem habitat to affect their shared ecosystem. Also, unknown is how such complex top-down effects may be mediated by the anti-predatory traits of prey and quality of the basal resource. We used the aquatic invertebrate food webs in tank bromeliads as a model system to investigate these questions. We manipulated the presence of a strictly aquatic predator (damselfly larvae) and a predator with both terrestrial and aquatic habitats (spider), and examined effects on survival of prey (detritivores grouped by anti-predator defence), detrital decomposition (of two plant species differing in litter quality), nitrogen flux and host plant growth. To evaluate the direct and indirect effects each predator type on multiple detritivore groups and ultimately on multiple ecosystem processes, we used piecewise structural equation models. For each response variable, we isolated the contribution of different detritivore groups to overall effects by comparing alternate model formulations. Alone, damselfly larvae and spiders each directly decreased survival of detritivores and caused multiple indirect negative effects on detritus decomposition, nutrient cycling and host plant growth. However, when predators co-occurred, the spider caused a negative non-consumptive effect on the damselfly larva, diminishing the net direct and indirect top-down effects on the aquatic detritivore community and ecosystem functioning. Both detritivore traits and detritus quality modulated the strength and mechanism of these trophic cascades. Predator interference was mediated by undefended or partially defended detritivores as detritivores with anti-predatory defences evaded consumption by damselfly larvae but not spiders. Predators and detritivores affected ecosystem decomposition and nutrient cycling only in the presence of high-quality detritus, as the low-quality detritus was consumed more by microbes than invertebrates. The complex responses of this system to predators from both recipient and adjacent ecosystems highlight the critical role of maintaining biodiversity components across multiple ecosystems.

As espécies em um ecossistema podem afetar indiretamente múltiplos componentes da biodiversidade e funções ecossistêmicas em ecossistemas adjacentes. A magnitude destes efeitos entre ecossistemas depende dos atributos dos organismos envolvidos nas interações, incluindo características do predador, da presa e do recurso basal. No entanto, não está claro como os predadores com habitat em múltiplos ecossistemas interagem com predadores de um ecossistema único, e como isso afeta o ecossistema partilhado entre eles. Além disso, não se sabe como esses efeitos complexos do tipo top­down podem ser mediados pelas características antipredatórias da presa e pela qualidade do recurso basal. Usamos as teias alimentares de invertebrados aquáticos de bromélias­tanque como um sistema modelo para investigar essas questões. Nós manipulamos a presença de um predador estritamente aquático (larvas de zigópteros) e um predador com habitats terrestre e aquático (aranha), e examinamos os efeitos na sobrevivência de presas (grupos de detritívoros com diferentes estratégias de defesa antipredatória), decomposição de detritos foliares (de duas espécies de plantas diferindo na qualidade foliar), fluxo de nitrogênio e crescimento da planta hospedeira. Para avaliar os efeitos diretos e indiretos de cada tipo de predador em múltiplos grupos de detritívoros e, finalmente, em múltiplos processos ecossistêmicos, utilizamos modelos de equações estruturais por partes (piecewiseSEM). Para cada variável resposta, isolamos a contribuição de diferentes grupos de detritívoros bem como seus efeitos globais, comparando modelos alternativos. Larvas de zigópteros e aranhas diminuíram diretamente a sobrevivência dos detritívoros e causaram múltiplos efeitos negativos indiretos na decomposição de detritos, na ciclagem de nutrientes e no crescimento da planta hospedeira. No entanto, quando os predadores coocorreram, a aranha causou um efeito negativo não consumível na larva de zigóptero, diminuindo os efeitos líquidos, diretos e indiretos, do tipo top­down na comunidade de detritívoros aquáticos e no funcionamento do ecossistema. Tanto os atributos antipredatórios dos detritívoros quanto a qualidade dos detritos modularam a força e o mecanismo dessas cascatas tróficas. A interferência do predador foi mediada por detritívoros indefesos ou com defesa parcial. Entretanto, os detritívoros com defesas antipredatórias escaparam do consumo por larvas de zigópteros, mas não por aranhas. Predadores e detritívoros afetaram a decomposição do ecossistema e a ciclagem de nutrientes apenas na presença de detritos de alta qualidade, uma vez que os detritos de baixa qualidade foram consumidos mais por micróbios do que por invertebrados. As respostas complexas deste sistema aos predadores tanto de ecossistemas receptores quanto adjacentes destacam o papel crítico da manutenção dos componentes da biodiversidade em múltiplos ecossistemas.