Patterns of avian tree usage in the primeval temperate forests of Bialowieza National Park.

Species distribution and resource utilization are a fundamental aspect of ecology. By analyzing the tree space usage by birds and determining the species composition of birds across different parts of trees, our study could shed light on the mechanisms shaping co-occurrence patterns in bird communities. Therefore, our study aimed to determine the species composition of birds across different parts of trees. We investigated whether species richness differs between positions on a tree and how these positions influence the probability of occurrence of the 10 most frequently observed bird species. To achieve this, we observed birds within permanent plots in Bialowieza National Park (BNP) and analyzed the distribution patterns of birds within six vertical and three horizontal sectors of trees. The compositional dissimilarity between tree sectors was assessed using detrended correspondence analysis. We employed generalized linear mixed-effects models to examine differences in species richness. The majority of the BNP bird community was associated with the branches, while other birds occupied the tree crown trunks and the understory trunks. Species richness was the highest on branches in the crown part of trees, followed by lower species richness on trunks associated with crowns, and the lowest richness was observed on branches and trunks in the understory. These results indicate that branches in the middle and lower parts of the crown serve as avian diversity hotspots on trees, likely due to the abundance of various food sources. The differing patterns of tree usage by specific bird species may suggest the avoidance of interspecific competition for resources. The study results of tree usage by bird species obtained in the primeval forest provides a reference point for studies conducted in human-altered woods.

Movement and Home Range of Amur Soft-Shell Turtle (Pelodiscus maackii) in the Ussuri River, Heilongjiang Province, China.

Comprehensively understanding the spatial ecology and habitat preferences of endangered species is essential for population restoration and conservation. We investigated the home range and movement of the endangered Amur soft-shell turtle (Pelodiscus maackii) in the Ussuri River, Heilongjiang Province, Northeastern China. The study involved tracking 19 Amur soft-shell turtles from late June to mid-October, 2022, resulting in complete and partial home range size data for eight subadults and two adults, respectively. The primary analysis focused on eight subadults, and the models that best described daily movement were identified. We also explored the potential factors influencing home range size. The mean movement rate ranged from 39.18 ± 20.04 m/day to 72.45 ± 29.36 m/day and was positively correlated with the linear home range and water temperature. The most enlightening estimation of home range was derived from a 95% kernel density estimate, utilizing likelihood cross-validation smoothing while adhering to constraints delineated by the river boundaries. The average size of the home range was determined to be 1.02 hectares and displayed no correlation with body size. Subadults tended to establish well-defined home ranges over time, whereas defining home ranges for adults proved challenging. This research addresses a gap regarding the ecology of the Amur soft-shell turtle and provides a foundation for future conservation plans.

From little things big things grow: enhancement of an acoustic telemetry network to monitor broad-scale movements of marine species along Australia's east coast.

BACKGROUND: Acoustic telemetry has become a fundamental tool to monitor the movement of aquatic species. Advances in technology, in particular the development of batteries with lives of > 10 years, have increased our ability to track the long-term movement patterns of many species. However, logistics and financial constraints often dictate the locations and deployment duration of acoustic receivers. Consequently, there is often a compromise between optimal array design and affordability. Such constraints can hinder the ability to track marine animals over large spatial and temporal scales. Continental-scale receiver networks have increased the ability to study large-scale movements, but significant gaps in coverage often remain. METHODS: Since 2007, the Integrated Marine Observing System's Animal Tracking Facility (IMOS ATF) has maintained permanent receiver installations on the eastern Australian seaboard. In this study, we present the recent enhancement of the IMOS ATF acoustic tracking infrastructure in Queensland to collect data on large-scale movements of marine species in the northeast extent of the national array. Securing a relatively small initial investment for expanding receiver deployment and tagging activities in Queensland served as a catalyst, bringing together a diverse group of stakeholders (research institutes, universities, government departments, port corporations, industries, Indigenous ranger groups and tourism operators) to create an extensive collaborative network that could sustain the extended receiver coverage into the future. To fill gaps between existing installations and maximise the monitoring footprint, the new initiative has an atypical design, deploying many single receivers spread across 2,100 km of Queensland waters. RESULTS: The approach revealed previously unknown broad-scale movements for some species and highlights that clusters of receivers are not always required to enhance data collection. However, array designs using predominantly single receiver deployments are more vulnerable to data gaps when receivers are lost or fail, and therefore "redundancy" is a critical consideration when designing this type of array. CONCLUSION: Initial results suggest that our array enhancement, if sustained over many years, will uncover a range of previously unknown movements that will assist in addressing ecological, fisheries, and conservation questions for multiple species.

The impact of within-host coinfection interactions on between-host parasite transmission dynamics varies with spatial scale.

Within-host interactions among coinfecting parasites can have major consequences for individual infection risk and disease severity. However, the impact of these within-host interactions on between-host parasite transmission, and the spatial scales over which they occur, remain unknown. We developed and apply a novel spatially explicit analysis to parasite infection data from a wild wood mouse (Apodemus sylvaticus) population. We previously demonstrated a strong within-host negative interaction between two wood mouse gastrointestinal parasites, the nematode Heligmosomoides polygyrus and the coccidian Eimeria hungaryensis, using drug-treatment experiments. Here, we show this negative within-host interaction can significantly alter the between-host transmission dynamics of E. hungaryensis, but only within spatially restricted neighbourhoods around each host. However, for the closely related species E. apionodes, which experiments show does not interact strongly with H. polygyrus, we did not find any effect on transmission over any spatial scale. Our results demonstrate that the effects of within-host coinfection interactions can ripple out beyond each host to alter the transmission dynamics of the parasites, but only over local scales that likely reflect the spatial dimension of transmission. Hence there may be knock-on consequences of drug treatments impacting the transmission of non-target parasites, altering infection risks even for non-treated individuals in the wider neighbourhood.

Dispersal evolution and eco-evolutionary dynamics in antagonistic species interactions.

Dispersal evolution modifies diverse spatial processes, such as range expansions or biological invasions of single species, but we are currently lacking a realistic vision for metacommunities. Focusing on antagonistic species interactions, we review existing theory of dispersal evolution between natural enemies, and explain how this might be relevant for classic themes in host-parasite evolutionary ecology, namely virulence evolution or local adaptation. Specifically, we highlight the importance of considering the simultaneous (co)evolution of dispersal and interaction traits. Linking such multi-trait evolution with reciprocal demographic and epidemiological feedbacks might change basic predictions about coevolutionary processes and spatial dynamics of interacting species. Future challenges concern the integration of system-specific disease ecology or spatial modifiers, such as spatial network structure or environmental heterogeneity.

Female Deer Movements Relative to Firearms Hunting in Northern Georgia, USA.

Perceived risk associated with hunters can cause white-tailed deer (Odocoileus virginianus) to shift their activity away from key foraging areas or alter normal movements, which are important considerations in managing hunting and its effects on a population. We studied the effects of seven firearms hunts on the movements of 20 female deer in two Wildlife Management Areas within the Chattahoochee National Forest of northern Georgia, USA, during the 2018-2019 and 2019-2020 hunting seasons. Deer populations and the number of hunters in our study area have declined significantly since the 1980s. In response, hunting regulations for the 2019-2020 hunting season eliminated opportunities for harvesting female deer. To evaluate the indirect effects of antlered deer hunting on non-target female deer, we calculated 90% utilization distributions (UDs), 50% UDs, and step lengths for pre-hunt, hunt, and post-hunt periods using the dynamic Brownian bridge movement model. Data included 30 min GPS locations for 44 deer-hunt combinations. Pre-hunt 50% UDs (x- = 7.0 ha, SE = 0.4 ha) were slightly greater than both hunt (x- = 6.0 ha, SE = 0.3 ha) and post-hunt (x- = 6.0 ha, SE = 0.2 ha) 50% UDs (F = 3.84, p = 0.03). We did not detect differences in step length, nor did we detect differences in size or composition of 90% UDs, among the periods. Overall, our results suggest that the low level of hunting pressure in our study area and lack of exposure to hunters led to no biologically significant changes in female deer movements. To the extent of the findings presented in this paper, adjustments to the management of hunting in our study area do not appear to be necessary to minimize hunting-related disturbances for female deer. However, managers should continue to consider female deer behavior when evaluating future changes to hunting regulations.

The Spatial Ecology of Nuisance Crocodiles: Movement Patterns of Relocated American Crocodiles (Crocodylus acutus) in Guanacaste, Costa Rica.

Anthropogenic alterations of the environment have increased, highlighting the need for human-wildlife coexistence and conflict mitigation. Spatial ecology, and the use of passive satellite movement technology in particular, has been used to identify patterns in human-wildlife conflict as a function of shared resources that present potential for dangerous situations. Here, we aim to remotely identify patterns indicative of human-crocodile conflict in Guanacaste, Costa Rica by exploring site fidelity and diverse modes of movement (i.e., land and water) across space between nuisance (relocated) and non-nuisance (wild) crocodiles. Advanced satellite remote sensing technology provided near-constant movement data on individuals at the regional scale. Telonics Iridium SeaTrkr-4370-4 transmitters were used with modified crocodilian fitting. Results indicate that relocated crocodiles exhibited large-scale movements relative to wild crocodiles. Nuisance relocated crocodiles either returned to the area of nuisance or potentially attempted to in short time frames. The results presented here highlight the need for alternative management strategies that facilitate relocation efficacy.

From forests to fields: investigating Culicoides (Diptera: Ceratopogonidae) abundance and diversity in cattle pastures and adjacent woodlands.

Culicoides Latreille (Diptera: Ceratopogonidae) biting midges are hematophagous flies that feed on wild and domestic ruminants. They can transmit arboviruses, such as bluetongue virus (BTV) and epizootic hemorrhagic disease virus (EHDV), which circulate in the United States. Larvae occupy a range of aquatic and semiaquatic habitats, and disperse short distances from their development sites. In the southeastern United States, there are limited studies on the abundance and diversity of Culicoides in wooded and adjacent livestock pasture habitats. In this study, we characterized Culicoides diversity and abundance within these distinct habitat types. BG-Sentinel and CDC miniature suction traps baited with CO2 or UV-light were placed in wooded and pasture habitats at 2 locations on a university beef farm in Savoy, Arkansas. Traps were set once per week for 9 wk during August-October of 2021 and 2022. Fifteen species were collected during this study, and the 2 most abundant species were Culicoides haematopotus Malloch and Culicoides stellifer Coquillett. There was a significant effect of site and location on C. haematopotus collections, and a significant effect and interaction of site and trap on C. stellifer collections. In the woods, significantly more C. stellifer were collected from CDC-UV traps, while in the pasture significantly more were collected in CDC-CO2 traps. These data suggest that C. stellifer, a putative vector of BTV and EHDV in the southeast, may be traveling into the pasture to host-seek, while C. haematopotus remains primarily in wooded areas. This study reveals community differences between these habitat types and implications for Culicoides control.

Heterogeneous dispersal networks to improve biodiversity science.

Dispersal has a key role in shaping spatial patterns of biodiversity, yet its spatial heterogeneity is often overlooked in biodiversity analyses and management strategies. Properly parameterised heterogeneous dispersal networks capture the complex interplay between landscape structure and species-specific dispersal capacities. However, this heterogeneity is recurrently neglected when studying the processes underlying biodiversity variation. To address this gap, we introduce a conceptual framework detailing the fundamental processes driving dispersal heterogeneity and its effects on biodiversity dynamics. We propose methods to parameterise heterogeneous dispersal networks, facilitating their integration into commonly used quantitative frameworks for biodiversity analyses. By considering the architecture of heterogeneous dispersal networks, we demonstrate their critical role in guiding biodiversity management strategies.

Tracking the source of antibiotic resistome in the stormwater network drainage in the presence of sewage illicit connections.

Stormwater pipes are illicitly connected with sewage in many countries, which means that sewage enters stormwater pipes and the drainage is discharged to surface water without any treatment. Sewage contains more pathogens and highly risky antibiotic resistance genes (ARGs) than surface runoff. Therefore, sewage may alter the microbial and ARG compositions in stormwater pipe drainage, which in turn leads to an increased risk of resistance in surface water. However, the effects of sewage on ARGs in the drainage of stormwater networks have not been systematically studied. This study characterized the microbial and ARG composition of several environmental compartments of a typical stormwater network and quantified their contributions to those in the drainage. This network transported ARGs and microorganisms from sewage, sediments in stormwater pipes, and surface runoff into the drainage and thus into the river. According to metagenomic analysis, multidrug resistance genes were most abundant in all samples and the numbers and relative abundance of ARGs in the drainage collected during wet weather were comparable to that of sewage. The results of SourceTracker showed that the relative contribution of sewage was double that of rainwater and surface runoff in the drainage during wet weather for both microorganisms and ARGs. Desulfovibrio, Azoarcus, and Sulfuritalea were connected with the greatest number of ARGs and were most abundant in the sediments of stormwater pipes. Furthermore, stochastic processes were found to dominate ARG and microbial assembly, as the effects of high hydrodynamic intensity outweighed the effects of environmental filtration and species interactions. The findings of this study can increase our understanding of ARGs in stormwater pipe drainage, a crucial medium linking ARGs in sewage to environmental ARGs.

Spatial ecology of the Giant Sea Bass, Stereolepis gigas, in a southern California kelp forest as determined by acoustic telemetry.

The fisheries history of the Giant Sea Bass, Stereolepis gigas (Telostei: Polyprionidae), is closely linked to its spatial ecology. Its overharvest is directly associated with formation of spatially distinct spawning aggregations during summer, while its subsequent recovery is hypothesized to be the result of spatially explicit gear restrictions. Understanding the spatial ecology of Giant Sea Bass is a key part of efforts to assess contemporary threats such as commercial harvest and incidental catch by recreational fisheries. In this study, we used acoustic telemetry to characterize Giant Sea Bass space use in the La Jolla kelp forest using an acoustic array that encompasses two marine protected areas (MPAs) and heavily trafficked recreational fishing grounds. Five of the seven fish we tagged remained in the La Jolla array for at least 6 months. Two fish were resident across multiple years, with one fish consistently detected for 4 years. Only one fish was detected in the broader network of regional acoustic receivers, moving north approximately 8 km to Del Mar. Most tagged fish had home ranges and core use areas indicating they spend considerable time outside MPAs, particularly in areas with high recreational fishing activity. During spawning season we detected fish less frequently in the La Jolla array and recorded higher movement rates. While the current MPA network in La Jolla by no means offers complete protection to this fish, it does appear to support long-term persistence of some individuals in a region of exceptionally high recreational fishing pressure.

Evaluating Different Factors That Affect the Nesting Patterns of European and Algerian Hedgehogs in Urban and Suburban Environments.

Small undisturbed patches in urban environments serve as important refuges for wildlife, e.g., hedgehogs. However, the effects of urbanization on certain biological aspects, like nesting behavior, remain unknown. We captured and tracked the movement of 30 male hedgehogs of two co-existing species: Algerian and European hedgehogs. The study was carried out in Valencia (eastern Spain). We distinguished six macrohabitats and five subhabitats. We analyzed the proportions of the macro and subhabitats where nests were found to calculate a resource selection function and fit GLMs. Hedgehog nests tended to concentrate in areas with little human disturbance and were built in hedges or under bush-like vegetation. We did not find any significant differences between species or other considered groups. We noted that nests were distributed around hard-to-find suitable habitat patches. We even recorded a case of two males from both species simultaneously using one nest. Our results suggest that hedgehog conservation in urban environments can be improved by the correct management of forest patches by conserving bush-like vegetation and improving the connectivity between suitable patches with ecological corridors.

Finding analytical approximations for discrete, stochastic, individual-based models of ecology.

Discrete time, spatially extended models play an important role in ecology, modelling population dynamics of species ranging from micro-organisms to birds. An important question is how 'bottom up', individual-based models can be approximated by 'top down' models of dynamics. Here, we study a class of spatially explicit individual-based models with contest competition: where species compete for space in local cells and then disperse to nearby cells. We start by describing simulations of the model, which exhibit large-scale discrete oscillations and characterize these oscillations by measuring spatial correlations. We then develop two new approximate descriptions of the resulting spatial population dynamics. The first is based on local interactions of the individuals and allows us to give a difference equation approximation of the system over small dispersal distances. The second approximates the long-range interactions of the individual-based model. These approximations capture demographic stochasticity from the individual-based model and show that dispersal stabilizes population dynamics. We calculate extinction probability for the individual-based model and show convergence between the local approximation and the non-spatial global approximation of the individual-based model as dispersal distance and population size simultaneously tend to infinity. Our results provide new approximate analytical descriptions of a complex bottom-up model and deepen understanding of spatial population dynamics.

The essential habitat role of a unique coastal inlet for a widely distributed apex predator.

Essential habitats support specific functions for species, such as reproduction, feeding or refuge. For highly mobile aquatic species, identifying essential habitats within the wider distribution range is central to understanding species ecology, and underpinning effective management plans. This study examined the movement and space use patterns of sevengill sharks (Notorynchus cepedianus) in Caleta Valdés (CV), a unique coastal habitat in northern Patagonia, Argentina. Seasonal residency patterns of sharks were evident, with higher detectability in late spring and early summer and lower during autumn and winter. The overlap between the residency patterns of sharks and their prey, elephant seals, suggests that CV functions as a seasonal feeding aggregation site for N. cepedianus. The study also found sexual differences in movement behaviour, with males performing abrupt departures from CV and showing increased roaming with the presence of more sharks, and maximum detection probability at high tide. These movements could be related to different feeding strategies between sexes or mate-searching behaviour, suggesting that CV may also be essential for reproduction. Overall, this study highlights the importance of coastal sites as essential habitats for N. cepedianus and deepens our understanding of the ecological role of this apex predator in marine ecosystems.

Diet and Spatial Ecology Influence Red-Legged Partridge Exposure to Pesticides Used as Seed Treatment.

Seed treatment with pesticides is an extended agricultural practice with a high risk to granivorous birds that consume those seeds. To characterize that risk, it is necessary to understand the ecological factors that determine the exposure chances of birds to treated seeds. We investigated how pesticide uptake by red-legged partridges was related to cultivated plant ingestion and to the use of recently sown fields. We analyzed pesticide residues in 144 fecal samples from 32 flocks and determined the plant diet composition using DNA metabarcoding. Habitat use was studied through the monitoring of 15 GPS-tagged partridges. We confirmed, through the analysis of seeds, that >80% of cereal fields from the area had seeds treated with triazole fungicides. Tebuconazole was detected in 16.6% of partridges' feces. During the sowing season, cultivated plants accounted for half of the plant diet, but no association was found between cultivated plant consumption and pesticide intake. GPS tracking revealed that tebuconazole was detected in feces when partridges had recently used sown fields, whereas nonexposed partridges showed no overlap with recently sown areas. Our results highlight the need to incorporate field ecology into the characterization of pesticide exposure to improve the efficacy of environmental risk assessment.

Don't mind if I do: Arctic humpback whales respond to winter foraging opportunities before migration.

Migration patterns are fundamentally linked to the spatio-temporal distributions of prey. How migrating animals can respond to changes in their prey's distribution and abundance remains largely unclear. During the last decade, humpback whales (Megaptera novaeangliae) used specific winter foraging sites in fjords of northern Norway, outside of their main summer foraging season, to feed on herring that started overwintering in the area. We used photographic matching to show that whales sighted during summer in the Barents Sea foraged in northern Norway from late October to February, staying up to three months and showing high inter-annual return rates (up to 82%). The number of identified whales in northern Norway totalled 866 individuals by 2019. Genetic sexing and hormone profiling in both areas demonstrate a female bias in northern Norway and suggest higher proportions of pregnancy in northern Norway. This may indicate that the fjord-based winter feeding is important for pregnant females before migration. Our results suggest that humpback whales can respond to foraging opportunities along their migration pathways, in some cases by continuing their feeding season well into winter. This provides an important reminder to implement dynamic ecosystem management that can account for changes in the spatio-temporal distribution of migrating marine mammals.

Block-pulse integrodifference equations.

We present a hybrid method for calculating the equilibrium population-distributions of integrodifference equations (IDEs) with strictly increasing growth, for populations that are confined to a finite habitat-patch. This method is based on approximating the growth function of the IDE with a piecewise-constant function, and we call the resulting model a block-pulse IDE. We explicitly write out analytic expressions for the iterates and equilibria of the block-pulse IDEs as sums of cumulative distribution functions. We characterize the dynamics of one-, two-, and three-step block-pulse IDEs, including formal stability analyses, and we explore the bifurcation structure of these models. These simple models display rich dynamics, with numerous fold bifurcations. We then use three-, five-, and ten-step block-pulse IDEs, with a numerical root finder, to approximate models with compensatory Beverton-Holt growth and depensatory, or Allee-effect, growth. Our method provides a good approximation for the equilibrium distributions for compensatory and depensatory growth and offers numerical and analytical advantages over the original growth models.

The role of seasonal migration in spatial population synchrony.

Spatially synchronized population dynamics are common in nature, and understanding their causes is key for predicting species persistence. A main driver of synchrony between populations of the same species is shared environmental conditions, which cause populations closer together in space to be more synchronized than populations further from one another. Most theoretical and empirical understanding of this driver considers resident species. For migratory species, however, the degree of spatial autocorrelation in the environment may change across seasons and vary by their geographic location along the migratory route or on a nonbreeding ground, complicating the synchronizing effect of the environment. Migratory species show a variety of different strategies in how they disperse to and aggregate on nonbreeding grounds, ranging from completely shared nonbreeding grounds to multiple different ones. Depending on the sensitivity to environmental conditions off the breeding grounds, we can expect that migration and overwintering strategies will impact the extent and spatial pattern of population synchrony on the breeding grounds. Here, we use spatial population-dynamic modeling and simulations to investigate the relationship between seasonal environmental autocorrelation and migration characteristics. Our model shows that the effects of environmental autocorrelation experienced off the breeding ground on population synchrony depend on the number and size of nonbreeding grounds, and how populations migrate in relation to neighboring populations. When populations migrated to multiple nonbreeding grounds, spatial population synchrony increased with increasing environmental autocorrelation between nonbreeding grounds. Populations that migrated to the same place as near neighbors had higher synchrony at short distances than populations that migrated randomly. However, synchrony declined less across increasing distances for the random migration strategy. The differences in synchrony between migration strategies were most pronounced when the environmental autocorrelation between nonbreeding grounds was low. These results show the importance of considering migration when studying spatial population synchrony and predicting patterns of synchrony and population viability under global environmental change. Climate change and habitat loss and fragmentation may cause range shifts and changes in migratory strategies, as well as changes in the mean and spatial autocorrelation of the environment, which can alter the scale and patterns observed in spatial population synchrony.

Catchment-based sampling of river eDNA integrates terrestrial and aquatic biodiversity of alpine landscapes.

Monitoring of terrestrial and aquatic species assemblages at large spatial scales based on environmental DNA (eDNA) has the potential to enable evidence-based environmental policymaking. The spatial coverage of eDNA-based studies varies substantially, and the ability of eDNA metabarcoding to capture regional biodiversity remains to be assessed; thus, questions about best practices in the sampling design of entire landscapes remain open. We tested the extent to which eDNA sampling can capture the diversity of a region with highly heterogeneous habitat patches across a wide elevation gradient for five days through multiple hydrological catchments of the Swiss Alps. Using peristaltic pumps, we filtered 60 L of water at five sites per catchment for a total volume of 1800 L. Using an eDNA metabarcoding approach focusing on vertebrates and plants, we detected 86 vertebrate taxa spanning 41 families and 263 plant taxa spanning 79 families across ten catchments. For mammals, fishes, amphibians and plants, the detected taxa covered some of the most common species in the region according to long-term records while including a few more rare taxa. We found marked turnover among samples from distinct elevational classes indicating that the biological signal in alpine rivers remains relatively localised and is not aggregated downstream. Accordingly, species compositions differed between catchments and correlated with catchment-level forest and grassland cover. Biomonitoring schemes based on capturing eDNA across rivers within biologically integrated catchments may pave the way toward a spatially comprehensive estimation of biodiversity.

Spatial metapopulation dynamics with local and global colonization.

We revisit a spatial metapopulation model on continuous space as a stochastic point pattern dynamics. In the model, local patches as points are distributed with a certain spatial configuration and status of each patch changes stochastically between occupied and empty: an occupied patch becomes empty by local extinction and an empty patch becomes occupied both by local and global colonization. We carry out simulation analysis and derive an analytical model in terms of singlet, pair and triplet probabilities that describe the stochastic dynamics. Using a simple closure that approximates triplet probabilities by singlet and pair probabilities, we show that equilibrium singlet and pair probabilities can be analytically derived. The derived equilibrium properties successfully describe simulation results under a certain condition where the range of local colonization and the proportion of global colonization play key roles. Our model is an extension of the classical non-spatial Levins model to a spatially explicit metapopulation model. We appeal the advantage of point pattern approach to study spatial dynamics in general ecology and call for the need to deepen our understanding of mathematical tools to explore point pattern dynamics.