Vervet monkeys use paths consistent with context‐specific spatial movement heuristics

Animal foraging routes are analogous to the computationally demanding “traveling salesman problem” (TSP), where individuals must find the shortest path among several locations before returning to the start. Humans approximate solutions to TSPs using simple heuristics or “rules of thumb,” but our knowledge of how other animals solve multidestination routing problems is incomplete. Most nonhuman primate species have shown limited ability to route plan. However, captive vervets were shown to solve a TSP for six sites. These results were consistent with either planning three steps ahead or a risk‐avoidance strategy. I investigated how wild vervet monkeys (Chlorocebus pygerythrus) solved a path problem with six, equally rewarding food sites; where site arrangement allowed assessment of whether vervets found the shortest route and/or used paths consistent with one of three simple heuristics to navigate. Single vervets took the shortest possible path in fewer than half of the trials, usually in ways consistent with the most efficient heuristic (the convex hull). When in competition, vervets' paths were consistent with different, more efficient heuristics dependent on their dominance rank (a cluster strategy for dominants and the nearest neighbor rule for subordinates). These results suggest that, like humans, vervets may solve multidestination routing problems by applying simple, adaptive, context‐specific “rules of thumb.” The heuristics that were consistent with vervet paths in this study are the same as some of those asserted to be used by humans. These spatial movement strategies may have common evolutionary roots and be part of a universal mental navigational toolkit. Alternatively, they may have emerged through convergent evolution as the optimal way to solve multidestination routing problems.     

Spatial foraging patterns and colony energy status in the African honey bee,Apis mellifera scutellata

The relationship between changes in foraging patterns (inferred from waggle dance activity) and colony energy status (inferred from brood rearing activity, food storage, and colony weight) was examined for the African honey bee during a period of relative resource abundance and resource dearth. When resources were more abundant mean foraging distances (about 400 m) and foraging areas (4–5 km²) were small, and colonies recruited to 12–19 different sites per day. Colony foraging ranges and sites visited increased slightly during the dearth period, yet foraging continued to be concentrated within less than 10 km². The degree to which fluctuations in foraging patterns were correlated with colony energy status varied with the availability of floral resources. During periods of relative forage abundance, increases in foraging range and number of sites visited were significantly correlated with increases in brood rearing and colony weight. In contrast, colonies examined during periods of resource dearth exhibited no correlations between foraging areas, foraging distances, and fluctuations in brood rearing, food storage, or colony weight. Thus, during dearth periods colonies may not be able to coordinate foraging patterns with changes in colony energy status.     

Foraging responses ofFormica truncorum (Hymenoptera; Formicidae); exploiting stable vs spatially and temporally variable resources

Summary Social organization allows a division of labor between reproduction and foraging, as well as a task allocation among foraging individuals. Therefore, a colony simultaneously exploiting various resources can use different ways of getting them. This work analyzes wood ant foraging tactics both at the collective and the individual level. The main issues are: (1) How does a wood ant colony respond to stable vs temporally and spatially variable resources in terms of worker force allocation? (2) How is retrieval of ephemeral but rich resources effected at the individual level? The results show a correspondence between resource stability and behavior. Ants visited stable resources continually and recruited to ephemeral ones. They also visited empty food sites at a low frequency suggesting territoriality and constant scanning. The results suggest that recruitment may involve defence of the resource, since only a small fraction of all the workers available carry the food home. Additional ants arriving at the resource are recruited from inside the nest, not by reallocating outdoor workers. The foragers also form two separate groups, one site-persistent and another site-flexible. These two forager groups may reflect specialization on two different kinds of diet: honeydew or insect prey (spatially and temporally stable vs unstable resources, respectively).Formica truncorum is thus an example of how sociality allows the use of two different foraging strategies simultaneously: one that is based on recruitment to ephemeral resources and another that is based on search persistance and memory.     

Mantled Howler (<Emphasis Type="Italic">Alouatta palliata</Emphasis>) Arboreal Pathway Networks: Relative Impacts of Resource Availability and Forest Structure

Numerous studies have concluded that primates move about their environments in a nonrandom manner, frequently traveling between consecutive foraging sites along relatively straight-line paths. However, primates do not always take the most direct path between resources, and a number of species have been observed to travel repeatedly along a network of the same arboreal pathways. In this study, I used spatially explicit techniques to examine quantitatively what mantled howler monkey groups on Barro Colorado Island, Panama, accomplish by selecting nonlinear paths between resources and by repeatedly using the same paths within an arboreal network. Results show that chosen arboreal paths between sites where foraging occurred have higher levels of resource availability and canopy connectivity than comparable straight-line paths between the same sites. When comparing the relative importance of these factors, autologistic models of pathway choice indicate that though canopy connectivity is related to the location of repeatedly used arboreal pathway networks, the most statistically significant predictor is resource availability (both on a path and within a visual detection distance of a path). These results provide support for the hypothesis that repeated use of arboreal pathway networks aids in resource monitoring and acquisition. In addition, statistical models developed from 1 primary focal group’s travel patterns had high predictive value when employed to generate likely locations for arboreal pathways in the home ranges of 3 neighboring groups. This finding has important implications for studies of primate habitat use and seed dispersal, as it suggests that different groups consistently use similar characteristics when deciding on travel paths.     

A novel ACO algorithm for optimization via reinforcement and initial bias

In this paper, we introduce the MAF-ACO algorithm, which emulates the foraging behavior of ants found in nature. In addition to the usual pheromone model present in ACO algorithms, we introduce an incremental learning component. We view the components of the MAF-ACO algorithm as stochastic approximation algorithms and use the ordinary differential equation (o.d.e.) method to analyze their convergence. We examine how the local stigmergic interaction of the individual ants results in an emergent dynamic programming framework. The MAF-ACO algorithm is also applied to the multi-stage shortest path problem and the traveling salesman problem. Research of Prof. V.S. Borkar was supported in part by grant no. III.5(157)/99-ET and a J.C. Bose Fellowship from the Department of Science and Technology, Government of India.     

Diet, foraging, and use of space in wild golden-headed lion tamarins

Lion tamarins (Callitrichidae: Leontopithecus) are small frugi-faunivores that defend large home ranges. We describe results from the first long-term investigation of wild golden-headed lion tamarins (L. chrysomelas; GHLTs). We present data about activity budgets, daily activity cycles, diet, daily path length, home range size, home range overlap, and territorial encounters for three groups of GHLTs that were studied for 1.5-2.5 years in Una Biological Reserve, Bahia State, Brazil, an area characterized by aseasonal rainfall. We compare our results to those from other studies of lion tamarins to identify factors that may influence foraging and ranging patterns in this genus. Ripe fruit, nectar, insects, and small vertebrates were the primary components of the GHLT diet, and gums were rarely eaten. Fruit comprised the majority of plant feeding bouts, and the GHLTs ate at least 79 different species of plants from 32 families. The most common foraging sites for animal prey were epiphytic bromeliads. The GHLTs defended large home ranges averaging 123 ha, but showed strong affinities for core areas, spending 50% of their time in approximately 11% of their home range. Encounters with neighboring groups averaged two encounters every 9 days, and they were always aggressive. Data about time budgets and daily activity cycles reveal that the GHLTs spent most of their time foraging for resources or traveling between foraging sites distributed throughout their home ranges. The GHLTs spent much less time consuming exudates compared to lion tamarins in more seasonal environments. Additionally, the GHLTs had much larger home ranges than golden lion tamarins (L. rosalia), and did not engage in territorial encounters as frequently as L. rosalia. GHLT ranging patterns appear to be strongly influenced by resource acquisition and, to a lesser extent, by resource defense.     

The attraction of the known: the importance of spatial familiarity in habitat selection in wapiti Cervus elaphus

Most studies of habitat selection by large herbivores focus on the resource availability and interactions with other species, but neglect the importance of an animal being familiar with an area due to past use. Yet, studies of the establishment and retention of territories, home ranges, birth sites, and feeding site choices in experimental settings have shown the importance of spatial familiarity at these scales. We used GPS locations of translocated wapiti Cervus elaphus , resource selection functions (RSF), and time-to-return to examine whether previous site use was important for selection of sites by wapiti in west-central Alberta. To construct RSFs, we used logistic regression that included spatial familiarity (presence of a previous GPS location within a 50-m radius) as well as estimates of herbaceous and shrub biomass, elevation, aspect, slope, and predation risk to wapiti from wolf predation, as dependent variables. We found that previous use had a strong positive relationship with subsequent site use, indicating that wapiti were not avoiding previously visited locations, as would be expected if memory of forage depletion (which we did not measure) determined response to familiar locations. Revisited sites were of higher quality, i.e. had more moderate terrain, higher forage, and lower predation risk, than sites that were not revisited, indicating that the selection of familiar locations was likely not the result of avoidance of unfamiliar locations. Finally, animals demonstrated preference for familiar locations that it had visited most recently, indicating that memory (which would decline with time) of higher site quality, rather than high quality alone, influenced selection for familiar locations. We conclude that spatial familiarity is important not only for large scale processes such as selection of home range and territory, but for smaller scale habitat selection and foraging as well.     

Winter habitat–space use in a large arctic herbivore facing contrasting forage abundance

We analysed how changes in resource levels influence foraging trade-offs in late winter by wild Svalbard reindeer. Forage plants, and particularly lichens, were less abundant at the overgrazed Brøggerhalvøya compared with the neighbouring Sarsøyra. Strong interactions occurred between habitat selection, home range size, and feeding crater selection. At Brøggerhalvøya, radiocollared females generally selected productive habitat (high summer NDVI; Normalised Difference Vegetation Index). “Immigrants” at Sarsøyra (dispersed from Brøggerhalvøya in early winter) had similar habitat preferences, probably due to past experience. In contrast, “residents” at Sarsøyra were more influenced by abiotic conditions, using habitat with low NDVI, but selecting for high-quality forage (lichens) when cratering. This suggests more quality-based selection at the expense of quantity when forage abundance increases. Habitat–space use relationships also differed between the animal categories, as home range size decreased with availability of preferred habitat. Thus, changes in forage abundance can strongly influence winter habitat–space use interactions in predator-free systems.     

Hierarchical resource selection by impala in a savanna environment

The factors that affect resource selection by a foraging herbivore can vary according to the resources or conditions associated with particular levels of organization in the environment, and to the scales over which the herbivore perceives and responds to those resources and conditions. To investigate the role of forage in this hierarchical process, we studied resource selection by a mixed‐feeding large herbivore, the impala (Aepyceros melampus). We focussed on three spatial scales: plant species, feeding station and feeding patch. In paired sites where impala were and were not observed, we identified the plant species from which animals fed, the attributes of the plants, and the characteristics of the broader site. Across all three scales, plant species available as forage was central in determining resource selection by impala. At the species level, that effect was modified by the nutritional quality (greenness) and whether it was during a period of forage abundance or scarcity (season). At the feeding‐station level, overall greenness and biomass of the station were important, but their effects were modified by the season. At the feeding‐patch level, broader‐scale factors such as the type of vegetation cover had an important influence on resource selection. The grass Panicum maximum was a preferred forage species and a key resource determining the locations of feeding impala. Our findings support the idea that selection by a foraging herbivore at fine scales (i.e. diet selection) can have consequences for broader‐scale selection that result in observed patterns of habitat use and animal distribution.     

Foraging in nature: Contrasting responses to resource heterogeneity at small‐ and large‐spatial scales

A key problem faced by foragers is how to forage when resources are distributed heterogeneously in space. This heterogeneity and associated trade‐offs may change with spatial scale. Furthermore, foragers may also have to optimize acquiring multiple resources. Such complexity of decision‐making while foraging is poorly understood. We studied the butterfly Ypthima huebneri to examine how foraging decisions of adults are influenced by spatial scale and multiple resources. We predicted that, at a small‐spatial scale, the time spent foraging in a patch should be proportional to resources in the patch, but at large‐spatial scales, due to limitations arising from large travel costs, this relationship should turn negative. We also predicted that both adult and larval resources should jointly affect foraging butterflies. To test these predictions, we laid eleven plots and sub‐divided them into patches. We mapped nectar and larval resources and measured butterfly behavior in these patches and plots. We found that adult foraging behavior showed contrasting relationships with adult resource density at small versus large‐spatial scales. At the smaller‐spatial scale, butterflies spent more time feeding in resource‐rich patches, whereas at the large‐scale, butterflies spent more time feeding in resource‐poor plots. Furthermore, both adult and larval resources appeared to affect foraging decisions, suggesting that individuals may optimize search costs for different resources. Overall, our findings suggest that the variation in foraging behavior seen in foragers might result from animals responding to complex ecological conditions, such as resource heterogeneity at multiple spatial scales and the challenges of tracking multiple resources.     

Territorial Advertisement and Mate Attraction in the Bat Pipistrellus pipistrellus

A South Swedish population of the vespertilionid bat Pipistrellus pipistrellus was studied by means of bat boxes from April to the beginning of October in 1984. The mating system of the pipistrelle bat is a “resource defence polygyny”. Three territorial males were studied to determine the relative importance of two potential resources, food and roost sites, for male reproductive success. The day roost was found to be the crucial resource for the male's chances to get access to females. A territorial male advertises the location of his day roost to the females by a songflight display. The male who spent most time in songflight display was visited by the greatest number of females.     

Factors influencing forage selection by the North American beaver (Castor canadensis)

Across their range, a large number of biotic and abiotic factors are known to influence the choice of browse plant and the foraging behaviour of the North American beaver (Castor canadensis). We used generalized linear mixed-effects models to investigate sets of variables that may influence the foraging choices of beaver: forage species, distance of forage from water, forage density, and site. Communities across the study sites in central British Columbia, Canada, were dominated by Salix sitchensis, Salix lucida, and Alnus spp. Density had no impact on forage selection, while site, distance from water, and species identity all influenced the foraging decisions of beaver. We postulated that these factors may be ordered hierarchically: large-scale factors, such as site, followed by the medium-scale distance from water, and species of plant at the finest scale. Forage items in some sites had a higher probability of being browsed than in others, while in all sites the probability of being browsed decreased with increasing distance from water. Beaver appeared to be foraging as “picky” generalists; of the 9 plant species examined, 3 species of Salix (S. scouleriana, S. drummondiana and S. sitchensis) were selected by beaver, Salix bebbiana was avoided, and 5 species were neither selected for nor against. Browse selection within the genus Salix implied that beaver were able to differentiate among closely related species. Detailed information on forage selection is a crucial first step in designing and interpreting models that predict large-scale distributional patterns of beaver.     

Mapping the navigational knowledge of individually foraging ants,Myrmecia croslandi

Ants are efficient navigators, guided by path integration and visual landmarks. Path integration is the primary strategy in landmark-poor habitats, but landmarks are readily used when available. The landmark panorama provides reliable information about heading direction, routes and specific location. Visual memories for guidance are often acquired along routes or near to significant places. Over what area can such locally acquired memories provide information for reaching a place? This question is unusually approachable in the solitary foraging Australian jack jumper ant, since individual foragers typically travel to one or two nest-specific foraging trees. We find that within 10 m from the nest, ants both with and without home vector information available from path integration return directly to the nest from all compass directions, after briefly scanning the panorama. By reconstructing panoramic views within the successful homing range, we show that in the open woodland habitat of these ants, snapshot memories acquired close to the nest provide sufficient navigational information to determine nest-directed heading direction over a surprisingly large area, including areas that animals may have not visited previously.     

Return Customers: Foraging Site Fidelity and the Effect of Environmental Variability in Wide-Ranging Antarctic Fur Seals

Strategies employed by wide-ranging foraging animals involve consideration of habitat quality and predictability and should maximise net energy gain. Fidelity to foraging sites is common in areas of high resource availability or where predictable changes in resource availability occur. However, if resource availability is heterogeneous or unpredictable, as it often is in marine environments, then habitat familiarity may also present ecological benefits to individuals. We examined the winter foraging distribution of female Antarctic fur seals, Arctocephalus gazelle, over four years to assess the degree of foraging site fidelity at two scales; within and between years. On average, between-year fidelity was strong, with most individuals utilising more than half of their annual foraging home range over multiple years. However, fidelity was a bimodal strategy among individuals, with five out of eight animals recording between-year overlap values of greater than 50%, while three animals recorded values of less than 5%. High long-term variance in sea surface temperature, a potential proxy for elevated long-term productivity and prey availability, typified areas of overlap. Within-year foraging site fidelity was weak, indicating that successive trips over the winter target different geographic areas. We suggest that over a season, changes in prey availability are predictable enough for individuals to shift foraging area in response, with limited associated energetic costs. Conversely, over multiple years, the availability of prey resources is less spatially and temporally predictable, increasing the potential costs of shifting foraging area and favouring long-term site fidelity. In a dynamic and patchy environment, multi-year foraging site fidelity may confer a long-term energetic advantage to the individual. Such behaviours that operate at the individual level have evolutionary and ecological implications and are potential drivers of niche specialization and modifiers of intra-specific competition.     

The effect of feeding enrichment methods on the behavior of captive Western lowland gorillas

Three feeding enrichment treatments were tested in an outdoor yard used by six Western lowland gorillas (Gorilla gorilla gorilla). In "Yard-toss," forage was thrown by hand over one third of the yard. In "Set-up," forage and browse were hand-scattered throughout the yard. "Set-up Enriched" was similar with the addition of either a hay- and forage-filled feeder or forage-filled boomer ball(s) suspended from a climbing structure. Each treatment was presented on 5 d. Behavior was recorded for 30 min before (baseline) and 30 min after the start of each treatment. All treatments led to more foraging and less inactivity compared with baseline (P80.05), but Yard-toss was the least effective, likely because resources were clumped and monopolized by dominant animals. In Set-up Enriched, dominant animals had the greatest increase in foraging (P=0.03), partly because they generally monopolized the suspended items, but this allowed others to forage at ground level. This separation of the animals likely explains why Set-Up Enriched led to more foraging than all other treatments (P80.05). Findings show that for these hierarchical animals, enrichment resources are most effective when distributed widely, including vertically, and that enrichment strategies must take social structure into account.     

Colony size does not predict foraging distance in the ant Temnothorax rugatulus: a puzzle for standard scaling models

Body size is often positively correlated with ecologically relevant traits such as fecundity, survival, resource requirements, and home range size. Ant colonies, in some respects, behave like organisms, and their colony size is thought to be a significant predictor of many behavioral and ecological traits similar to body size in unitary organisms. In this study, we test the relationship between colony size and field foraging distance in the ant species Temnothorax rugatulus. These ants forage in the leaf litter presumably for small arthropod prey. We found colonies did not differ significantly in their foraging distances, and colony size is not a significant predictor of foraging distance. This suggests that large colonies may not exhaust local resources or that foraging trips are not optimized for minimal distance, and thus that food may not be the limiting resource in this species. This study shows T. rugatulus are behaving in ways that differ from existing models of scaling.     

Fine-scale resource selection and behavioral tradeoffs of eastern wild turkey broods

Resource heterogeneity across the landscape prompts animals to make behavioral tradeoffs to survive and reproduce. Behavioral thermoregulation can buffer organisms from thermal extremes but may conflict with other essential activities such as predator avoidance or foraging, and necessitate tradeoffs among resource requirements. We evaluated patterns of habitat selection relative to thermal conditions, forage availability, and concealment cover for female eastern wild turkeys (Meleagris gallopavo silvestris) with broods to assess potential tradeoffs among resource requirements. We quantified air temperature (°C), vegetation characteristics (e.g., visual obstruction), and arthropod biomass (g/m²) at locations used by broods across 5 study sites in the southeastern United States during May–July 2019–2020. We used conditional logistic regression to estimate brooding female resource selection at the second (home range) and third (within home range) orders. Specifically, we identified differences in selection between brooding and non-brooding females (second order), and factors influencing selection of sites used by brooding females during the day (when loafing and foraging) and night (roosting; third order). Brooding females selected sites with cooler temperatures (β = −0.22; 95% CI = −0.338–−0.102) and greater ground cover vegetation (β = 0.02; 95% CI = 0.013–0.033) than non-brooding females. Additionally, biomass of large prey (Orthoptera) was positively related to ambient temperature, suggesting that use of thermal refuge by brooding females may limit availability of large prey. Brooding females appeared to balance the tradeoff between thermal refuge and forage availability by altering habitat selection patterns within home ranges. Brooding females selected for herbaceous areas that provided greater biomass of large arthropods during the day, and avoided areas dominated by woody vegetation during both the day and night. We did not observe brooding females using locations where woody cover exceeded 27% of understory vegetation. Thermal refuge is an important component of brood habitat, but within thermally suitable areas brooding females can select sites with greater availability of large prey to meet energetic demands of broods. Evaluation of multiple spatial scales is key when assessing tradeoffs among resource needs and determining the potential of behavioral thermoregulation to buffer an organism's thermal environment and allow persistence in a warming climate.     

Ant Navigation: Fractional Use of the Home Vector

Home is a special location for many animals, offering shelter from the elements, protection from predation, and a common place for gathering of the same species. Not surprisingly, many species have evolved efficient, robust homing strategies, which are used as part of each and every foraging journey. A basic strategy used by most animals is to take the shortest possible route home by accruing the net distances and directions travelled during foraging, a strategy well known as path integration. This strategy is part of the navigation toolbox of ants occupying different landscapes. However, when there is a visual discrepancy between test and training conditions, the distance travelled by animals relying on the path integrator varies dramatically between species: from 90% of the home vector to an absolute distance of only 50 cm. We here ask what the theoretically optimal balance between PI-driven and landmark-driven navigation should be. In combination with well-established results from optimal search theory, we show analytically that this fractional use of the home vector is an optimal homing strategy under a variety of circumstances. Assuming there is a familiar route that an ant recognizes, theoretically optimal search should always begin at some fraction of the home vector, depending on the region of familiarity. These results are shown to be largely independent of the search algorithm used. Ant species from different habitats appear to have optimized their navigation strategy based on the availability and nature of navigational information content in their environment.     

Long-term home range stability provides foraging benefits in western gorillas

Objectives

Long-term home range stability presumably emerges because familiarity with an area improves fitness through increased foraging efficiency, reduced predation risk, or reduced costs of intergroup aggression. While the use of spatial memory by primates has been widely demonstrated, few studies have examined whether long-term space use creates opportunities for interannual reuse of spatial knowledge. Here we examine the ranging behavior of western gorillas (Gorilla gorilla) to assess the degree of long-term site fidelity and the foraging consequences of reuse of space.

Methods

We measured interannual home range overlap over a 10-year period for a single group of gorillas at the Mondika Research Center, using both grid-based and kernel density estimation. By plotting the total area used over time, we identified periods of home-range stability and expansion. We compared foraging and ranging behavior in familiar versus unfamiliar areas, considering fruit trees visited, dietary diversity, and daily path length, to determine whether the lack of spatial knowledge in unfamiliar areas was associated with foraging costs.

Results

Average interannual home range overlap by the group remained high throughout the study. During periods of home range expansion, daily path lengths increased but not the number of fruit trees visited, suggesting that reduced familiarity with the area led to decreased foraging efficiency because individuals lacked prior knowledge of where to find resources.

Discussion

Western gorillas at Mondika exhibit long-term home range stability, presumably reflecting a strategy that relies on the use of spatial memory to increase foraging efficiency that is favored by their reliance on ephemeral fruit resources.

     

Effects of predatory risk and resource renewal on the timing of foraging activity in a gerbil community

The foraging decisions of animals are often influenced by risk of predation and by the renewal of resources. For example, seed-eating gerbils on sand dunes in the Negev Desert of Israel prefer to forage in the bush microhabitat and during darker hours due to risk of predation. Also, daily renewal of seed resource patches and timing of nightly foraging activity in a depleting environment play important roles in species coexistence. We examined how these factors influence the timing of gerbil foraging by quantifying foraging activity in seed resource patches that we experimentally renewed hourly during the night. As in previous work, gerbils showed strong preference for the safe bush microhabitat and foraged less in response to high levels of illumination from natural moon light and from artificial sources. We demonstrate here for the first time that gerbils also responded to temporal and spatial heterogeneity in predatory risk through their timing of activity over the course of each night. Typically, gerbils concentrated their activity early in the night, but this changed with moon phase and in response to added illumination. These results can be understood in terms of the nature of patch exploitation by gerbils and the role played by the marginal value of energy in determining the cost of predation. They further show the dynamic nature of gerbil foraging decisions, with animals altering foraging efforts in response to time, microhabital, moon phase, illumination, and resource availability.