Maximizing feeding efficiency and minimizing time exposed to predators: a trade-off in the black-capped chickadee

Summary Animals often must feed away from protective cover, sometimes at a considerable risk of being preyed upon. Feeding at the maximum rate while away from cover may simultaneously minimize the time spent exposed to predators, but this is not always the case. Under some circumstances, carrying prey items to protective cover before they are consumed will minimize the time spent exposed to predators, whereas feeding at maximum efficiency (staying to eat prey where they are found) will actually increase the time spent exposed to predators. Whether or not there is a conflict between maximizing foraging efficiency and minimizing exposure time, depends upon the travel time to cover relative to the handling time of a prey item; short handling times and/or long travel times are associated with the no-conflict situation, whereas the conflict situation is associated with long handling times and/or short travel times to cover. Free-ranging chickadees foraging at an artificial patch at various distances from cover can distinguish between these two foraging situations. When there is no conflict, they stay and eat at the patch. Their behavior in the conflict situation indicates that they are tradingoff foraging considerations against the risk of predation. When the cost of carrying is low and the benefit gained is high, the chickadees elect to carry items to cover; they tend to stay and eat at the patch when the relative magnitudes of costs and benefits are reversed.     

Foraging-efficiency-predation-risk trade-off in the grey squirrel

Many animals must often respond to environmental patterns that simulatneously influence both foraging efficiency and predation risk. We noted that grey squirrels (Sciurus carolinensis) sometimes immediately consume food items in areas of relatively great exposure to predators, and at other times carry food items to the safety of a tree prior to consumption. We outlined a hypothesis that the squirrels were somehow trading-off energy intake rate against predation risk. A simple model shows that maximal energetic efficiency is associated with immediate consumption, whereas (under the field conditions studied) carrying items to the safety of trees provides for minimal exposure to predation. Our analysis of the model predicts that the tendency to carry a food item should decrease with distance of food from cover (travel time) and increase with item size (handling time). To test our predictions, we presented free-roaming grey squirrels with patches containing a fixed number of identical food items. We estimated the proportion of items carried to trees before consumption for 12 different combinations of distance to the nearest tree and item size. The results support our hypothesis and indicate that a simple behavioural criterion based solely on foraging rate or time exposed to predators is insufficient to explain the variation in the data.     

The Little Miss Muffet Effect: Quantifying the Effect of Predation Risk on Foraging Patch Choice by Houseflies (<Emphasis Type="Italic">Musca domestica</Emphasis>)

The ability of the ideal free distribution (IFD) to predict patch choice of female houseflies (Musca domestica) was determined by examining their distribution between two patches containing unequal amounts of food. The effect of predation risk was then quantified in energetic terms by examining fly distribution between patches of equal food, with one containing spiders. Results were used to predict how much extra food must be added to the risky patch to offset the risk of predation. Flies were found to conform fairly closely to the IFD. Predation risk had a major effect on their distribution, with fewer flies feeding in the presence of predators as risk increased. Addition of extra food to the risky patch was successful in offsetting the risk of predation. These results suggest that the effect of risk on housefly foraging behavior can be quantified in energy terms, providing a common currency for predicting the effects of resources and predation risk on habitat use.     

Relative Roles of Grey Squirrels,Supplementary Feeding,and Habitat in Shaping Urban Bird Assemblages

Non-native species are frequently considered to influence urban assemblages. The grey squirrel Sciurus carolinensis is one such species that is widespread in the UK and is starting to spread across Europe; it predates birds’ nests and can compete with birds for supplementary food. Using distance sampling across the urbanisation intensity gradient in Sheffield (UK) we test whether urban grey squirrels influence avian species richness and density through nest predation and competition for supplementary food sources. We also assess how urban bird assemblages respond to supplementary feeding. We find that grey squirrels slightly reduced the abundance of breeding bird species most sensitive to squirrel nest predation by reducing the beneficial impact of woodland cover. There was no evidence that grey squirrel presence altered relationships between supplementary feeding and avian assemblage structure. This may be because, somewhat surprisingly, supplementary feeding was not associated with the richness or density of wintering bird assemblages. These associations were positive during the summer, supporting advocacy to feed birds during the breeding season and not just winter, but explanatory capacity was limited. The amount of green space and its quality, assessed as canopy cover, had a stronger influence on avian species richness and population size than the presence of grey squirrels and supplementary feeding stations. Urban bird populations are thus more likely to benefit from investment in improving the availability of high quality habitats than controlling squirrel populations or increased investment in supplementary feeding.     

捕食风险对不同饥饿状态下艾虎取食行为的影响

在室内条件下,将大鵟作为艾虎的天敌动物,通过双通道选择实验确定6 只成体艾虎在3 个捕食风险水平和4 种饥饿状态条件下的取食行为,探讨艾虎在取食过程中对饥饿风险与捕食风险的权衡策略。研究结果表明:在无捕食风险存在时,艾虎被剥夺食物0 d 和1 d 后对食物量不同的两个斑块中的取食量和利用频次均无明显不同(P > 0. 05),但对高食物量斑块的利用时间均明显高于低食物量斑块的(P <0.05),而艾虎被剥夺食物2 d和3 d后对高食物量斑块中的取食量和利用时间均明显高于低食物量斑块中的(P < 0.05),但在利用频次上均无明显差异(P > 0.05)。在面临低风险时,艾虎在4 种饥饿状态下均只利用无天敌动物存在的低食物量斑块,而基本不利用有天敌动物存在的高食物量斑块。在面临高风险时,艾虎不得不利用有天敌动物存在的食物斑块,被剥夺食物0 d 时艾虎对无风险、无食物量斑块的利用时间基本相同于对高风险、有食物量斑块的利用时间(P>0.05),而被剥夺食物1d、2 d 和3 d 后艾虎对高风险、有食物量斑块的利用时间明显高于无风险、无食物量斑块的(P< 0. 05)。在相同风险条件下,随着饥饿程度增加,艾虎在斑块中的取食量均明显增加(P< 0.05),而对斑块的利用时间和利用频次明显降低(P<0.05)。在相同的饥饿状态下,不同风险水平时,艾虎在斑块中的取食量无明显的差异(P>0.05),但在低风险和高风险时对斑块的利用时间和频次均明显低于无风险时的(P <0.05)。以上结果说明艾虎能够根据食物摄取率和自身的能量需求在捕食风险和饥饿风险之间做出权衡,当饥饿风险小于捕食风险时,艾虎趋于躲避捕食风险,当饥饿风险大于捕食风险时,艾虎趋于面对捕食风险,所采用的取食策略是减少活动时间和能量消耗,最大程度地提高单位时间内获得的能量。     

Foraging Patch Selection in Winter: A Balance between Predation Risk and Thermoregulation Benefit

In winter, foraging activity is intended to optimize food search while minimizing both thermoregulation costs and predation risk. Here we quantify the relative importance of thermoregulation and predation in foraging patch selection of woodland birds wintering in a Mediterranean montane forest. Specifically, we account for thermoregulation benefits related to temperature, and predation risk associated with both illumination of the feeding patch and distance to the nearest refuge provided by vegetation. We measured the amount of time that 38 marked individual birds belonging to five small passerine species spent foraging at artificial feeders. Feeders were located in forest patches that vary in distance to protective cover and exposure to sun radiation; temperature and illumination were registered locally by data loggers. Our results support the influence of both thermoregulation benefits and predation costs on feeding patch choice. The influence of distance to refuge (negative relationship) was nearly three times higher than that of temperature (positive relationship) in determining total foraging time spent at a patch. Light intensity had a negligible and no significant effect. This pattern was generalizable among species and individuals within species, and highlights the preponderance of latent predation risk over thermoregulation benefits on foraging decisions of birds wintering in temperate Mediterranean forests.     

The anatomy of a range expansion: changes in cranial morphology and rates of energy extraction for North American red squirrels from different latitudes

Species with expanding ranges provide unique opportunities to examine environmentally induced adaptations in ecological traits and anatomical characteristics. Since the late 1800s, the North American red squirrel (Tamiasciurus hudsonicus) has expanded its range into the central hardwoods of the United States in conjunction with increasing agricultural fragmentation. We examined whether red squirrels from the central hardwoods (west‐central Indiana, USA) displayed differences in foraging behaviors and morphology relative to red squirrels from conifer‐dominated environments (upper peninsula of Michigan, USA), a biome in which red squirrels evolved. Specifically, we measured rates of energy extraction, variation in cranial morphology, and diet preference between red squirrels from both regions. In addition, we compared foraging behaviors of red squirrels from the central hardwoods to those of a competitor that coevolved with nut‐producing trees, the gray squirrel (Sciurus carolinensis). Red squirrels from Indiana and Michigan differed significantly in the efficiency with which they used food items, with individuals from each region extracting calories at a more rapid rate for items that were common in their region. The enhanced efficiency of southern red squirrels feeding on black walnuts (Juglans nigra) was correlated with geographic differences in cranial morphology; skulls of southern squirrels were larger, with longer jaws and higher metrics associated with greater mandibular force than northern squirrels. Contrary to our expectations, red squirrels from Indiana and Michigan did not differ qualitatively in preferences for food items, suggesting that diet choice may be governed by perishability of food items rather than by rates of energy extraction. Gray squirrels were more efficient than Indiana red squirrels in using all food items, and differed only slightly from red squirrels with regard to preference for food items. Measures of efficiency of resource use, after accounting for species‐specific metabolic requirements, suggest that red squirrels are unlikely to compensate ecologically for declining gray squirrel populations in fragmented portions of the central hardwoods, with potentially adverse effects for forest regeneration and succession. Our results demonstrate that invading species can display significant flexibility in adapting to new environments, but they may not be flexible enough to exploit resources in a manner comparable to native species.     

Effects of stoat's presence and auditory cues indicating its presence on tree seedling predation by meadow voles

Predators may control the impact of herbivores on their plant resources by 1) decreasing herbivore numbers, 2) imposing predation risk affecting foraging behavior. The goal of the present study was to examine the effects of a predator and auditory cues indicating its presence on the rate of tree seedling (Acer rubrum, Betula lenta) consumption by meadow voles (Microtus pennsylvanicus). The first of our experiments involved introduction of a stoat (Mustela erminea) into an enclosed vole population and the second a playback of recordings of vole distress calls, movements of a stoat and its vocalizations. In both experiments we manipulated vegetation cover and the availability of food next to the experimental seedlings to assess the effects of microhabitat under the different risk situations. The results of the first unreplicated experiment suggested an increased rate of seedling predation in the presence of the stoat. Consistent with these results, the playback of predator sounds in the second replicated experiment caused an increased rate of seedling predation compared to control plots with no recordings. A mowed circle around a seedling station, representing increased risk of predation on the voles, decreased seedling consumption. This effect was modest in the playback treatment. We suggest the results to be due to displacement behavior by the voles exposed to prolonged risk and conflicting demands of foraging and avoiding predators. Alternatively, as suggested by the model of Lima and Bednekoff, prolonged risk of predation forced the voles to decrease their levels of vigilance during low‐risk playback breaks. The modest inhibitory effect of cover removal on seedling predation in the playback treatment is consistent with this interpretation. The results confirm recent evidence for trophic cascades mediated by behavioral interactions between predator and prey. They are novel in suggesting that the presence of predation risk can increase the inhibitory effects of consumers on their resources.     

A kinematic investigation into the feeding behavior of the Goliath grouper <Emphasis Type="Italic">Epinephelus itajara</Emphasis>

Goliath grouper Epinephelus itajara suffered significant overfishing in the United States until they were protected from harvest in 1990. As the population recovers, interactions between Goliath grouper and anglers have increased, and are often reported to management agencies as complaints after grouper predation upon hooked fish. Goliath grouper are generally characterized as opportunistic predators capable of consuming a wide variety of prey types; however, minimal data are available regarding the prey capture behavior of this species. Kinematic analyses of adult Goliath grouper feeding events demonstrated the capacity of individuals to modulate feeding behavior based upon the mobility and position of ‘prey’ items. Mobile epibenthic food (tethered swimming fish) elicited larger maximum gapes, faster times to food capture, shorter times to mouth closing, and more rapid total bite durations than food items that were not moving (cut dead bait). Feeding sequences involving mobile food items were characterized by a significantly higher degree of ram feeding behavior, while immobile food elicited primarily suction feeding and were preceded by a slower and closer approach to the food item prior to the onset of mouth opening. The findings are discussed in light of predation upon angled species and demonstrate the ability of Goliath grouper to adjust their feeding strategy based upon prey type and condition. This behavior likely allows for the exploitation of a wide variety of prey and provides an expansive dietary breadth for these opportunistic predators.     

Nest predation in forest birds: influence of predator type and predator's habitat quality

We used the introduction of a generalist nest predator, the red squirrel Tamiasciurus hudsonicus, and of a large herbivore, the Sitka black-tailed deer Odocoileus hemionus sitkensis, to the islands of Haida Gwaii (Queen Charlotte Islands, British Columbia, Canada) to study how predator assemblage and habitat quality and structure influenced nest predation in forest birds. We compared losses of natural nests to predators on islands with and without squirrels. We selected nine islands with or without squirrel or deer and used 506 artificial nests put on the ground or in shrubs to further analyse variation of nest predation with predator assemblage and habitat quality for the predators. For both natural and artificial nests predation risk was higher in presence of squirrels. But predation risk varied within island categories. In presence of squirrels it was highest in stands with mature conifers where it fluctuated from year to year, in response to fluctuations in squirrel abundance. Vegetation cover around the nest had little effect on nest predation by squirrels. Where squirrels were absent, nest predation concentrated near predictable food sources for corvids, the main native predators, and increased with decreasing vegetation cover, suggesting that removal of the vegetation by deer increased the risk of predation by native avian nest predators that use visual cues. Predation risk in these forests therefore varies in space and time with predator composition and with quality of the habitat from the predators' perspective. This temporal and spatial variation in predation risk should promote trade-offs in the response of birds to nest predation, rather than fine-tuned adaptations to a given predation pattern.     

Foraging of multimammate mice, Mastomys natalensis, under different predation pressure: cover, patch-dependent decisions and density-dependent GUDs

Patch use under predation risk often results in a change of feeding behaviour in the prey animals. However, such changes only appear if the animals are able to assess under which predation pressure they live. We investigated patch use of Mastomys natalensis under different conditions of avian predation pressure.
In replicated maize field plots in Morogoro, Tanzania, avian predators were allowed under natural conditions (control), attracted with perches and nest boxes or kept out with nets. During four one‐week periods in late 1999, we measured rodent feeding decisions with the giving‐up density (GUD) method. Trays with known amounts of millet seeds in sand were placed in pairs, one of them under a cover, the other one in the open. M. natalensis mice were expected to give up sooner in the open trays than in those with cover. We hypothesised that M. natalensis mice could assess the ambient predation pressure leading to larger difference in GUD between covered and non‐covered trays in the plots where predators were attracted. We also made video recordings of the rodent activity at a pair of trays in each treatment. The GUD‐values were significantly lower for the covered trays but predation pressure did not affect this difference. The video observations showed that in the control and netted plots the animals visited trays equally frequently regardless of the cover, while the visits in the predator‐attracted plots occurred significantly more often in the covered trays. We conclude that M. natalensis can assess the ambient predation pressure and adapt its behaviour at a feeding patch. However, the variation in predation pressure in our experiment was not obvious from the GUD. Moreover, we found a strong relation between rodent density and GUD, which may mask variations in perceived predation pressure. Similar GUD values may be reached in different ways and we present models to investigate whether animals’ decision to forage at a food patch is only affected by the seed density at that patch, not by that at a neighbour patch.     

Variation in Note Composition of Chick‐a‐dee Calls is Associated with Signaler Flight in Carolina Chickadees,Poecile carolinensis

Chick‐a‐dee calls are used in a wide range of social contexts in Poecile (chickadee) species. These calls comprise a number of distinct note types. Earlier naturalistic observational studies suggested that the ‘C’ note type was used frequently in calls of Carolina chickadees (P. carolinensis) in the context of flight. We conducted three field studies with Carolina chickadees to test in more experimentally manipulative ways whether Chick‐a‐dee calls with more ‘C’ notes were associated with flight. The three studies differed in how they elicited flight behavior from chickadees, as well as in the likely arousal levels experienced by the birds. First, we captured chickadees and released them, recording any calls they produced in flight or when later perched after escape. Second, we approached chickadees that were foraging near the ground in field settings and recorded any calls they produced when perched compared to when they were in flight. Third, from a distant blind, we recorded chickadees flying to and from feeding stations, in which the closest perching substrate/cover was at least 2 m away from the feeding station, thus requiring flight. In all three studies, calls contained more ‘C’ notes when birds were in flight compared to when they were perched. This work expands our understanding of variation in note composition of chick‐a‐dee calls beyond the contexts of food and predator detection to the context of movement. Studies are now needed to test whether such variation in chick‐a‐dee calls brings about group cohesion.     

Diets of native and introduced tree squirrels in Washington

Competition for food resources can be a primary mechanism for displacement of native species by introduced species. Investigation of dietary partitioning between potential competitors and spatiotemporal variability in food resources can reveal contested food items and intensity of competition. Introduced eastern gray squirrels (Sciurus carolinensis) have been implicated as competitors with western gray squirrels (S. griseus) and Douglas’ squirrels (Tamiasciurus douglasii), but little is known about interactions among these species. We collected foraging observations and fecal pellets of sympatric gray and Douglas’ squirrels trapped and radio-tracked in western Washington, USA, from 2007 to 2012. We compared food resources consumed across species, seasons, and years to evaluate dietary overlap and the potential for competitive interactions. All squirrel species ate hypogeous fungi throughout all seasons; spores of several genera were present in 272 of 275 fecal samples and observed in all months. Rhizopogon, Geopora, and Melanogaster spp. occurred in most fecal pellets for all squirrels, but the diet of the Douglas’ squirrels had slightly lower richness of hypogeous fungi than gray squirrels. Although foraging observations suggest that eastern and western gray squirrels shared important dietary items such as acorns, strong differences in consumption of seeds of a common conifer may facilitate their coexistence. Our results suggest that dietary overlap among arboreal squirrels could lead to competitive interactions during periods of food scarcity, but subtle differences may be sufficient to permit long-term coexistence. Tree squirrels in Washington would likely benefit from forest management practices that promote or sustain robust crops of hypogeous fungi. © 2019 The Wildlife Society.     

Effects of Predation Risk and Food Availability on the Activity,Habitat Use,Feeding Behavior and Mating Behavior of a Pond Water Strider,Gerris marginatus (Hemiptera)

By experimentally manipulating predation risk and food availability, we explored the effects of these factors on refuge use, general activity, feeding activity, and mating activity of adults of the pond water strider Gerris marginatus. Predation risk (from adult backswimmers Notonecta undulata) profoundly influenced the behavior of G. marginatus, reducing mating activity, general activity in males and females, decreasing feeding activity in females and increasing the use of refuge by both sexes. Surprisingly, variations in food availability failed to produce significant effects on any of these behavior patterns. Risk x food interactions were also not statistically significant for any gerrid behavior. The significant responses to predators all changed in directions that should decrease the risk of predation for G. marginatus.     

艾虎对猎物和捕食者气味的反应

在室内条件下通过双通道选择实验确定艾虎(Mustelaeversmanni)在通过不同气味源的场所中的取食量以及利用时间和利用频次,由此分析捕食者气味和猎物气味对艾虎取食行为的影响。研究结果表明,在无任何气味源存在时,艾虎在对照气味和对照气味组这两个场所中的取食量、活动时间和活动频次基本一致。在猎物高原鼠兔(Ochotonacurzoniae)气味和对照气味组及高原鼢鼠(Myospalaxbaileyi)气味和对照气味组的两组实验中,艾虎在通过猎物气味的场所中的取食量、活动时间和活动频次与对照气味场所中比较也无明显的差异,但是在天敌动物赤狐(Vulpesvulpes)气味存在的条件下,艾虎明显减少在通过赤狐气味的场所中的取食量、活动时间和活动频次。同时,当间接捕食风险天敌动物赤狐气味存在时,艾虎明显增加了总的取食量,减少了在巢箱中居住的时间,而将更多的时间停留在无风险的场所中,这说明艾虎能够利用天敌动物赤狐的气味,采用增加取食量和减少高风险区域活动时间的策略来降低被捕食风险。     

Habitat use and feeding behavior of thirteen species of benthic stream fishes

Synopsis The densities, habitat use, and feeding behaviors of 13 fish species belonging to the benthic invertebrate-feeding guild were studied by snorkeling at three localities in the Little River of eastern Tennessee, U.S.A. Resource partitioning occurred by habitat, feeding behavior and time of activity. Differences were also found at the generic level.Cottus was a nocturnal feeder, whereasPercina andEtheostoma were, for the most part, diurnally active.Percina moved about rapidly and spent most of its time above the bottom. In contrast,Etheostoma varied considerably in the amount of time spent under cover, spent little time above the bottom, and exhibited low levels of swimming activity. Nearly all species sought cover at night, suggesting they may be particularly sensitive to predation at night. Species with small adult sizes (Etheostoma, Cottus andP. evides) were concentrated in shallow water habitats, whereas species with large adult sizes (Percina) were more abundant in deep water habitats. The habitat use data are consistent with the hypothesis that size-selective predation by centrarchid bass may cause smaller fish to avoid deep water areas. Large species should have a lower risk of predation due to their size and behavior.     

Is social coordination during escape flights a general phenomenon in birds?

Escape from predatory attack as a socially coordinated group is observed in many social animals, including birds, especially those in more open habitats where the group itself may be the only source of protection from an attacking predator. For many social birds, however, woody vegetative cover is the main refuge from attack, but such birds might nevertheless benefit from social coordination during escape flights to cover. Such benefits could reflect the confusion effect, selfish herd effect, or the simple dilution of risk. We examined this possibility of coordinated escape in mixed flocks of wintering passerellid sparrows (Passerellidae). These free-living birds fed in a patch of food flanked on opposite sides by two refuges composed of woody cover. Under such conditions, coordination in escape behavior should be expressed as a tendency to escape together as a group to the same cover location. Such behavior, however, was not the rule. During spontaneous flushes to cover, a group of escaping birds stayed together only when one cover location was clearly closer than the other. With cover equidistant from the food patch, escaping flocks tended to split about evenly between cover locations. Birds in close proximity prior to an escape flight did not show enhanced escape coordination, nor did those feeding at significant distances from protective cover. Evidence of escape coordination was observed in small groups (two–four birds), but even in such groups, flock splitting during escape was generally the rule. Flock splitting during attacks might reflect some sort of strategic decision-making process that lessens the risk of capture, but the most parsimonious explanation is that (all else equal) birds head for the nearest refuge, largely irrespective of the behavior of their flockmates. Our results thus provide little evidence of flock-wide social coordination during escape flights in cover-dependent birds.     

Mammalian predator scent, vegetation cover and tree seedling predation by meadow voles

Herbivores are thought to respond to the increased risk of attack by predators during foraging activities by concentrating feeding in safe habitats and by reducing feeding in the presence of predators. We tested these hypotheses by comparing tree seedling predation by meadow voles within large outdoor enclosures treated either with scent of large mammalian predators (red fox, bobcat, coyote) or a control scent (vinegar). In addition, we compared the distribution of voles in relation to naturally occurring variation in vegetation cover and the tendency of voles to attack tree seedlings planted in small patches with cover manipulation (intact, reduced or removed cover). Predator scent did not affect the rate or spatial distribution of tree seedling predation by voles, nor did it affect giving up densities (a surrogate of patch quitting harvest rate), survival rates, body size or habitat distribution of voles. In both predator scent and vinegar treatments voles preferred abundant vegetation providing good cover, which was also the site of almost all tree seedling predation. We conclude that large mammalian predator scent does not influence the perception by voles of the general safety of habitat, which is more strongly affected by the presence of cover.     

Predation risk and habitat complexity modify intermediate predator feeding rates and energetic efficiencies in a tri‐trophic system

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Effect of predation risk on selectivity in heteromyid rodents

Variations in predation risk affect the costs of foraging and may therefore warrant different foraging decisions. One class of models ("higher requisite profit") predicts that foragers should become more selective when predation risk increases, as low-profitability items that do not cover the increased costs are dropped from the diet. An alternative class of models ("reduced finickiness") predicts that foragers should become less selective when predation risk increases, because selectivity requires more extensive assessment and/or search behaviour, prolonging exposure to risk. We assessed the selectivity of foraging heteromyid rodents (Merriam's kangaroo rats, Dipodomys merriami, and pocket mice, Chaetodipus spp.) by comparing differences in "giving up densities" (GUD: the quantity of cryptic food left in a patch by animals for whom the diminishing marginal gains from foraging have dropped below the threshold for continued search) for foods of different value as a measure of selectivity in patches varying in predation risk. Data collected over two field seasons revealed that heteromyids were more selective when predation risk was highest; away from the protective cover of shrubs during the full moon. These findings support the predictions of higher requisite profit models.