Perception of Interannual Covariation and Strategies for Risk Reduction among Mikea of Madagascar

This paper begins with the hypothesis that Mikea, participants in a mixed foraging–fishing–farming–herding economy of southwestern Madagascar, may attempt to reduce interannual variance in food supply caused by unpredictable rainfall by following a simple rule-of-thumb: Practice an even mix of activities that covary positively with rainfall and activities that covary negatively with rainfall. Results from a historical matrix participatory exercise confirm that Mikea perceive that foraging and farming outcomes covary positively or negatively with rainfall. This paper further considers whether Mikea learn about covariation through personal observation and memory recall (individual learning) or through socially transmitted ethnotheory (social learning). Dual inheritance theory models by Boyd and Richerson (1988) predict that individual learning is more effective in spatially and temporally variable environments such as the Mikea Forest. In contrast, the psychological literature suggests that individuals judge covariation poorly when memory of past events is required, unless they share a socially learned theory that a covariation should exist (Nisbett and Ross 1980). Results suggest that Mikea rely heavily on shared ethnotheory when judging covariation, but individuals continually strive to improve their judgment through individual observation.     

Behavioral states in space and time: understanding landscape use by an invasive mammal

Animal movement models can be used to understand species behavior and assist with implementation of management activities. We explored behavioral states of an invasive wild pig (Sus scrofa) population that recently colonized central Michigan, USA, 2014–2018. To quantify environmental factors related to wild pig movement ecology and spatio-temporal landscape use, we predicted wild pig behavioral states relative to land cover type, landscape structure (i.e., edge and patch cohesion), and weather conditions. We used global positioning system (GPS)-collars and monitored 8 wild pigs from 2014–2018. We fit local convex hulls and calculated movement metrics revealing 3 wild pig behavioral states (resting, exploratory, and relocating) and constructed a 3-level model to predict behavioral state probabilities relative to biotic and abiotic conditions. Probabilities of exploratory and resting behaviors were higher nearer to riparian and open herbaceous cover types (oftentimes emergent marsh), indicating that these cover types provided security cover during activity and bedding. Hard mast cover types had a strong positive association with relocating behaviors. More cohesive patches of agriculture and shrub cover types were associated with higher probabilities of exploratory behaviors, while resting was more likely in continuous patches of agriculture (mostly mid-summer corn). The probability of exploratory behaviors increased exponentially with warming ambient temperature. Our results may be used by managers to develop control strategies conducive to landscape and environmental conditions where the likelihood of encountering wild pigs is highest or targeting wild pigs when in a behavioral state most vulnerable to a particular removal technique.     

The social cognition of social foraging: partner selection by underlying valuation

Humans and other animals have a variety of psychological abilities tailored to the demands of asocial foraging, that is, foraging without coordination or competition with other conspecifics. Human foraging, however, also includes a unique element: the creation of resource pooling systems. In this type of social foraging, people contribute when they have excess resources and receive provisioning when in need. Is this behavior produced by the same psychology as asocial foraging? If so, foraging partners should be judged by the same criteria used to judge asocial patches of resources: the net energetic benefits they provide. The logic of resource pooling speaks against this. Maintaining such a system requires the ability to judge others not on their short-term returns, but on the psychological variables that guide their behavior over the long term. We test this idea in a series of five studies using an implicit measure of categorization. Results showed that (a) others are judged by the costs they incur (a variable not relevant to asocial foraging), whereas (b) others are not judged by the benefits they provide when benefits provided are unrevealing of underlying psychological variables (despite this variable being relevant to asocial foraging). These results are suggestive of a complex psychology designed for both social and asocial foraging.     

A Device to Study the Behavioral Responses of Zooplankton to Food Quality and Quantity

In order to explore the behavioral mechanisms underlying aggregation of foragers on local resource patches, it is necessary to manipulate the location, quality and quantity of food patches. This requires careful control over the conditions in the foraging arena, which may be a challenging task in the case of aquatic resource-consumer systems, like that of freshwater zooplankton feeding on suspended algal cells. We present an experimental tool designed to aid behavioral ecologists in exploring the consequences of resource characteristics for zooplankton aggregation behavior and movement decisions under conditions where the boundaries and characteristics (quantity and quality) of food patches can be standardized. The aggregation behavior of Daphnia magna and D. galeata x hyalina was tested in relation to i) the presence or absence of food or ii) food quality, where algae of high or low nutrient (phosphorus) content were offered in distinct patches. Individuals of both Daphnia species chose tubes containing food patches and D. galeata x hyalina also showed a preference towards food patches of high nutrient content. We discuss how the described equipment complements other behavioral approaches providing a useful tool to understand animal foraging decisions in environments with heterogeneous resource distributions.     

A dynamic habitat selection game

A patch selection game is formulated and analyzed. Organisms can forage in one of H patches. Each patch is characterized by the cost of foraging, the density and value of food, the predation risk, and the density of conspecifics. The presence of conspecifics affects the finding and sharing of food, and the predation risk. Optimal foraging theory can be viewed as a "1-person" game against nature in which the optimal patch choice of a specific organism is analyzed assuming that the number of conspecifics in other patches is fixed. In the general game theoretic approach, the behavior of conspecifics is included in the determination of the distinguished organism's strategy. An iterative algorithm is used to compute the solution of the "n-person" game or dynamic ESS, which differs from the optimal foraging theory solution. Experiments to test the proposed theory using rodents and seed trays are briefly discussed.     

Use of native woodlands and traditional olive groves by foraging bats on a Mediterranean island: consequences for conservation

We recorded bat activity on Zakynthos island (Greece) to test the hypotheses that (1) olive ( Olea europea ) groves and native woodlands provide comparable foraging habitat for insectivorous bats, (2) lower foraging activity occurs in olive groves treated with insecticide chemicals. We acoustically sampled bat activity (passes per minute) in four wooded habitats (organic and non-organic olive groves, oak woodland ( Quercus ilex and Quercus coccifera ) and pine ( Pinus halepensis ) woodland from June to August 2005. Habitat type did not affect overall bat activity. A single application of insecticide chemicals annually did not affect activity over traditional olive groves. Habitat use on the island differed in several ways from that reported in studies at mainland sites. Most strikingly, pine woodland supported higher bat activity than expected relative to other habitat types, and we recorded unexpectedly high levels of M. capaccinii activity in woodland habitats. We suggest that traditional olive groves buffer some bat species from the effects of deforestation. Conservation plans for Mediterranean bats should consider the biodiversity value of these groves along with the need to conserve small woodland patches. Finally, understanding island-specific patterns of habitat use is essential to bat conservation on small off-shore islands.     

Root and tuber phytoliths and starch grains document manioc (Manihot esculenta) arrowroot (Maranta arundinacea) and llerén (Calathea sp.) at the real alto site Ecuador

Although roots and tubers are dietary staples in many parts of the world, their use is difficult to document archaeo logically because their organic remains are often poorly preserved in archaeological sedi ments. Here we describe the first diagnostic phytoliths from the underground storage organs of the important New World agricultural crops manioc or yuca(Manihot esculenta Crantz), arrowroot(Maranta arundinacea L.), and llerén(Calathea allouia [Aubl.] Lindl.) and demonstrate their usefulness for identifying prehistoric root and tuber processing with a study of stone artifacts from a Valdivia 3 (2800–2400 B.C., calibrated) household at Real Alto, Ecuador. Gelatinized starch (heat-altered) and unaltered starch from maize (Zea mays L.), arrowroot, and manioc were also found on these stone tools. Our data document early evidence for manioc in Ecuador’s coastal lowlands. In combination, these phytoliths and starch residues provide evidence that both raw and cooked foods were processed in this early mixed agricultural economy.     

Energy benefits and emergent space use patterns of an empirically parameterized model of memory‐based patch selection

Many species frequently return to previously visited foraging sites. This bias towards familiar areas suggests that remembering information from past experience is beneficial. Such a memory‐based foraging strategy has also been hypothesized to give rise to restricted space use (i.e. a home range). Nonetheless, the benefits of empirically derived memory‐based foraging tactics and the extent to which they give rise to restricted space use patterns are still relatively unknown. Using a combination of stochastic agent‐based simulations and deterministic integro‐difference equations, we developed an adaptive link (based on energy gains as a foraging currency) between memory‐based patch selection and its resulting spatial distribution. We used a memory‐based foraging model developed and parameterized with patch selection data of free‐ranging bison Bison bison in Prince Albert National Park, Canada. Relative to random use of food patches, simulated foragers using both spatial and attribute memory are more efficient, particularly in landscapes with clumped resources. However, a certain amount of random patch use is necessary to avoid frequent returns to relatively poor‐quality patches, or avoid being caught in a relatively poor quality area of the landscape. Notably, in landscapes with clumped resources, simulated foragers that kept a reference point of the quality of recently visited patches, and returned to previously visited patches when local patch quality was poorer than the reference point, experienced higher energy gains compared to random patch use. Furthermore, the model of memory‐based foraging resulted in restricted space use in simulated landscapes and replicated the restricted space use observed in free‐ranging bison reasonably well. Our work demonstrates the adaptive value of spatial and attribute memory in heterogeneous landscapes, and how home ranges can be a byproduct of non‐omniscient foragers using past experience to minimize temporal variation in energy gains.     

People's study time allocation and its relation to animal foraging

In this article we suggest a relation between people's metacognitively guided study time allocation strategies and animal foraging. These two domains are similar insofar as people use specific metacognitive cues to assist their study time allocation just as other species use cues, such as scent marking. People decline to study items that they know they already know, just as other species use a win-shift strategy - avoiding already visited and depleted patches - in foraging. People selectively study the easiest as-yet-unlearned items first, before turning to more difficult items just as other species take the ‘just right’ size and challenge of prey—the so-called Goldilocks principle. People use a stop rule by which they give up on one item and turn to another when the returns diminish just as others species use a stop rule that guides shifting from one patch to another. The value that each item is assigned on the criterion test, if known during study, influences which items people choose to study and how long they study them just as knowledge of the nutritional or energy value of the food influences choices and perseverance in foraging. Finally, study time allocation strategies can differ in their effectiveness depending upon the expertise of the student just as some species forage close to optimally while others do not.     

Unpacking chimpanzee (Pan troglodytes) patch use: Do individuals respond to food patches as predicted by the marginal value theorem?

The marginal value theorem is an optimal foraging model that predicts how efficient foragers should respond to both their ecological and social environments when foraging in food patches, and it has strongly influenced hypotheses for primate behavior. Nevertheless, experimental tests of the marginal value theorem have been rare in primates and observational studies have provided conflicting support. As a step towards filling this gap, we test whether the foraging decisions of captive chimpanzees (Pan troglodytes) adhere to the assumptions and qualitative predictions of the marginal value theorem. We presented 12 adult chimpanzees with a two-patch foraging environment consisting of both low-quality (i.e., low-food density) and high-quality (i.e., high-food density) patches and examined the effect of patch quality on their search behavior, foraging duration, marginal capture rate, and its proxy measures: giving-up density and giving-up time. Chimpanzees foraged longer in high-quality patches, as predicted. In contrast to predictions, they did not depress high-quality patches as thoroughly as low-quality patches. Furthermore, since chimpanzees searched in a manner that fell between systematic and random, their intake rates did not decline at a steady rate over time, especially in high-quality patches, violating an assumption of the marginal value theorem. Our study provides evidence that chimpanzees are sensitive to their rate of energy intake and that their foraging durations correlate with patch quality, supporting many assumptions underlying primate foraging and social behavior. However, our results question whether the marginal value theorem is a constructive model of chimpanzee foraging behavior, and we suggest a Bayesian foraging framework (i.e., combining past foraging experiences with current patch sampling information) as a potential alternative. More work is needed to build an understanding of the proximate mechanisms underlying primate foraging decisions, especially in more complex socioecological environments.     

The failure of the introduction of wild boar Sus scrofa in the island of Cyprus: a case study

Wild boar (Sus scrofa L.) were introduced in the island of Cyprus in 1990, when five animals were imported from Greece for game farming. In 1994, wild boars were illegally released in Lemesos (Limassol) Forest and in 1996 in the Troodos National Forest Park. Soon the population increased and dispersed throughout the park. In 1997, the government of Cyprus decided to eradicate wild boar because of the danger of transmitting diseases to livestock and to prevent possible environmental destruction. To control the wild boar, hunting was permitted and game wardens were instructed to eliminate the free-ranging animals. In 2004, no animals were observed in localities where they had been seen before. Surveys in September 2004 (Troodos National Park) and January/February 2005 (Troodos Forest, Pafos Forest, and Lemesos Forest) revealed no signs of recent wild boar presence. The reasons for the possible failure of wild boar to establish in Cyprus are discussed.     

Foraging costs,hunting success and its implications for African wild dog (Lycaon pictus) conservation inside and outside a protected area

When selecting a habitat, animals utilize habitat in which they yield the highest rate of energy. Differences in foraging costs and hunting success are therefore likely to affect habitat choice. In a previous study, we showed that African wild dog (Lycaon pictus) packs with territories inside Hwange National Park (HNP), over the course of several years, moved their territories into the buffer zone outside HNP, where reproductive success was higher but anthropogenic mortality exceeded natality. In this study, based on long‐term radio‐telemetry data from 22 African wild dog packs, we analysed whether differences in foraging costs and hunting success could have contributed to this territorial drift. Taking seasonality and pack size into account, we determined foraging costs (foraging distance and chase distance) and hunting success (successful or failed chase) inside and outside HNP. Although we observed no difference in foraging costs, hunting success was higher outside HNP, which is likely to have contributed to the territorial drift into the buffer zone outside the protected area. This study shows the importance of taking factors affecting hunting success into account in the conservation strategy of African wild dogs.     

Sex differences in giraffe foraging behavior at two spatial scales

We test predictions about differences in the foraging behaviors of male and female giraffes (Giraffa camelopardalis tippelskirchi Matchie) that derive from a hypothesis linking sexual size dimorphism to foraging behavior. This body-size hypothesis predicts that males will exhibit specific behaviors that increase their dry-matter intake rate relative to females. Foraging behavior was examined at two hierarchical levels corresponding to two spatial and temporal scales, within patches and within habitats. Patches are defined as individual trees or shrubs and habitats are defined as collections of patches within plant communities. Males were predicted to increase dry-matter intake rate within patches by taking larger bites, cropping bites more quickly, chewing less, and chewing faster. Within habitats, males were expected to increase intake rate by increasing the proportion of foraging time devoted to food ingestion as opposed to inter-patch travel time and vigilance. The predictions were tested in a free-ranging population of giraffes in Mikumi National Park, Tanzania. Males spent less total time foraging than females but allocated a greater proportion of their foraging time to forage ingestion as opposed to travel between patches. There was no sex difference in rumination time but males spent more time in activities other than foraging and rumination, such as walking. Within patches, males took larger bites than females, but females cropped bites more quickly and chewed faster. Males had longer per-bite handling times than females but had shorter handling times per gram of intake. Within habitats, males had longer average patch residence times but there was no significant sex difference in inter-patch travel times. There was no overall difference between sexes in vigilance while foraging, although there were significant sex by habitat and sex by season interactions. Although not all the predictions were confirmed, overall the results agree qualitatively with the body-size hypothesis. Sex-related differences in foraging behavior led to greater estimated intake rates for males at the within-patch and within-habitat scales. Received: 20 November 1995 / Accepted: 5 November 1996     

The Behavioral Ecology of Color Vision: Considering Fruit Conspicuity,Detection Distance and Dietary Importance

Primate color vision is well suited for investigating the genetic basis of foraging behavior owing to a clear genotype–phenotype linkage. Finding fruits amid tropical foliage has long been proffered as an adaptive explanation for primate trichromacy, yet there is a dearth of systematic evaluations of frugivory as an ecological selective force. We studied the behavioral ecology of wild capuchins (Cebus capucinus) in northwestern Costa Rica across the annual cycle and modeled the ability of three dichromatic and three trichromatic phenotypes to discriminate fruits from leaves, a task that represents long-distance search for food patches in a tropical forest. Models of the trichromatic phenotypes could correctly discriminate approximately three-quarters of the total capuchin dietary fruits from leaves, including some fruits subjectively classified as having “cryptic” (greenish-brownish) hues. In contrast, models of dichromatic phenotypes could discriminate fewer than one-third of the fruits. This pattern held when we restricted our analysis to only the most heavily consumed diet items, preferred foods, or seasonally critical species. We in addition highlight the potential of fruit species with small patch sizes to confer an advantage to trichromats, as these resources are anticipated to provide a high finder’s reward. Our results are consistent with the hypothesis that long-distance detection of fruit patches exerts a selective pressure on trichromacy in neotropical primates, and suggest that greenish-brownish fruits might have played an underappreciated role in the evolution of primate color vision.     

不同天气条件下青头潜鸭越冬期行为的响应模式

2018年11—12月在河南民权国家湿地公园,采用定点观察法、扫描取样法对越冬青头潜鸭在不同天气条件下的行为响应模式进行研究。结果表明: 在晴天,静息、飞行、运动是青头潜鸭越冬期的优势行为(65.5%),觅食、修整是次要行为(31.9%);行为节律表现为上午觅食、中午静息、下午觅食修整;飞行通常发生在觅食高峰前,运动行为多与其他行为相伴,其时间分配与觅食呈正响应,与静息呈负响应。相较于晴天,阴雨天青头潜鸭增加了对静息和觅食的时间分配,静息、觅食和运动为阴雨天的主要行为(76.5%),修整、飞行为次要行为(20.3%);觅食和静息的高峰时段后移,傍晚时达到高峰,飞行和修整时间显著减少。与晴天相比,雾霭天青头潜鸭静息、觅食、运动和修整的时间分配增加,飞行行为减少;觅食高峰后移至中午和下午,静息高峰后移至下午;静息、运动和觅食为雾霭天的优势行为(70.6%),修整和飞行为次要行为(27.5%)。研究表明,不同天气条件下,民权湿地公园青头潜鸭的越冬期行为在时间分配、行为节律、优势行为方面存在一定的响应;阴雨天和雾霭天青头潜鸭会分配更多的静息和觅食时间,并通过增加能量摄入和减少耗能行为应对恶劣天气。     

Learning rules for social foragers: implications for the producer-scrounger game and ideal free distribution theory

In population games, the optimal behaviour of a forager depends partly on courses of action selected by other individuals in the population. How individuals learn to allocate effort in foraging games involving frequency-dependent payoffs has been little examined. The performance of three different learning rules was investigated in several types of habitats in each of two population games. Learning rules allow individuals to weigh information about the past and the present and to choose among alternative patterns of behaviour. In the producer-scrounger game, foragers use producer to locate food patches and scrounger to exploit the food discoveries of others. In the ideal free distribution game, foragers that experience feeding interference from companions distribute themselves among heterogeneous food patches. In simulations of each population game, the use of different learning rules induced large variation in foraging behaviour, thus providing a tool to assess the relevance of each learning rule in experimental systems. Rare mutants using alternative learning rules often successfully invaded populations of foragers using other rules indicating that some learning rules are not stable when pitted against each other. Learning rules often closely approximated optimal behaviour in each population game suggesting that stimulus-response learning of contingencies created by foraging companions could be sufficient to perform at near-optimal level in two population games.     

To dare or not to dare? Risk management by owls in a predator–prey foraging game

In a foraging game, predators must catch elusive prey while avoiding injury. Predators manage their hunting success with behavioral tools such as habitat selection, time allocation, and perhaps daring—the willingness to risk injury to increase hunting success. A predator’s level of daring should be state dependent: the hungrier it is, the more it should be willing to risk injury to better capture prey. We ask, in a foraging game, will a hungry predator be more willing to risk injury while hunting? We performed an experiment in an outdoor vivarium in which barn owls (Tyto alba) were allowed to hunt Allenby’s gerbils (Gerbillus andersoni allenbyi) from a choice of safe and risky patches. Owls were either well fed or hungry, representing the high and low state, respectively. We quantified the owls’ patch use behavior. We predicted that hungry owls would be more daring and allocate more time to the risky patches. Owls preferred to hunt in the safe patches. This indicates that owls manage risk of injury by avoiding the risky patches. Hungry owls doubled their attacks on gerbils, but directed the added effort mostly toward the safe patch and the safer, open areas in the risky patch. Thus, owls dared by performing a risky action—the attack maneuver—more times, but only in the safest places—the open areas. We conclude that daring can be used to manage risk of injury and owls implement it strategically, in ways we did not foresee, to minimize risk of injury while maximizing hunting success.     

Altruistic cooperation during foraging by the Ache, and the evolved human predisposition to cooperate

This paper presents quantitative data on altruistic cooperation during food acquisition by Ache foragers. Cooperative activities are defined as those that entail a cost of time and energy to the donor but primarily lead to an increase in the foraging success of the recipient. Data show that Ache men and women spend about 10% of all foraging time engaged in altruistic cooperation on average, and that on some days they may spend more than 50% of their foraging time in such activities. The most time-consuming cooperative activity for both sexes is helping during the pursuit of game animals, a pattern that is probably linked to the widespread sharing of game by Ache foragers. Cooperative food acquisition and subsequent food redistribution in hunter-gatherer societies are critical behaviors that probably helped shape universal, evolved, cooperative tendencies that are well illustrated in modern experimental economics. This paper was originally presented at a Max Planck symposium on cooperation in Leipzig in June 1999. This work was partially funded by a grant from the L.S.B. Leakey Foundation and NSF grant BNS 9727656. Kim Hill is a professor of anthropology in the Human Evolutionary Ecology (HEE) program at the University of New Mexico. His primary research interests include hunter-gatherer behavioral ecology, life history theory, food acquisition strategies, food sharing, cooperation, and biodiversity conservation in lowland South America. He has done fieldwork with Nahautl, Ache, Guarani, Hiwi, Mashco Piro, Matsiguenga, and Yora indigenous peoples of Central and South America.     

Virgins in the wild: mating status affects the behavior of a parasitoid foraging in the field

In haplodiploid organisms, virgin females can produce offspring, albeit only sons. They may therefore face a trade-off between either: (1) searching for hosts and producing sons immediately; or (2) searching for mates and perhaps producing both sons and daughters later in life. Although this trade-off raises a theoretical interest, it has not been approached experimentally. The objective of this article is thus to document the effect of mating status on the foraging behavior of a haplodiploid parasitoid. For this, we recorded the behavior of virgin and mated female Lysiphlebus testaceipes (Hymenoptera: Braconidae) after being released, in the field, on a colony of their aphid hosts. Half of the virgin females were mated by a wild male after less than 10 min of foraging. Evidently, virgin females attract males while foraging on host patches, so that the two activities are not mutually exclusive. Nonetheless, virgin females stayed motionless more often and for longer periods than mated females. Consequently, they attacked aphids at a lower rate, and in turn, attacked fewer aphids on each patch. Moreover, contrary to mated females, virgins did not aggregate their progeny on large patches. We conclude that in L. testaceipes, the trade-off may not be as hypothesized. By dispersing across patches more than mated females, virgins could promote future mating opportunities for their sons and increase their inclusive fitness. However, by moving too frequently, females may lose immediate mating opportunities for themselves and the immediate advantage of producing offspring of both sexes. The observed behavior of virgin L. testaceipes females on host patches could reflect an optimal solution to such a trade-off.     

Cattle select African savanna termite mound patches less when sharing habitat with wild herbivores

African savanna termite mounds function as nutrient‐rich foraging hotspots for different herbivore species, but little is known about their effects on the interaction between domestic and wild herbivores. Understanding such effects is important for better management of these herbivore guilds in landscapes where they share habitats. Working in a central Kenyan savanna ecosystem, we compared selection of termite mound patches by cattle between areas cattle accessed exclusively and areas they shared with wild herbivores. Termite mound selection index was significantly lower in the shared areas than in areas cattle accessed exclusively. Furthermore, cattle used termite mounds in proportion to their availability when they were the only herbivores present, but used them less than their availability when they shared foraging areas with wild herbivores. These patterns were associated with reduced herbage cover on termite mounds in the shared foraging areas, partly indicating that cattle and wild herbivores compete for termite mound forage. However, reduced selection of termite mound patches was also reinforced by higher leafiness of Brachiaria lachnantha (the principal cattle diet forage species) off termite mounds in shared than in unshared areas. Taken together, these findings suggest that during wet periods, cattle can overcome competition for termite mounds by taking advantage of wildlife‐mediated increased forage leafiness in the matrix surrounding termite mounds. However, this advantage is likely to dissipate during dry periods when forage conditions deteriorate across the landscape and the importance of termite mounds as nutrient hotspots increases for both cattle and wild herbivores. Therefore, we suggest that those managing for both livestock production and wildlife conservation in such savanna landscapes should adopt grazing strategies that could lessen competition for forage on termite mounds, such as strategically decreasing stock numbers during dry periods.