Patterns Of Foraging Activity Research Articles

Divergent Circadian Foraging Strategies in Response to Diurnal Predation Versus Persistent Rain in Asian Weaver Ant, Oecophylla smaragdina, Suggest Possible Energetic Trade-offs.

The study of chronobiology of foraging behavior in social insects offers valuable models for the investigation of circadian rhythms. We scored hourly nest entries and exits of Oecophylla smaragdina (Asian weaver ant) workers in 9 active non-polydomous nests on days with and without rain and with and without a primarily diurnal predator present. After determining that Oecophylla display a high nest fidelity, we focused exclusively on analyzing nest entry counts: we found a significant decrease in overall entry counts of individual ants on rainy days compared with non-rainy days (p < 0.0001). They usually maintain a typical diurnal pattern of foraging activity; however, that regularity was often distorted during rainy periods but appeared to quickly revert to typical patterns following rain. This lack of compensatory foraging activity following a period of rain supports the hypothesis that these ants have enough food reserves to withstand a pure masking-induced suppression of foraging activity. Predation through bird anting, too, decreased foraging activity but appeared to cause a reversal in foraging activity timing from diurnal to nocturnal foraging. Daily periodicity of foraging was significantly disrupted in most nests during rain; however, daily foraging periodicity was disrupted in only one nest due to presence of predators. Thus, rain and predation both exert significant impacts on the overall foraging activity of Asian weaver ants, but while persistent pressure from rain seemed to primarily cause masking (diminution) of circadian foraging activity, predation restricted to the daytime resulted in phase-inversion to nocturnal foraging activity, with little diminution. This is consistent with different energetic strategies being used in response to different pressures by this species.

Urban abiotic stressors drive changes in the foraging activity and colony growth of the black garden ant Lasius niger

Changes in habitat characteristics are known to have profound effects on biotic communities and their functional traits. In the context of an urban-rural gradient, urbanisation drastically alters abiotic characteristics, e.g., by increasing environmental temperatures and through light pollution. These abiotic changes significantly impact the functional traits of organisms, particularly insects. Furthermore, changes in habitat characteristics also drive changes in the behavioural traits of animals, allowing them to adapt and thrive in new environments. In our study, we focused on the synanthropic ant species Lasius niger as a model organism. We conducted nocturnal field observations and complemented them with laboratory experiments to investigate the influence of night warming (NW) associated with Urban Heat Islands (UHI), light pollution (ALAN), and habitat type on ant foraging behaviour. In addition, we investigated the influence of elevated temperatures on brood development and worker mortality. Our findings revealed that urban populations of L. niger were generally more active during the night compared to their rural counterparts, although the magnitude of this difference varied with specific city characteristics. In laboratory settings, higher temperatures and continuous illumination were associated with increased activity level in ants, again differing between urban and rural populations. Rural ants exhibited more locomotion compared to their urban counterparts when maintained under identical conditions, which might enable them to forage more effectively in a potentially more challenging environment. High temperatures decreased the developmental time of brood from both habitat types and increased worker mortality, although rural colonies were more strongly affected. Overall, our study provides novel insights into the influence of urban environmental stressors on the foraging activity pattern and colony development of ants. Such stressors can be important for the establishment and spread of synanthropic ant species, including invasive ones, and the biotic homogenization of anthropogenic ecosystems.

Seasonal pattern of foraging activity in Apis cerana (indica) for assessment of colony performance index and weight gain in Kashmir valley

Seasonal pattern of foraging activity in Apis cerana (indica) for assessment of colony performance index and weight gain in Kashmir valley

Reef effect of offshore structures on the occurrence and foraging activity of harbour porpoises

With increasing numbers of offshore structures being installed and decommissioned, a better understanding of their effect on marine predators is timely. There is some evidence that oil and gas platforms may attract marine mammals, acting as artificial reefs. However, it is unclear whether different man-made structure designs have similar effects or whether artificial structures modify the diel patterns of occurrence and foraging of marine mammals. Here, we used passive acoustics to investigate the occurrence and foraging activity of harbour porpoises (Phocoena phocoena) around four artificial structures of different age and complexity. We deployed an array of echolocation click detectors (CPODs) in 2021, along a gradient of distances to these structures and assessed the extent to which porpoises were attracted to them and their effect on porpoises’ diel patterns of occurrence and foraging activity. The probability of porpoise occurrence and foraging activity decreased with distance from offshore structures. A significant increase in porpoise occurrence and foraging was detected during night-time compared to daytime around all four offshore structures (&amp;lt; 200 m). Comparing pre- and post-installation porpoise detections, the daily patterns of occurrence and foraging activity shifted from a weak diel pattern before the structure was installed, to a strong nocturnal pattern when the structure was present. These findings provide evidence that marine mammals are attracted to man-made structures and that porpoises modify their diel patterns of occurrence and foraging activity around them. This research suggests that offshore structures play an important role as foraging areas for some marine mammals and provides key information for decommissioning considerations and the planning of decommissioning activities.

Foraging Activity of Tetragonula laeviceps Workers for Natural Resources and Nest Materials at a Polyculture Cropland in Batusangkar, Tanah Datar Regency, West Sumatra

Tetragonula laeviceps is the most widely distributed stingless bee species in tropics. The observation on foraging activity of stingless bee workers of Tetragonula laeviceps had been done from September to December 2020 at a polyculture cropland in Batusangkar, Tanah Datar Regency, West Sumatra. The study aimed to determine the pattern of foraging activity of this species at a cropland with vegetation mixture and to identify the relationships between environmental factors i.e., temperature, humidity, and light intensity, to the foraging activity. The stingless bee workers were observed hourly for fifteen minutes from 6 am to 6 pm at their nest entrance, focusing on what they carry when leaving and entering colony. The observation was conducted on seven consecutive days within a week. The natural resources that stingless bee workers brought into their nest were identified and noted along with the number of workers carried certain natural resource. The analysis on observation data showed that the going-out of stingless bee workers for resources peaked at 9 am. The resource carried the most by stingless bee workers into their nest were pollen. Statistical analysis revealed that 78% of foraging activities for resources and nest material in Tetragonula laeviceps were influenced by environmental factors.

Seasonal Phenology and Colony Longevity Patterns in a Predatory Social Wasp

Social wasps play critical ecological roles in an ecosystem, providing a diversity of services and some disservices. Yellowjacket wasps (Vespula spp.) in particular are well known for shaping arthropod communities via predation and competition for resources. In part due to their sociality and large colony sizes, Vespula can have a profound ecological impact on local communities. Such effects can be magnified when colonies exhibit a perennial life history, in which a colony will overwinter, persist for more than one year, and become orders of magnitude larger in size compared to typical annual colonies. Despite growing interest in the factors that influence colony success, we currently lack the ability to predict when yellowjackets may have a high-abundance or outbreak year. This highlights the need for a critical understanding of the phenological patterns of foraging activity, colony distribution, and senescence. Here, we quantify the seasonal activity and foraging rates of 123 colonies of the western yellowjacket, V. pensylvanica, in its native range over 4 consecutive years. Average colony longevity was about 1 month longer than previously reported for this species, and colonies with later peaks in activity and higher average traffic rates persisted longer into the winter. Longer-lived colonies tended to cluster together within a year, but not between years. We found 3 perennial colonies (2.4% of all colonies), and they exhibited tenfold higher peak traffic rates compared to annual colonies. By identifying temporal and spatial patterns in survivorship and colony longevity, we gain insight into the factors associated with prolonged survival time and increased likelihood of overwintering in yellowjacket wasps.

Behavioural biology and ecology of Blackbuck (Antelope cervicapra) : A Review

Blackbuck (Antelope cervicapra) belongs to the Bovidae family and subfamily Antilopinae and is distributed in India, Pakistan, Nepal and United Arab Emirates. It can live in wide variety of habitats, including grassland, bush land, scrubland and dry thorn habitats. Sexual dimorphism is very clear. Males have blackish or dark brown coat colours, while females have a yellowish coat colour. Males have a shorter lifespan as compared to females. Generally, blackbuck forage in the daytime but sometimes also forage at night and their foraging activity patterns are influenced by environmental factors and seasonal variations. They use to prefer wide variety of food materials, like leaves, grasses, cereals, vegetables, shrubs and varieties of crops. During the breeding season, males become territorial. Herd size is also affected by the seasons and environmental factors. Exotic species invasion, habitat loss, anthropogenic activity, overgrazing and agricultural expansion, urbanization and industrialization all reduce suitable habitat for blackbuck survival. Illegal hunting, poaching, habitat loss, road killing and lack of genetic variation are also responsible for the dramatic decline in the population of blackbuck. Study of genetic variation, captive breeding and the establishment of a blackbuck rescue centre might be helpful for enhancing the population and survival of blackbucks.

Actividad forrajera y de mantenimiento de nido de la hormiga cortadora de hojas Acromyrmex lobicornis (Hymenoptera: Formicidae) en un viñedo orgánico inserto en el desierto de Monte en Mendoza, Argentina

Exclusive to the Neotropical region, leaf cutter ants are considered agricultural pests, although they can also have a positive effect on plants. In Mendoza vineyards, vegetal biodiversity is minimal, therefore they cut off this plant as a feeding resource. Acromyrmex lobicornis Emery it is mostly nocturnal and forages according to temperature. Foraging and maintenance activity was measured monthly in 14 colonies, throughout a full day at fourhour intervals, from October 2019 to March 2020. Acromyrmex lobicornis showed different seasonal patterns of foraging activity. Foraging intensity was highest throughout January and February, intermediate during December, and lowest in October and November. A bimodal feeding pattern was observed, foraging both day and night avoiding the hottest hours. The maximum collection of fragments was observed in the range of 10 and 19 °C, less between 20 - 39 °C, minimum between 40 - 49 °C and null between 0 - 9 °C. The nest - maintenance activity was maximum between 20 - 29 °C, less between 30 - 39 ° C, minimum between 40 - 49 °C and null between 0 - 20 °C.

Ant Communities and Ecosystem Services in Organic Versus Conventional Agriculture in the U.S. Corn Belt.

Reducing the use of synthetic fertilizers and pesticides can limit negative impacts of agriculture on insects and is a crucial step towards sustainable agriculture. In the United States, organic agriculture has the potential to reduce greenhouse gas emissions, pollutant runoff, and biodiversity loss in the Midwestern Corn Belt-an area extending over 500,000 km2 devoted to intensive production of corn Zea mays (Linnaeus 1753) (Poales: Poaceae), often in rotation with soy Glycine max (Linnaeus 1753) (Fabales: Fabaceae) or wheat Triticum aestivum (Linnaeus 1753) (Poales: Poaceae). Working in 30-yr-long landscape experiments in this region, we tested for impacts of conventional versus organic agriculture on ant communities (Hymenoptera: Formicidae) and potential ecosystem services they provide. Organic fields supported higher ant diversity and a slightly more species-rich ant assemblage than conventionally managed fields but did not otherwise differ in community composition. Despite similar community composition, organic and conventional fields differed in seasonal patterns of ant foraging activity and potential for natural pest suppression. Conventional plots experienced higher overall ant foraging activity, but with the timing skewed towards late in the growing season such that 75% of ant foraging occurred after crop harvest in a wheat year and was therefore unavailable for pest suppression. Organic fields, in contrast, experienced moderate levels of ant foraging activity throughout the growing season, with most foraging occurring during crop growth. Organic fields thus supported twice as much pest suppression potential as conventional fields. Our results highlight the importance of timing in mediating ecosystem services in croplands and emphasize the value of managing landscapes for multiple services rather than yield alone.

Neighbor Colonies Affect Level of Foraging in the Generalist Ant Pheidole oxyops (Hymenoptera: Formicidae)

Pheidole oxyops Forel (Hymenoptera: Formicidae) is a generalist ant that can display passive and active forage strategies. Moreover, this species can forage actively for a variety of food resources from plant debris to dead arthropods. In this study we tested whether abiotic factors (e.g., daytime, temperature, and residual soil accumulation), as well as neighboring colonies, could influence the patterns of foraging activity. Forage activity was observed in morning, afternoon, and evening sessions. Also, we recorded the frequency of individuals, direction, and ground surface temperature during foraging events. Our results showed that forage direction was not affected by residual soil accumulation and neighboring P. oxyops colonies. The rate of departures from nests decreased as ground surface temperature increased, but nest exits increased significantly with the occurrence of nearby P. oxyops colonies. We observed also that P. oxyops appeared to maximize forage activities during the evening period that would reduce their exposure to potential predators, competitors, and any adverse effects associated with direct solar radiation.

Timing and Duration of Observation Periods of Foraging Behavior in Natural Grasslands

The goals of this study were to evaluate the length of time grazing which should be monitored over a 24-h period to predict the grazing behavior of beef heifers within a season and determine the patterns of foraging activity over 24 h. A database was constructed between 2010 and 2012 for beef heifers managed under rotational grazing in a natural grassland. Grazing, rumination, and other activity times were assessed visually during 24 h on 15 occasions. Data were classified according to climatic seasons, generating 12 replicates in summer, 18 in spring, 24 in autumn, and 36 in winter. Treatments were the evaluation of four distinct periods: from sunrise to sunset (DAY-SUN), daylight duration from dawn to nightfall (DAYLIGHT), DAYLIGHT plus 2 h (DAYLIGHT+2), DAYLIGHT to midnight (DAYLIGHT to 0), and the entire 24 h period (CONTROL). Differences for grazing, rumination, and other activities were found in all seasons for the evaluation periods. Sampling sufficiency was reached only with the DAYLIGHT to 0 and CONTROL for all four climatic seasons. The DAYLIGHT to 0 treatment covered 75% of a 24-h period and 95% of the mean foraging time took place during this time interval. Considering grazing distribution during a day, in the warm seasons, the major grazing period during mornings occurred earlier than in the cool seasons, and in cool seasons, the grazing peak was observed during the afternoon. Visual observations from dawn until midnight represented the total grazing time and natural behavior of heifers and could be used to represent grazing activities for the entire day.

Patterns of foraging activity and fidelity in a southeast Asian flying fox

BackgroundImproved understanding of the foraging ecology of bats in the face of ongoing habitat loss and modification worldwide is essential to their conservation and maintaining the substantial ecosystem services they provide. It is also fundamental to assessing potential transmission risks of zoonotic pathogens in human-wildlife interfaces. We evaluated the influence of environmental and behavioral variables on the foraging patterns of Pteropus lylei (a reservoir of Nipah virus) in a heterogeneous landscape in Cambodia.MethodsWe employed an approach based on animal-movement modeling, which comprised a path-segmentation method (hidden Markov model) to identify individual foraging-behavior sequences in GPS data generated by eight P. lylei. We characterized foraging localities, foraging activity, and probability of returning to a given foraging locality over consecutive nights. Generalized linear mixed models were also applied to assess the influence of several variables including proxies for energetic costs and quality of foraging areas.ResultsBats performed few foraging bouts (area-restricted searches) during a given night, mainly in residential areas, and the duration of these decreased during the night. The probability of a bat revisiting a given foraging area within 48 h varied according to the duration previously spent there, its distance to the roost site, and the corresponding habitat type. We interpret these fine-scale patterns in relation to global habitat quality (including food-resource quality and predictability), habitat-familiarity and experience of each individual.ConclusionsOur study provides evidence that heterogeneous human-made environments may promote complex patterns of foraging-behavior and short-term re-visitation in fruit bat species that occur in such landscapes. This highlights the need for similarly detailed studies to understand the processes that maintain biodiversity in these environments and assess the potential for pathogen transmission in human-wildlife interfaces.

Evidence of resource-defense polygyny in an endangered subtropical bat, Eumops floridanus

Understanding sociality and animal behavior is critical for developing effective conservation strategies. Many tropical bat species form harems, where dominant males play key social roles by defending groups of females directly (female-defense polygyny) and/or the resources that females need (resource-defense polygyny). The Florida bonneted bat (Eumops floridanus) is an endangered subtropical species suspected to form harems, but our understanding of its social structure, reproduction, and behavior is rudimentary. In this study, we evaluated demographic variation in morphological and behavioral characteristics of Florida bonneted bats to test the hypothesis that this species forms harem groups and exhibits resource-defense polygyny at roost sites. We used a 4-year dataset of 341 individuals uniquely marked with passive integrated transponders (PIT tags), coupled with tri-annual capture records, to track activity patterns of bats at five roosts fitted with PIT tag readers. We identified the likely dominant males and other demographic groups in each roost using morphometric characteristics and reproductive status. We assessed differences between sexes and among status categories in three primary metrics: amount of activity at the roost, time of emergence, and initial foray duration per night. Dominant males consistently were the most active individuals at roosts and spent the least amount of time away from roosts during forays, relative to females and other males. Females spent more time away from roosts than males and shared similar foraging activity patterns regardless of status. Our findings suggest that Florida bonneted bats form small harem groups that are active year round. Male bats exhibit characteristics of resource-defense polygyny at roost sites and a size-biased hierarchy, with the largest reproductively active males appearing to defend the roost at the expense of time spent foraging. We suggest that the roost site represents a critical, limited and defendable resource for male Florida bonneted bats to gain access to females, which has important implications for the conservation and enhancement of roost sites. Our study highlights the importance of accounting for differences in behavior across demographic groups and social roles when considering resource needs for imperiled species.

Don’t stay out too long! Thermal tolerance of the stingless bees Melipona subnitida decreases with increasing exposure time to elevated temperatures

In hot climates, foraging bees risk overheating during food collection. Here, we investigated influence of ambient temperature on the foraging activity in Melipona subnitida, a stingless bee species that naturally occurs in the hottest and driest regions of the Brazilian tropical dry forest. We observed a decrease in round-trip duration of pollen foragers with increasing ambient temperature. Foraging duration of nectar collectors, however, was not affected by the thermal conditions of the environment. When exposed to heat stress in incubators, the upper thermal limits of foragers decreased with increasing exposure time. Moreover, individuals with previous access to liquid food showed higher critical and lethal temperatures than unfed bees. Our study revealed plastic changes in the upper thermal limit of M. subnitida, which help to understand the foraging activity pattern of this stingless bee species.

Foraging activity patterns of critically endangered white-bellied herons (Ardea insignis) in Punatsangchu and Mangdechu river basins of Bhutan ―Conservationists for White-bellied Herons―

Foraging activity patterns of critically endangered white-bellied herons (Ardea insignis) in Punatsangchu and Mangdechu river basins of Bhutan ―Conservationists for White-bellied Herons―

Body Size and Its Relation to the Foraging Schedules of Social Wasps.

The foraging behavior of social wasps may vary throughout the day in response to different factors, including environmental variation and interspecific interactions. Body size is known to play a critical role in determining daily activity patterns in ecological communities. However, the body size characteristics of most species of social wasps is poorly known, as well the relationship between body size and foraging schedules. In this study, we evaluated the extent to which body size determines the patterns of daily activity in a community of social wasps. We found a high temporal overlap in most of the species pairs tested. The pattern of daily foraging activity fits a bimodal distribution for the majority of the species. Moreover, we found a relationship between body and foraging time; in general, smaller species tended to begin foraging in the early hours of the morning, in contrast to larger wasps, which began foraging later. These results suggest that patterns of foraging activity in social wasps are likely to be the result of complex interactions between many different factors, including body size.

Does social thermal regulation constrain individual thermal tolerance in an ant species?

In ants, social thermal regulation is the collective maintenance of a nest temperature that is optimal for individual colony members. In the thermophilic ant Aphaenogaster iberica, two key behaviours regulate nest temperature: seasonal nest relocation and variable nest depth. Outside the nest, foragers must adapt their activity to avoid temperatures that exceed their thermal limits. It has been suggested that social thermal regulation constrains physiological and morphological thermal adaptations at the individual level. We tested this hypothesis by examining the foraging rhythms of six populations of A. iberica, which were found at different elevations (from 100 to 2,000m) in the Sierra Nevada mountain range of southern Spain. We tested the thermal resistance of individuals from these populations under controlled conditions. Janzen's climatic variability hypothesis (CVH) states that greater climatic variability should select for organisms with broader temperature tolerances. We found that the A. iberica population at 1,300m experienced the most extreme temperatures and that ants from this population had the highest heat tolerance (LT50=57.55°C). These results support CVH's validity at microclimatic scales, such as the one represented by the elevational gradient in this study. Aphaenogaster iberica maintains colony food intake levels across different elevations and mean daily temperatures by shifting its rhythm of activity. This efficient colony-level thermal regulation and the significant differences in individual heat tolerance that we observed among the populations suggest that behaviourally controlled thermal regulation does not constrain individual physiological adaptations for coping with extreme temperatures.

Thermal tolerance varies with dim‐light foraging and elevation in large carpenter bees (Hymenoptera: Apidae: Xylocopini)

1. Thermal tolerance has a strong predictive power for understanding the ecology and distribution of organisms, as well as their responses to changes in land use and global warming. However, relatively few studies have assessed thermal tolerances for bees.2. The present study aimed to determine whether the critical thermal maximum (CTmax) of carpenter bees (Apidae: genus Xylocopa Latreille) varies with different patterns of foraging activity and elevation. In addition, the influence of body size, body water content and relative age was examined with respect to their CTmax and differences in thoracic temperature (Tth) among species were evaluated.3. The CTmax of one crepuscular (Xylocopa olivieri) and two diurnal species (Xylocopa violacea and Xylocopa iris) of carpenter bees was assessed at sea level on the Greek island of Lesvos. To detect variation as a result of elevation, the CTmax of a population of X. violacea at 625 m.a.s l. was assessed and compared with that from sea level.4. Xylocopa olivieri displayed a similar CTmax to that of X. violacea but lower than that of X. iris. Body size, body water content, and relative age did not affect CTmax. In X. violacea, CTmax decreased with elevation and all three species have high Tth independent of ambient temperatures.5. The results of the present study are consistent with variations in CTmax predicted by broad spatial and temporal patterns reported for other insects, including honey and bumble bees. The implications of the results are discussed aiming to understand the differences in the foraging pattern of these bees.

Secondary types of food in the diet of a small-sized mammalian herbivore: from species ecology to habitat inferences

A macroscopic analysis of stomach contents is undertaken in the common vole, Microtus arvalis, obscurus chromosomal form in order to assess the patterns of non-herbivorous foraging activity of a herbivore distributed in different Eurasian biomes. The samples are collected in the Ural Mountains at midsummer (8 localities, 60° 38′ N to 51° 07′ N), and in winter (one locality, 56° 37′ N). Based on the frequency of occurrence and quantitative abundance in stomachs, photoassimilating parts of vascular plants are considered as the primary food category and inflorescences, roots, seeds, mycorrhizal fungi, arthropods, chitin fragments, colorless amorphous matter, and mosses are recognized as the secondary food categories. Although no alternatives to the primary food are found, some of the supplements substitute one another either geographically (fungi in the north versus mature seeds in the south) or by the vertical layers of habitats (seeds and arthropods in the herbaceous layer versus roots and fungi in the shallow layers of soil). Non-staple food consumption in different latitudinal zones at midsummer reflects spatiotemporal dynamics of food availability. From taiga to steppes of the Urals, the common vole is sporadically included in soil foraging but the consumption of mycorrhizal fungi (Glomerales) is only revealed within the zones of dark coniferous forests. The results suggest that the extents of inclusion of a herbivore in soil food webs might vary among ecosystems. In a paleoecological perspective, the analysis of stomach contents in a herbivorous micromammal might reveal fine-scale details of the habitats and provides insights into the stages of the growing season.

THE BEHAVIOR OF YOUNG SALMON (SALMO SALAR L.) IN THE RIVER LIZHMA (LAKE ONEGO) IN THE SUMMER SEASON

It was found that each age group of young salmon has its own daily foraging activity pattern, mainly regulated by illumination. At the latitude of the Lizhma River (62°22ʹ40ʺ N, 34°29ʹ57ʺ E), which drains to Lake Onego, the foraging activity duration in June and July is 16 h ± 30 min. In twilight hours and in the dark (below 200 lx) it was absent in all age groups. Young-of-the-year (0+) maintained a constant level of foraging activity during the day, making about 900 darts at food. Parr aged 1+ and 2+ foraged the most actively in the morning (18,000 lx), after which the activity declined until 15 h (70,000 lx)and remained low until 19 h (21,000 lx). Then it increased slightly until dusk. The number of darts at food was about 350 per day. For 3+ parr, the activity was the lowest in the morning hours, gradually growing afterwards to reach a peak at around 15 hours, after which a gradual decline was observed. During the day they make about 300 darts at food. The foraging intensity for 0+ fish averages 63 darts per hour. Fish aged 1+ make less than 30 darts/h; 2+ make about 40 darts/h; 3+ salmon make about 25 darts/h. Foraging activity can be reduced by dense clouds and competition. Young fish aged 0+ and 1+ cannot compete with larger parr and other predators for fodder-rich sites. 2+ and 3+ parr occupy the most advantageous feeding microsites, where they perform up to 88 % of all daily darts for food in less than 50 % active time, and thus satisfy their daily nutritive demand. As a result, 2+ and 3+ parr would be more often seen inactive in “shelter” microsites compared to 0+ and 1+ individuals.