Spatiotemporal smoothing of water quality in a complex riverine system with physical barriers.

Freshwater ecosystems offer a variety of ecosystem services, and water quality is essential information for understanding their environment, biodiversity, and functioning. Interpolation by smoothing methods is a widely used approach to obtain temporal and/or spatial patterns of water quality from sampled data. However, when these methods are applied to freshwater systems, ignoring terrestrial areas that act as physical barriers may affect the structure of spatial autocorrelation and introduce bias into the estimates. In this study, we applied stochastic partial differential equation (SPDE) smoothing methods with barriers to spatial interpolation and spatiotemporal interpolation on water quality indices (chemical oxygen demand, phosphate phosphorus, and nitrite nitrogen) in a freshwater system in Japan. Then, we compared the estimation bias and accuracy with those of conventional non-barrier models. The results showed that the estimation bias of spatial interpolations of snapshot data was improved by considering physical barriers (5.8 % for (chemical oxygen demand, 22.5 % for phosphate phosphorus, and 21.6 % for nitrite nitrogen). The prediction accuracy was comparable to that of the non-barrier model. These were consistent with the expectation that accounting for physical barriers would capture realistic spatial correlations and reduce estimation bias, but would increase the variance of the estimates due to the limited information that can be gained from the neighbourhood. On the other hand, for spatiotemporal smoothing, the barrier model was comparable to the non-barrier model in terms of both estimation bias and prediction accuracy. This may be due to the availability of information in the time direction for interpolation. These results demonstrate the advantage of considering barriers when the available data are limited, such as snapshot data. SPDE smoothing methods can be widely applied to interpolation of various environmental and biological indices in river systems and are expected to be powerful tools for studying freshwater systems spatially and temporally.

Diverse flower-visiting responses among pollinators to multiple weather variables in buckwheat pollination.

Response diversity to environmental change among species is important for the maintenance of ecosystem services, but response diversity to changes in multiple environmental parameters is largely unexplored. Here, we examined how insect visitations to buckwheat flowers differ among species groups in response to changes in multiple weather variables and landscape structures. We found differences in responses to changes in weather conditions among insect taxonomic groups visiting buckwheat flowers. Beetles, butterflies, and wasps were more active in sunny and/or high-temperature conditions, whereas ants and non-syrphid flies showed the opposite pattern. When looking closely, the different response pattern among insect groups was itself shown to be different from one weather variable to another. For instance, large insects were responsive to temperatures more than small insects while smaller insects were responsive to sunshine duration more than large insects. Furthermore, responses to weather conditions differed between large and small insects, which agreed with the expectation that optimal temperature for insect activity depends on body size. Responses to spatial variables also differed; large insects were more abundant in fields with surrounding forests and mosaic habitats, whereas small insects were not. We suggest that response diversity at multiple spatial and temporal niche dimensions should be a focus of future studies of the biodiversity-ecosystem service relationships.

Role of landscape context in Toxoplasma gondii infection of invasive definitive and intermediate hosts on a World Heritage Island.

Free-ranging cats are invasive species threatening biodiversity worldwide. They may also impose an environmental risk to humans and livestock through the transmission of zoonotic diseases. We investigated antibody levels against Toxoplasma gondii in free-ranging cats and black rats (definitive and representative intermediate hosts) by ELISA and determined their relationships with landscape environmental factors on Tokunoshima Island, Japan, the Natural World Heritage site. We found a higher seroprevalence (>70%) in both cats and black rats in landscapes where the cattle barn density was high. This was consistent with higher density of rats revealed in our trapping survey. The spatial scale of landscape factors affecting infection was broader in cats (1 km buffer radius) than in black rats (100 m buffer radius). Both cats and rats showed an increasing trend in optical density (OD) values with increasing body weight and landscape cattle barn density, suggesting that the antibody concentration increases as the chance of exposure to T. gondii in the environment increases. Thus, management actions to stop humans from feeding cats and to control rat populations without using cats are both necessary to reduce the human health risk as well as to conserve endangered species on the island.

Positive and negative effects of land abandonment on butterfly communities revealed by a hierarchical sampling design across climatic regions.

Land abandonment may decrease biodiversity but also provides an opportunity for rewilding. It is therefore necessary to identify areas that may benefit from traditional land management practices and those that may benefit from a lack of human intervention. In this study, we conducted comparative field surveys of butterfly occurrence in abandoned and inhabited settlements in 18 regions of diverse climatic zones in Japan to test the hypotheses that species-specific responses to land abandonment correlate with climatic niches and habitat preferences. Hierarchical models that unified species occurrence and habitat preferences revealed that negative responses to land abandonment were associated with species that have cold climatic niches and use open habitats, suggesting that species negatively impacted by land abandonment will decline more due to future climate warming. Maps representing species gains and losses due to land abandonment, which were created from the model estimates, showed similar geographical patterns, but some areas exhibited high species losses relative to gains. Our hierarchical modelling approach was useful for scaling up local-scale effects of land abandonment to a macro-scale assessment, which is crucial to developing spatial conservation strategies in the era of depopulation.

The significance of region-specific habitat models as revealed by habitat shifts of grey-faced buzzard in response to different agricultural schedules.

To determine large scales habitat suitability for focal species, habitat models derived from one region are often extrapolated to others. However, extrapolation can be inappropriate due to regional variation of habitat selection. Accounting for the ecological mechanisms causing such variation is necessary to resolve this problem. We focused on grey-faced buzzards in agricultural landscapes of Japan, which show geographically different habitat selection. To determine whether this variation is caused by the difference in climatic conditions at geographical scales or the difference in agricultural practices at smaller regional scales, we surveyed distributions of buzzards and their major prey (frogs/orthopterans) in regions differing in rice-transplanting schedules within the same climatic zone. We found that buzzards preferred paddy-forest landscapes in the early transplanting regions, but grassland-forest landscapes in the late transplanting regions. Frogs were more abundant in the early transplanting regions due to flooded paddies, while the abundance of orthopterans did not differ. The regional variation in habitat selection of buzzards may be due to different prey availabilities caused by different agricultural schedules. We propose that habitat suitability assessments of organisms inhabiting agricultural landscapes should consider differences in production systems at regional scales and such regional partitioning is effective for accurate assessments.

Prevalence of serum antibodies to Toxoplasma gondii in free-ranging cats on Tokunoshima Island, Japan.

The prevalence of Toxoplasma gondii infection in free-ranging cats on Tokunoshima Island was assessed by testing 125 serum samples using anti-T. gondii IgG indirect enzyme-linked immunosorbent assay. The overall seropositivity rate was 47.2% (59/125). Seropositivity rates in cats with body weight >2.0 kg (57.4%) were significantly higher than in those with body weight ≤2.0 kg (12.5%, P<0.01). Analysis of the number of seropositive cats by settlement revealed the presence of possibly-infected cats in 17 of 23 settlements, indicating the widespread prevalence of T. gondii on the island. This is the first study to show the seroprevalence of T. gondii in free-ranging cats on Tokunoshima Island. The information revealed in this paper will help to prevent the transmission of T. gondii among cats and also in both wild and domestic animals and humans on the island.

Spatial optimization of invasive species control informed by management practices.

Optimization of spatial resource allocation is crucial for the successful control of invasive species under a limited budget but requires labor-intensive surveys to estimate population parameters. In this study, we devised a novel framework for the spatially explicit optimization of capture effort allocation using state-space population models from past capture records. We applied it to a control program for invasive snapping turtles to determine effort allocation strategies that minimize the population density over the whole area. We found that spatially heterogeneous density dependence and capture pressure limit the abundance of snapping turtles. Optimal effort allocation effectively improved the control effect, but the degree of improvement varied substantially depending on the total effort. The degree of improvement by the spatial optimization of allocation effort was only 3.21% when the total effort was maintained at the 2016 level. However, when the total effort was increased by two, four, and eight times, spatial optimization resulted in improvements of 4.65%, 8.33%, and 20.35%, respectively. To achieve the management goal for snapping turtles in our study area, increasing the current total effort by more than four times was necessary, in addition to optimizing the spatial effort. The snapping turtle population is expected to reach the target density one year after the optimal management strategy is implemented, and this rapid response can be explained by high population growth rate coupled with density-dependent feedback regulation. Our results demonstrated that combining a state-space model with optimization makes it possible to adaptively improve the management of invasive species and decision-making. The method used in this study, based on removal records from an invasive management program, can be easily applied to monitoring data for wildlife and pest control management using traps in a variety of ecosystems.

Simultaneous estimation of seasonal population density, habitat preference and catchability of wild boars based on camera data and harvest records.

Analyses of life history and population dynamics are essential for effective population control of wild mammals. We developed a model for the simultaneous estimation of seasonal changes in three parameters-population density, habitat preference and trap catchability of target animals-based on camera-trapping data and harvest records. The random encounter and staying time model, with no need for individual recognition, is the core component of the model-by combining this model with the catch-effort model, we estimated density at broad spatial scales and catchability by traps. Here, the wild boar population in central Japan was evaluated as a target population. We found that the estimated population density increased after the birth period and then decreased until the next birth period, mainly due to harvesting. Habitat preference changed seasonally, but forests having abandoned fields nearby were generally preferred throughout the season. These patterns can be explained by patterns of food availability and resting or nesting sites. Catchability by traps also changed seasonally, with relatively high values in the winter, which probably reflected changes in the attractiveness of the trap bait due to activity changes in response to food scarcity. Based on these results, we proposed an effective trapping strategy for wild boars, and discussed the applicability of our model to more general conservation and management issues.

Plant community assembly in suburban vacant lots depends on earthmoving legacy, habitat connectivity, and current mowing frequency.

In suburban regions, vacant lots potentially offer significant opportunities for biodiversity conservation. Recently, in Japan, due to an economic recession, some previously developed lands have become vacant. Little is known, however, about the legacy of earlier earthmoving, which involves topsoil removal and ground leveling before residential construction, on plant community composition in such vacant lots. To understand (dis)assembly processes in vacant lots, we studied 24 grasslands in a suburban region in Japan: 12 grasslands that had experienced earthmoving and 12 that had not. We surveyed plant community composition and species richness, and clarified compositional turnover (replacement of species) and nestedness (nonrandom species loss) by distance-based ß-diversities, which were summarized by PCoA analysis. We used piecewise structural equation modeling to examine the effects of soil properties, mowing frequency, past and present habitat connectivities on compositional changes. As a result, past earthmoving, mowing frequency, soil properties, and past habitat connectivity were found to be the drivers of compositional turnover. In particular, we found legacy effects of earthmoving: earthmoving promoted turnover from native grassland species to weeds in arable lands or roadside by altering soil properties. Mowing frequency also promoted the same turnover, implying that extensive rather than intensive mowing can modify the negative legacy effects and maintain grassland species. Decrease in present habitat connectivity marginally enhanced nonrandom loss of native grassland species (nestedness). Present habitat connectivity had a positive effect on species richness, highlighting the important roles of contemporary dispersal. Our study demonstrates that community assembly is a result of multiple processes differing in spatial and temporal scales. We suggest that extensive mowing at local scale, as well as giving a high conservation priority to grasslands with high habitat connectivity at regional scale, is the promising actions to maintain endangered native grassland species in suburban landscapes with negative legacy effects of earthmoving.

The moth specialist spider Cyrtarachne akirai uses prey scales to increase adhesion.

Contaminants decrease adhesive strength by interfering with substrate contact. Spider webs adhering to moths present an ideal model to investigate how natural adhesives overcome contamination because moths' sacrificial layer of scales rubs off on sticky silk, facilitating escape. However, Cyrtarachninae spiders have evolved gluey capture threads that adhere well to moths. Cyrtarachne capture threads contain large glue droplets oversaturated with water, readily flowing but also prone to drying out. Here, we compare the spreading and adhesion of Cyrtarachne akirai glue on intact mothwings, denuded cuticle and glass to the glue of a common orb-weaving spider, Larinioides cornutus, to understand how C. akirai glue overcomes dirty surfaces. Videos show that C. akirai's glue spreading accelerates along the underlying moth cuticle after the glue seeps beneath the moth scales-not seen on denuded cuticle or hydrophilic glass. Larinioides cornutus glue droplets failed to penetrate the moth scales, their force of adhesion thus limited by the strength of attachment of scales to the cuticle. The large size and low viscosity of C. akirai glue droplets function together to use the three-dimensional topography of the moth's scales against itself via capillary forces. Infrared spectroscopy shows C. akirai glue droplets readily lose free-flowing water. We hypothesize that this loss of water leads to increased viscosity during spreading, increasing cohesive forces during pull-off. This glue's two-phase behaviour shows how natural selection can leverage a defensive specialization of prey against themselves and highlights a new design principle for synthetic adhesives for adhering to troublesome surfaces.

Estimating range expansion of wildlife in heterogeneous landscapes: A spatially explicit state-space matrix model coupled with an improved numerical integration technique.

Dispersal as well as population growth is a key demographic process that determines population dynamics. However, determining the effects of environmental covariates on dispersal from spatial-temporal abundance proxy data is challenging owing to the complexity of model specification for directional dispersal permeability and the extremely high computational loads for numerical integration. In this paper, we present a case study estimating how environmental covariates affect the dispersal of Japanese sika deer by developing a spatially explicit state-space matrix model coupled with an improved numerical integration technique (Markov chain Monte Carlo with particle filters). In particular, we explored the environmental drivers of inhomogeneous range expansion, characteristic of animals with short dispersal. Our model framework successfully reproduced the complex population dynamics of sika deer, including rapid changes in densely populated areas and distribution fronts within a decade. Furthermore, our results revealed that the inhomogeneous range expansion of sika deer seemed to be primarily caused by the dispersal process (i.e., movement barriers in fragmented forests) rather than population growth. Our state-space matrix model enables the inference of population dynamics for a broad range of organisms, even those with low dispersal ability, in heterogeneous landscapes, and could address many pressing issues in conservation biology and ecosystem management.

Supersaturation with water explains the unusual adhesion of aggregate glue in the webs of the moth-specialist spider, Cyrtarachne akirai.

Orb webs produced by araneoid spiders depend upon aggregate glue-coated capture threads to retain their prey. Moths are challenging prey for most spiders because their scales detach and contaminate the glue droplets, significantly decreasing adhesion. Cyrtarachne are moth-specialist orb-weaving spiders whose capture threads adhere well to moths. We compare the adhesive properties and chemistry of Cyrtarachne aggregate glue to other orb-weaving spiders to test hypotheses about their structure, chemistry and performance that could explain the strength of Cyrtarachne glue. We show that the unusually large glue droplets on Cyrtarachne capture threads make them approximately 8 times more adhesive on glass substrate than capture threads from typical orb-weaving species, but Cyrtarachne adhesion is similar to that of other species after normalization by glue volume. Glue viscosity reversibly changes over 1000-fold in response to atmospheric humidity, and the adhesive strength of many species of orb spiders is maximized at a viscosity of approximately 105-106 cst where the contributions of spreading and bulk cohesion are optimized. By contrast, viscosity of Cyrtarachne aggregate glue droplets is approximately 1000 times lower at maximum adhesive humidity, likely facilitating rapid spreading across moth scales. Water uptake by glue droplets is controlled, in part, by hygroscopic low molecular weight compounds. NMR showed evidence that Cyrtarachne glue contains a variety of unknown low molecular weight compounds. These compounds may help explain how Cyrtarachne produces such exceptionally large and low viscosity glue droplets, and also why these glue droplets rapidly lose water volume after brief ageing or exposure to even slightly dry (e.g. < 80% RH) conditions, permanently reducing their adhesion. We hypothesize that the combination of large glue droplet size and low viscosity helps Cyrtarachne glue to penetrate the gaps between moth scales.

Silk structure rather than tensile mechanics explains web performance in the moth-specialized spider, Cyrtarachne.

Orb webs intercept and retain prey so spiders may subdue them. Orb webs are composed of sticky, compliant spirals of capture silk spun across strong, stiff major ampullate silk threads. Interplay between differences in the mechanical properties of these silks is crucial for prey capture. Most orb webs depend upon insects contacting several radial and capture threads for successful retention. Moths, however, escape quickly from most orb webs due to the sacrificial scales covering their bodies. Cyrtarachne orb webs are unusual as they contain a reduced number of capture threads and moths stick unusually well to single threads. We aimed to determine how the tensile properties of the capture spiral and radial threads spun by Cyrtarachne operate in retention of moth prey. A NanoBionix UTM was used to quantify the material properties of flagelliform and major ampullate threads to test if Cyrtarachne's reduced web architecture is accompanied by improvements in tensile performance of its silk. Silk threads showed tensile properties typical of less-specialized orb-weavers, with the exception of high extensibility in radial threads. Radial thread diameters were 62.5% smaller than flagelliform threads, where commonly the two are roughly similar. We utilized our tensile data to create a finite element model of Cyrtarachne's web to investigate energy dissipation during prey impact. Large cross-sectional area of the flagelliform threads played a key role in enabling single capture threads to withstand prey impact. Rather than extraordinary silk, Cyrtarachne utilizes structural changes in the size and attachment of silk threads to facilitate web function.

Temporal trends in arthropod abundances after the transition to organic farming in paddy fields.

Organic farming aims to reduce the effect on the ecosystem and enhance biodiversity in agricultural areas, but the long-term effectiveness of its application is unclear. Assessments have rarely included various taxonomic groups with different ecological and economic roles. In paddy fields with different numbers of years elapsed since the transition from conventional to organic farming, we investigated changes in the abundance of insect pests, generalist predators, and species of conservation concern. The abundance of various arthropods exhibited diverse trends with respect to years elapsed since the transition to organic farming. Larval lepidopterans, Tetragnatha spiders, and some planthoppers and stink bugs showed non-linear increases over time, eventually reaching saturation, such as the abundance increasing for several years and then becoming stable after 10 years. This pattern can be explained by the effects of residual pesticides, the lag time of soil mineralization, and dispersal limitation. A damselfly (Ischnura asiatica) did not show a particular trend over time, probably due to its rapid immigration from source habitats. Unexpectedly, both planthoppers and some leafhoppers exhibited gradual decreases over time. As their abundances were negatively related to the abundance of Tetragnatha spiders, increased predation by natural enemies might gradually decrease these insect populations. These results suggest that the consideration of time-dependent responses of organisms is essential for the evaluation of the costs and benefits of organic farming, and such evaluations could provide a basis for guidelines regarding the length of time for organic farming to restore biodiversity or the economic subsidy needed to compensate for pest damage.

Habitat modification by invasive crayfish can facilitate its growth through enhanced food accessibility.

BACKGROUND: Invasive ecosystem engineers can facilitate their invasions by modifying the physical environment to improve their own performance, but this positive feedback process has rarely been tested empirically except in sessile organisms. The invasive crayfish Procambarus clarkii is an ecosystem engineer that destroys aquatic macrophytes, which provide a physical refuge for animal prey, and this destruction is likely to enhance vulnerability to predators. Using two series of mesocosm experiments, we tested the hypothesis that the invasive crayfish increases its feeding efficiency on animal prey by reducing submerged macrophytes, thus increasing its individual growth rate in a positive density-dependent manner. RESULTS: In the first experiment, increasing crayfish density reduced both macrophytes and animal prey (dragonfly and chironomid larvae) and, importantly, increased the growth rate of individual crayfish, in accordance with our expectation. In the second experiment, we used artificial macrophytes to clarify whether the physical architecture of macrophytes itself protects animal prey and limits crayfish growth rate. Increasing the artificial macrophyte quantity not only increased the survival of animal prey, but also retarded the crayfish growth rate. CONCLUSIONS: We conclude that macrophytes strengthen bottom-up control of crayfish, but this effect can be relaxed by increasing the density of crayfish via reduction in macrophytes. This positive feedback process may explain the crayfish outbreaks and regime shifts occasionally observed in invaded freshwater ecosystems.

Human interest meets biodiversity hotspots: A new systematic approach for urban ecosystem conservation.

Creating a win-win relationship between biodiversity and human well-being is one of the major current challenges for environmental policy. One way to approach this challenge is to identify sites with both high biodiversity and high human interest in urban areas. Here, we propose a new systematic approach to identify such sites by using land prices and biodiversity indexes for butterflies and birds from a nationwide perspective. As a result, we found sites that are valuable to humans and to other organisms, including national red-list species, and they are located in sites with cultural heritages and near seaside. By referencing the habitat features and landscape characteristics of these sites, we can establish high quality environments that provide a benefit to both humans and biodiversity in urban landscapes.

Context Dependent Effect of Landscape on the Occurrence of an Apex Predator across Different Climate Regions.

In studies of habitat suitability at landscape scales, transferability of species-landscape associations among sites are likely to be critical because it is often impractical to collect datasets across various regions. However, limiting factors, such as prey availability, are not likely to be constant across scales because of the differences in species pools. This is particularly true for top predators that are often the target for conservation concern. Here we focus on gray-faced buzzards, apex predators of farmland-dominated landscapes in East Asia. We investigated context dependency of "buzzard-landscape relationship", using nest location datasets from five sites, each differing in landscape composition. Based on the similarities of prey items and landscape compositions across the sites, we determined several alternative ways of grouping the sites, and then examined whether buzzard-landscape relationship change among groups, which was conducted separately for each way of grouping. As a result, the model of study-sites grouping based on similarities in prey items showed the smallest ΔAICc. Because the terms of interaction between group IDs and areas of broad-leaved forests and grasslands were selected, buzzard-landscape relationship showed a context dependency, i.e., these two landscape elements strengthen the relationship in southern region. The difference in prey fauna, which is associated with the difference in climate, might generate regional differences in the buzzard-landscape associations.

Time-scale dependency of host plant biomass- and trait-mediated indirect effects of deer herbivory on a swallowtail butterfly.

Despite recent attempts to quantify the relative strength of density- and trait-mediated indirect effects, rarely has the issue been properly addressed at the population level. Most research is based on short-term small-scale experiments in which behavioural and/or physiological responses prevail. Here, we estimated the time-scales during which density- and trait-mediated effects manifest, as well as the strength of these effects, using an interaction chain with three organisms (deer-plant-butterfly). A hierarchical Bayesian model was performed by using a long-term data set of deer density in the Boso Peninsula, central Japan (where local densities differ spatially and temporally) as well as densities of the swallowtail butterfly Byasa alcinous and its host plant Aristolochia kaempferi. The time-scale effect of deer on plant quantity and quality was estimated according to the degree of carry-over effects. The negative influence on leaf density showed a temporal saturation pattern over the long term, while the positive influence on leaf quality due to resprouting of leaves after deer browsing showed no clear temporal trend. The net indirect effect changed from positive to negative with time, with the negative density-mediated effect becoming prominent in the long term. Our novel approach is widely applicable in assessing the dynamic impacts of wildlife if the spatio-temporal variability of expansion and/or invasion history is known.

Molecular phylogeny of moth-specialized spider sub-family Cyrtarachninae, which includes bolas spiders.

The evolutionary process of the unique web architectures of spiders of the sub-family Cyrtarachninae, which includes the triangular web weaver, bolas spider, and webless spider, is thought to be derived from reduction of orbicular 'spanning-thread webs' resembling ordinal orb webs. A molecular phylogenetic analysis was conducted to explore this hypothesis using orbicular web spiders Cyrtarachne, Paraplectana, Poecilopachys, triangular web spider Pasilobus, bolas spiders Ordgarius and Mastophora, and webless spider Celaenia. The phylogeny inferred from partial sequences of mt-COI, nuclear 18S-rRNA and 28S-rRNA showed that the common ancestor of these spiders diverged into two clades: a spanning-thread web clade and a bolas or webless clade. This finding suggests that the triangular web evolved by reduction of an orbicular spanning web, but that bolas spiders evolved in the early stage, which does not support the gradual web reduction hypothesis.

Adjustment of web-building initiation to high humidity: a constraint by humidity-dependent thread stickiness in the spider Cyrtarachne.

Cyrtarachne is an orb-weaving spider belonging to the subfamily Cyrtarachninae (Araneidae) which includes triangular-web-building Pasilobus and bolas spiders. The Cyrtarachninae is a group of spiders specialized in catching moths, which is thought to have evolved from ordinary orb-weaving araneids. Although the web-building time of nocturnal spiders is in general related to the time of sunset, anecdotal evidence has suggested variability of web-building time in Cyrtarachne and its closely related genera. This study has examined the effects of temperature, humidity, moonlight intensity, and prey (moths) availability on web-building time of Cyrtarachne bufo, Cyrtarachne akirai, and Cyrtarachne nagasakiensis. Generalized linear mixed model (GLMM) have revealed that humidity, and not prey availability, was the essential variable that explained the daily variability of web-building time. Experiments measuring thread stickiness under different humidities showed that, although the thread of Cyrtarachne was found to have strong stickiness under high humidity, low humidity caused a marked decrease of thread stickiness. By contrast, no obvious change in stickiness was seen in an ordinary orb-weaving spider, Larinia argiopiformis. These findings suggest that Cyrtarachne adjusts its web-building time to favorable conditions of high humidity maintaining strong stickiness, which enables the threads to work efficiently for capturing prey.