Resilience of spider communities affected by a range of silvicultural treatments in a temperate deciduous forest stand.

To secure the ecosystem services forests provide, it is important to understand how different management practices impact various components of these ecosystems. We aimed to uncover how silvicultural treatments affected the ground-dwelling spider communities during the first five years of a forest ecological experiment. In an oak-hornbeam forest stand, five treatments, belonging to clear-cutting, shelterwood and continuous cover forestry systems, were implemented using randomised complete block design. Spiders were sampled by pitfall traps, and detailed vegetation, soil and microclimate data were collected throughout the experiment. In the treatment plots spider abundance and species richness increased marginally. Species composition changes were more pronounced and treatment specific, initially diverging from the control plots, but becoming more similar again by the fifth year. These changes were correlated mostly to treatment-related light intensity and humidity gradients. The patchy implementation of the treatments induced modest increase in both gamma and beta diversity of spiders in the stand. Overall, spiders gave a prompt and species specific response to treatments that was by the fifth year showing signs of relatively quick recovery to pre-treatment state. At the present fine scale of implementation the magnitude of changes was not different among forestry treatments, irrespective of their severity.

Environmental drivers of forest biodiversity in temperate mixed forests - A multi-taxon approach.

Harmonization of timber production and forest conservation is a major challenge of modern silviculture. For the establishment of ecologically sustainable forest management, the management-related environmental drivers of multi-taxon biodiversity should be explored. Our study reveals those environmental variables related to tree species diversity and composition, stand structure, litter and soil conditions, microclimate, landscape, and land-use history that determine species richness and composition of 11 forest-dwelling organism groups. Herbs, woody regeneration, ground-floor and epiphytic bryophytes, epiphytic lichens, terricolous saprotrophic, ectomycorrhizal, and wood-inhabiting macrofungi, spiders, carabid beetles, and birds were sampled in West Hungarian mature mixed forests. The correlations among the diversities and compositions of different organism groups were also evaluated. Drivers of organism groups were principally related to stand structure, tree species diversity and composition, and microclimate, while litter, soil, landscape, and land-use historical variables were less influential. The complex roles of the shrub layer, deadwood, and the size of the trees in determining the diversity and composition of various taxa were revealed. Stands with more tree species sustained higher stand-level species richness of several taxa. Besides, stands with different dominant tree species harbored various species communities of organism groups. Therefore, landscape-scale diversity of dominant tree species may enhance the diversity of forest-dwelling communities at landscape level. The effects of the overstory layer on forest biodiversity manifested in many cases via microclimate conditions. Diversity of organism groups showed weaker relationship with the diversity of other taxa than with environmental variables. According to our results, the most influential drivers of forest biodiversity are under the direct control of the actual silvicultural management. Heterogeneous stand structure and tree species composition promote the different organism groups in various ways. Therefore, the long-term maintenance of the structural and compositional heterogeneity both at stand and landscape scale is an important aspect of ecologically sustainable forest management.

Landscape structure affects the sunflower visiting frequency of insect pollinators.

Mass-flowering crop monocultures, like sunflower, cannot harbour a permanent pollinator community. Their pollination is best secured if both managed honey bees and wild pollinators are present in the agricultural landscape. Semi-natural habitats are known to be the main foraging and nesting areas of wild pollinators, thus benefiting their populations, whereas crops flowering simultaneously may competitively dilute pollinator densities. In our study we asked how landscape structure affects major pollinator groups' visiting frequency on 36 focal sunflower fields, hypothesising that herbaceous semi-natural (hSNH) and sunflower patches in the landscape neighbourhood will have a scale-dependent effect. We found that an increasing area and/or dispersion of hSNH areas enhanced the visitation of all pollinator groups. These positive effects were scale-dependent and corresponded well with the foraging ranges of the observed bee pollinators. In contrast, an increasing edge density of neighbouring sunflower fields resulted in considerably lower visiting frequencies of wild bees. Our results clearly indicate that the pollination of sunflower is dependent on the composition and configuration of the agricultural landscape. We conclude that an optimization of the pollination can be achieved if sufficient amount of hSNH areas with good dispersion are provided and mass flowering crops do not over-dominate the agricultural landscape.

Simplified and rapid staining of leafhopper salivary sheaths in plant tissues for electrical penetration graph waveform correlations.

Herbivorous insects in the order Hemiptera use piercing-sucking mouthparts to utilize plant sap. Among them salivary sheath feeders penetrate into the plant by their flexible stylets to reach vascular elements. Manoeuvering stylets in plant tissues is aided by the creation of salivary sheaths, which solidify from proteinous gelling saliva and remain as lasting artefacts in the plant tissues. Studying their structure reveals hidden details of the feeding behaviour and the transmission of pathogens in case of vector insects. One important aspect of studying salivary sheaths is that it can be used to confirm the biological function of electropenetrography (EPG) waveform patterns. Previously, complex and vaguely documented histological methods have been used to observe salivary sheath structure. Building on existing methodologies, we report a simplified histological procedure where each step was optimized to offer a rapid process that does not require special equipment, can be applied to many samples, has good success rate and a low cost of errors in terms of time and materials. We describe the procedure, using a Psammotettix alienus - barley model system, in three steps. (i) Clarification of entire plant parts and pre-staining salivary sheaths with aqueous fuchsin. This step allows to identify salivary sheath starting points on the surface. (ii) Knowing salivary sheath location, using hand sectioning, produce a single c. 60 µm section that contains the entire salivary sheath. (iii) Counterstain the section with methylene green and, after further clarification, study under light microscope in a glycerol - ethanol embedding solution, without fixed mounting.

Consequences of rapid development owing to cohort splitting: just how costly is it to hurry?

In cohort splitting, diverging sub-cohorts may show substantial differences in their growth and developmental rates. Although in the past, causes and adaptive value of cohort splitting were studied in detail, individual-level consequences of cohort splitting are still rather overlooked. Life history theory predicts that considerably increased growth and developmental rates should be traded off against other costly life history traits. However, it is not clear whether one should expect such associations in adaptive developmental plasticity scenarios, because natural selection might have promoted genotypes that mitigate those potential costs of rapid development. To address these contrasting propositions, we assessed life history traits in the wolf spider Pardosa agrestis, both collected from natural habitat and reared in laboratory. We found that some traits are negatively associated with developmental rates in spiders collected from the wild, but these associations were relaxed to a considerable extent in laboratory-reared specimens. In general, we observed no consistent trend for the presence of developmental costs, although some results might suggest higher relative fecundity costs in rapidly developing females. Our study provides a detailed approach to the understanding of individual-level consequences of cohort splitting, and to the associations between key life history traits in adaptive developmental plasticity scenarios.

Consuming alternative prey does not influence the DNA detectability half-life of pest prey in spider gut contents.

BACKGROUND: Key natural enemy-pest interactions can be mapped in agricultural food webs by analysing predator gut content for the presence of a focal pest species. For this, PCR-based approaches are the most widely used methods providing the incidence of consumption of a focal pest in field sampled predators. To interpret such data the rate of prey DNA decay in the predators' gut, described by DNA detectability half-life (t 1/2), is needed. DNA decay may depend on the presence of alternative prey in the gut of generalist predators, but this effect has not been investigated in one of the major predatory arthropod groups, spiders. METHODS: In a laboratory feeding experiment, we determined t 1/2 of the key cereal pest virus vector leafhopper Psammotettix alienus in the digestive tracts of its natural enemy, the spider Tibellus oblongus. We followed the fate of prey DNA in spiders which received only the focal prey as food, or as an alternative prey treatment they also received a meal of fruit flies after leafhopper consumption. After these feeding treatments, spiders were starved for variable time intervals prior to testing for leafhopper DNA in order to establish t 1/2. RESULTS: We created a PCR protocol that detects P. alienus DNA in its spider predator. The protocol was further calibrated to the digestion speed of the spider by establishing DNA decay rate. Detectability limit was reached at 14 days, where c. 10% of the animals tested positive. The calculated t 1/2 = 5 days value of P. alienus DNA did not differ statistically between the treatment groups which received only the leafhopper prey or which also received fruit fly. The PCR protocol was validated in a field with known P. alienus infestation. In this applicability trial, we showed that 12.5% of field collected spiders were positive for the leafhopper DNA. We conclude that in our model system the presence of alternative prey did not influence the t 1/2 estimate of a pest species, which makes laboratory protocols more straightforward for the calibration of future field data.

Taxon-specific responses to different forestry treatments in a temperate forest.

There are only few studies that explore the ecological consequences of forest management on several organism groups. We studied the short-term effects of four forestry treatments including preparation cutting, clear-cutting, retention tree group and gap-cutting in a temperate managed forest on the assemblage structure of understory plants, enchytraeid worms, spiders and ground beetles. Here we show, that the effect of treatments on the different facets of assemblage structure was taxon-specific. Clear-cutting and retention tree group strongly impoverished enchytraeids assemblages. Even if the species richness and cover of plants increased in clear-cutting and gap-cutting, their species composition moderately changed after treatments. For spiders only their species composition was influenced by the treatments, while the response of ground beetles was slightly affected. Short-term effect of forest management interventions on biodiversity might be compensated by the dispersal (spiders, ground beetles) and resilience (plants) of organism groups, however sedentary soil organism showed high sensitivity.

Natural enemies partially compensate for warming induced excess herbivory in an organic growth system.

Predators can limit prey abundance and/or levels of activity. The magnitudes of these effects are contingent on predator and prey traits that may change with environmental conditions. Aberrant thermal regimes could disrupt pest suppression through asymmetric effects, e.g. heat-sensitive predator vs. heat-tolerant prey. To explore potential effects of warming on suppressing pests and controlling herbivory in a vegetable crop, we performed laboratory experiments exposing an important pest species to two spider predator species at different temperatures. Heat tolerance was characterised by the critical thermal maxima parameter (CTM50) of the cucumber beetle (Diabrotica undecimpunctata), wolf spider (Tigrosa helluo), and nursery web spider (Pisaurina mira). Cucumber beetles and wolf spiders were equally heat tolerant (CTM50 > 40 °C), but nursery web spiders had limited heat tolerance (CTM50 = 34 °C). Inside mesocosms, beetle feeding increased with temperature, wolf spiders were always effective predators, nursery web spiders were less lethal at high temperature (38 °C). Neither spider species reduced herbivory at ambient temperature (22 °C), however, at warm temperature both species reduced herbivory with evidence of a dominant non-consumptive effect. Our experiments highlight the contingent nature of predator-prey interactions and suggest that non-consumptive effects should not be ignored when assessing the impact of temperature change.

Cool Headed Individuals Are Better Survivors: Non-Consumptive and Consumptive Effects of a Generalist Predator on a Sap Feeding Insect.

Non-consumptive effects (NCEs) of predators are part of the complex interactions among insect natural enemies and prey. NCEs have been shown to significantly affect prey foraging and feeding. Leafhopper's (Auchenorrhyncha) lengthy phloem feeding bouts may play a role in pathogen transmission in vector species and also exposes them to predation risk. However, NCEs on leafhoppers have been scarcely studied, and we lack basic information about how anti-predator behaviour influences foraging and feeding in these species. Here we report a study on non-consumptive and consumptive predator-prey interactions in a naturally co-occurring spider-leafhopper system. In mesocosm arenas we studied movement patterns during foraging and feeding of the leafhopper Psammotettix alienus in the presence of the spider predator Tibellus oblongus. Leafhoppers delayed feeding and fed much less often when the spider was present. Foraging movement pattern changed under predation risk: movements became more frequent and brief. There was considerable individual variation in foraging movement activity. Those individuals that increased movement activity in the presence of predators exposed themselves to higher predation risk. However, surviving individuals exhibited a 'cool headed' reaction to spider presence by moving less than leafhoppers in control trials. No leafhoppers were preyed upon while feeding. We consider delayed feeding as a "paradoxical" antipredator tactic, since it is not necessarily an optimal strategy against a sit-and-wait generalist predator.

Can field-scale habitat diversification enhance the biocontrol potential of spiders?

The study investigated the effect of strip management in alfalfa on the abundance and diversity of spiders. In strip management, narrow strips were left unmown in the crop, which were then shifted at next mowing. In an experimental field, out of the six 50 x 50-m blocks, three received strip management and three were managed traditionally (ie the whole block was mown). Our main hypothesis was that unmown strips will contain a more diverse and abundant spider assemblage, which will increase spider numbers in the mown parts of the crop. Over the 3 years of the study, unmown strips contained a spider assemblage of more than 50% higher abundance than in control alfalfa. Species diversity was also greater in the strips, and the presence of specific indicator species could be shown. We found that five dominant species comprised over 75% of spiders in the strips, and they were the same species as those that are dominant in alfalfa, where they represent 85% of all spiders. In contrast, a neighbouring meadow, which was a control in the study, also had high diversity, but different dominant species. The main result of the experiment was negative in that elevated abundance of spiders in unmown strips did not raise spider numbers in the mown strips of alfalfa, where abundance was only marginally higher than in the control alfalfa. We can speculate that better ecological conditions in unmown strips attract and conserve spiders from neighbouring areas and from the crop at mowing. We suggest that if diversification is more interspersed within the crop, we have a higher chance of increasing the number of natural enemies and bringing them closer to pests, so that they can interact more intensively.

A spider population in flux: selection and abandonment of artificial web-sites and the importance of intraspecific interactions in Lephthyphantes tenuis (Araneae: Linyphiidae) in wheat.

Lepthyphantes tenuis, a small sheet-webbuilding linyphiid spider is one of the most abundant spider species of cereal fields in Europe. In the present study we examined the process of web-site selection and web-site tenacity by adult females of this species in a winter wheat field. Spiders were selective in their choice of web-site. Different immigration rates into various manipulated web-sites, in field and laboratory, suggested that structural support and suitable micro-climate (high humidity) are the most important factors in the selection. Small holes dug in the ground were the most favoured web-sites. Web-site occupation was influenced by the presence of other conspecific spiders. Territorial contests occurred between spiders attempting to occupy the same web, these almost invariably led to the take-over of the web when the intruder was heavier. Interference, but also a certain level of tolerance, between spiders within the same web-site but in different webs was suggested by direct and indirect evidence. Many holes supported two or even three spiders in vertically stratified webs. Leaving probability of marked spiders was significantly higher in multiply occupied holes than in holes with a single web. Comparison with the results of a no-interference stochastic model showed that multiple occupancy in nature is less frequent than predicted by the model. There was further evidence for weak extra-web-interference between spiders in that multiple occupancy was even less frequent and overall occupancy was lower in web-sites which were packed close to each other. However, a level of tolerance for crowding is shown by the fact that closely packed hole colonies supported a spider density 13 times higher than in natural web-sites in the field. A marking experiment was carried out to gain information on web-site tenacity (i.e. the length of time a spider spends in a web-site) and abandonment. The average duration of tenacity was less than 2 days. A random loss function gave a good fit to the tenacity distribution and suggested that spiders abandoned web-sites randomly with a fixed leaving probability of c. 0.5. Individual webs were often used consecutively by more than one spider, and some spiders built more than one web in the same web-site. Calculations showed that abandonment is the most frequent leaving mode, while take-over by contest between spiders and disappearance due to destruction were some-what less frequent and equally likely modes of ending tenacity. It is suggested that the apparent contradiction between the selectiveness and competitiveness of spiders for web-sites and the relatively short tenacity observed can be resolved by hypothesising that spiders leave websites soon because they apply the strategy of spreading risk: spiders by frequently moving from one web-site to another distribute their reproductive efforts across several localities. This hypothesis is further supported by changes in web-site preference and ballooning behaviour at the onset of the reproductive stage in L. tenuis.