Social information affects Canada goose alert and escape responses to vehicle approach: implications for animal-vehicle collisions.

BACKGROUND: Animal-vehicle collisions represent substantial sources of mortality for a variety of taxa and can pose hazards to property and human health. But there is comparatively little information available on escape responses by free-ranging animals to vehicle approach versus predators/humans. METHODS: We examined responses (alert distance and flight-initiation distance) of focal Canada geese (Branta canadensis maxima) to vehicle approach (15.6 m·s-1) in a semi-natural setting and given full opportunity to escape. We manipulated the direction of the vehicle approach (direct versus tangential) and availability of social information about the vehicle approach (companion group visually exposed or not to the vehicle). RESULTS: We found that both categorical factors interacted to affect alert and escape behaviors. Focal geese used mostly personal information to become alert to the vehicle under high risk scenarios (direct approach), but they combined personal and social information to become alert in low risk scenarios (tangential approach). Additionally, when social information was not available from the companion group, focal birds escaped at greater distances under direct compared to tangential approaches. However, when the companion group could see the vehicle approaching, focal birds escaped at similar distances irrespective of vehicle direction. Finally, geese showed a greater tendency to take flight when the vehicle approached directly, as opposed to a side step or walking away from the vehicle. CONCLUSIONS: We suggest that the perception of risk to vehicle approach (likely versus unlikely collision) is weighted by the availability of social information in the group; a phenomenon not described before in the context of animal-vehicle interactions. Notably, when social information is available, the effects of heightened risk associated with a direct approach might be reduced, leading to the animal delaying the escape, which could ultimately increase the chances of a collision. Also, information on a priori escape distances required for surviving a vehicle approach (based on species behavior and vehicle approach speeds) can inform planning, such as location of designated cover or safe areas. Future studies should assess how information from vehicle approach flows within a flock, including aspects of vehicle speed and size, metrics that affect escape decision-making.

Speed kills: ineffective avian escape responses to oncoming vehicles.

Animal-vehicle collisions cause high levels of vertebrate mortality worldwide, and what goes wrong when animals fail to escape and ultimately collide with vehicles is not well understood. We investigated alert and escape behaviours of captive brown-headed cowbirds (Molothrus ater) in response to virtual vehicle approaches of different sizes and at speeds ranging from 60 to 360 km h(-1). Alert and flight initiation distances remained similar across vehicle speeds, and accordingly, alert and flight initiation times decreased at higher vehicle speeds. Thus, avoidance behaviours in cowbirds appeared to be based on distance rather than time available for escape, particularly at 60-150 km h(-1); however, at higher speeds (more than or equal to 180 km h(-1)) no trend in response behaviour was discernible. As vehicle speed increased, cowbirds did not have enough time to assess the approaching vehicle, and cowbirds generally did not initiate flight with enough time to avoid collision when vehicle speed exceeded 120 km h(-1). Although potentially effective for evading predators, the decision-making process used by cowbirds in our study appears maladaptive in the context of avoiding fast-moving vehicles. Our methodological approach and findings provide a framework to assess how novel management strategies could affect escape rules, and the sensory and cognitive abilities animals use to avoid vehicle collisions.

Animal reactions to oncoming vehicles: a conceptual review.

Animal-vehicle collisions (AVCs) are a substantial problem in a human-dominated world, but little is known about what goes wrong, from the animal's perspective, when a collision occurs with an automobile, boat, or aircraft. Our goal is to provide insight into reactions of animals to oncoming vehicles when collisions might be imminent. Avoiding a collision requires successful vehicle detection, threat assessment, and evasive behaviour; failures can occur at any of these stages. Vehicle detection seems fairly straightforward in many cases, but depends critically on the sensory capabilities of a given species. Sensory mechanisms for detection of collisions (looming detectors) may be overwhelmed by vehicle speed. Distractions are a likely problem in vehicle detection, but have not been clearly demonstrated in any system beyond human pedestrians. Many animals likely perceive moving vehicles as non-threatening, and may generally be habituated to their presence. Slow or minimal threat assessment is thus a likely failure point in many AVCs, but this is not uniformly evident. Animals generally initiate evasive behaviour when a collision appears imminent, usually employing some aspect of native antipredator behaviour. Across taxa, animals exhibit a variety of behaviours when confronted with oncoming vehicles. Among marine mammals, right whales Eubalaena spp., manatees Trichechus spp., and dugongs Dugong dugon are fairly unresponsive to approaching vehicles, suggesting a problem in threat assessment. Others, such as dolphins Delphinidae, assess vehicle approach at distance. Little work has been conducted on the behavioural aspects of AVCs involving large mammals and automobiles, despite their prevalence. Available observations suggest that birds do not usually treat flying aircraft as a major threat, often allowing close approach before taking evasive action, as they might in response to natural predators. Inappropriate antipredator behaviour (often involving immobility) is a major source of AVCs in amphibians and terrestrial reptiles. Much behavioural work on AVCs remains to be done across a wide variety of taxa. Such work should provide broad phylogenetic generalizations regarding AVCs and insights into managing AVCs.

Effects of vehicle speed on flight initiation by Turkey vultures: implications for bird-vehicle collisions.

The avoidance of motorized vehicles is a common challenge for birds in the modern world. Birds appear to rely on antipredator behaviors to avoid vehicles, but modern vehicles (automobiles and aircraft) are faster than natural predators. Thus, birds may be relatively ill-equipped, in terms of sensory capabilities and behaviors, to avoid vehicles. We examined the idea that birds may be unable to accurately assess particularly high speeds of approaching vehicles, which could contribute to miscalculations in avoidance behaviors and ultimately cause collisions. We baited turkey vultures (Cathartes aura) to roads with animal carcasses and measured flight initiation distance and effective time-to-collision in response to a truck driving directly towards vultures from a starting distance of 1.13 km and at one of three speeds: 30, 60, or 90 kph (no vultures were struck). Flight initiation distance of vultures increased by a factor of 1.85 as speed increased from 30 to 90 kph. However, for 90-kph approaches there was no clear trend in flight initiation distance across replicates: birds appeared equally likely to initiate escape behavior at 40 m as at 220 m. Time-to-collision decreased by a factor of 0.62 with approach speeds from 30 to 90 kph. Also, at 90 kph, four vehicle approaches (17%) resulted in near collisions with vultures (time-to-collision ≤ 1.7 s), compared to none during 60 kph approaches and one during 30 kph approaches (4%). Our findings suggest that antipredator behaviors in turkey vultures, particularly stimulus processing and response, might not be well tuned to vehicles approaching at speeds ≥ 90 kph. The possible inability of turkey vultures to react appropriately to high-speed vehicles could be common among birds, and might represent an important determinant of bird-vehicle collisions.

Wintering birds avoid warm sunshine: predation and the costs of foraging in sunlight.

Wintering birds can gain significant thermal benefits by foraging in direct sunlight. However, exposure to bright sunlight might make birds easier to detect by predators and may also cause visual glare that can reduce a bird's ability to monitor the environment. Thus, birds likely experience a trade-off between the thermal benefits and predation-related costs of foraging in direct sunlight. To examine this possible thermoregulation-predation trade-off, we monitored the behavior of mixed-species flocks of wintering emberizid sparrows foraging in alternating strips of sunlight and shade. On average, these sparrows routinely preferred to forage in the shade, despite midday air temperatures as much as 30 °C below their thermoneutral zone. This preference for shade was strongest at relatively high temperatures when the thermal benefits of foraging in sunlight were reduced, suggesting a thermoregulation-predation trade-off. Glare could be reduced if birds faced away from the sun while feeding in direct sunlight, but we found that foraging birds tended to face southward (the direction of the sun). We speculate that other factors, such as the likely direction of predator approach, may explain this southerly orientation, particularly if predators use solar glare to their advantage during an attack. This interpretation is supported by the fact that birds had the weakest southerly orientation on cloudy days. Wintering birds may generally avoid foraging in direct sunlight to minimize their risk of predation. However, given the thermal benefits of sunshine, such birds may benefit from foraging in habitats that provide a mosaic of sunlit and shaded microhabitats.

Nocturnal hypothermia impairs flight ability in birds: a cost of being cool.

Many birds use regulated drops in night-time body temperature (Tb) to conserve energy critical to winter survival. However, a significant degree of hypothermia may limit a bird's ability to respond to predatory attack. Despite this likely energy-predation trade-off, the behavioural costs of avian hypothermia have yet to be examined. We thus monitored the nocturnal hypothermia of mourning doves (Zenaida macroura) in a laboratory setting in response to food deprivation. Nocturnal flight tests were used to quantify the flight ability of hypothermic doves. Many hypothermic doves (39% of tests) could not fly while carrying a small weight, but could do so after quickly warming up to typical daytime Tb. Doves that were unable to fly during their first test were more hypothermic than those that could fly, with average Tb reductions of 5.3°C and 3.3°C, respectively, but there was no overall indication of a threshold Tb reduction beyond which doves were consistently incapable of flight. These results suggest that energy-saving hypothermia interferes with avian antipredator behaviour via a reduction in flight ability, likely leading to a trade-off between energy-saving hypothermia and the risk of predation.

Do predators influence the behaviour of bats?

Many aspects of animal behaviour are affected by real-time changes in the risk of predation. This conclusion holds for virtually all taxa and ecological systems studied, but does it hold for bats? Bats are poorly represented in the literature on anti-predator behaviour, which may reflect a lack of nocturnal predators specialized on bats. If bats actually experience a world with minimal anti-predator concerns, then they will provide a unique contrast within the realm of vertebrate ecology. Alternatively, such predator-driven behaviour in bats may not yet be fully understood, given the difficulties in working with these highly mobile and nocturnal animals. We provide a wide-ranging exploration of these issues in bat behaviour. We first cover the basic predator-prey information available on bats, both on potential predators and the ways in which bats might perceive predators and respond to attacks. We then cover work relevant to key aspects of bat behaviour, such as choice of daytime roosts, the nature of sleep and torpor, evening roost departures, moonlight avoidance, landscape-related movement patterns, and habitat selection. Overall, the evidence in favour of a strong influence of predators on bat behaviour is equivocal, with the picture clouded by contradictory results and a lack of information on potential predators and the perception of risk by bats. It seems clear that day-active bats run a considerable risk of being killed by diurnal raptors, which are able to capture bats with relative ease. Thus, bats taking advantage of a pulse of insects just prior to sunset are likely taking risks to gain much-needed energy. Further, the choice of daytime roosts by bats is probably strongly influenced by roost safety. Few studies, however, have directly addressed either of these topics. As a group, insectivorous temperate-zone bats show no clear tendency to avoid apparently risky situations, such as activity on moonlit nights. However, some observations are consistent with the idea that predation risk affects choice of movement paths and feeding areas by temperate-zone bats, as well as the timing of roost departures. The behaviour of tropical bats, on the other hand, seems more generally influenced by predators; this is especially true for tropical nectarivores and frugivores, but also for insectivorous bats. Presumably there are more serious predators on bats in the tropics (e.g. specialized raptors or carnivorous bats), but the identity of these predators is unclear. More information is needed to assess fully the influence of predators on bat behaviour. There is much need for work on the ways in which bats perceive predators via auditory, visual, and olfactory cues, and whether bats have some knowledge of the risks posed by different predators. Also needed is information on how predators attack bats and how bats react to attacking predators. Difficult to obtain, but of critical value, will be information on the nature of the predation risk experienced by bats while away from roosts and during the full darkness of night.

Sleep locally, act globally.

In most animals, sleep is considered a global brain and behavioral state. However, recent intracortical recordings have shown that aspects of non-rapid eye movement (NREM) sleep and wakefulness can occur simultaneously in different parts of the cortex in mammals, including humans. Paradoxically, however, NREM sleep still manifests as a global behavioral shutdown. In this review, the authors examine this paradox from an evolutionary perspective. On the basis of strategic modeling, they suggest that in animals with brains composed of heavily interconnected and functionally interdependent units, a global regulator of sleep maintains the behavioral shutdown that defines sleep and thereby ensures that local use-dependent functions are performed in a safe and efficient manner. This novel perspective has implications for understanding deficits in human cognitive performance resulting from sleep deprivation, sleep disorders such as sleepwalking, changes in consciousness that occur during sleep, and the function of sleep itself.

Testing the terrain hypothesis: Canada geese see their world laterally and obliquely.

The distribution of ganglion cells in the retina determines the specific regions of the visual field with high visual acuity, and thus reflects the perception of a species' visual environment. The terrain hypothesis proposes that animals living in open areas should have a horizontal visual streak across the retina with high ganglion cell density to increase visual acuity along the horizon. We tested this hypothesis in Canada geese (Branta canadensis) by assessing retinal topography, visual field configuration, and scanning behavior. We found that geese have an oblique rather than a horizontal visual streak across the retina: from a dorsal-nasal to a ventral-temporal position. Geese showed narrow blind areas, which increased the range of their lateral vision, and a relatively large degree of eye movement. Canada geese have relatively wide binocular fields and can see their bill tips. Goose head movement rates were low compared to species with a single fovea, and head movement rates increased in visually obstructed habitats. Canada geese have high acuity across their retina, which would allow them to simultaneously scan the ground and the sky when the head is up and parallel to the ground, as well as align the visual streak with the horizon when the head is tilted downwards. Their visual streak, along with their large eye size, may reduce the need for large amplitude head movements during vigilance bouts in visually unobstructed habitats. Overall, the visual system of geese combines features related to the detection of predators/conspecifics in open areas (visual streak, large lateral field, reduced head movements) as well as visual specializations that would allow for monitoring both the ground and sky (oblique streak) and for extracting and handling of food items (wide binocular fields, visualization of the bill tip).

Mothers forewarn offspring about predators: a transgenerational maternal effect on behavior.

Predation risk has long been known to exert a strong influence on behavior, but no study to date has determined whether predators influence offspring antipredator behavior via maternal effects. Here, we provide a unique example of a transgenerational maternal effect in antipredator behavior that takes the form of a "warning" about predators that female fall field crickets Gryllus pennsylvanicus transmit to their offspring. Specifically, the offspring of gravid crickets exposed to a wolf spider Hogna helluo exhibit greater antipredator immobility in response to Hogna chemical cues than do offspring of nonexposed females. These "forewarned" crickets exhibit greater survival in the presence of Hogna than do those not forewarned. Accordingly, gravid crickets from areas with significant Hogna populations produce offspring that are more responsive to Hogna cues than do those from nearby Hogna-free areas. Such transgenerational maternal effects may be more common than currently realized.

Stress and aversive learning in a wild vertebrate: the role of corticosterone in mediating escape from a novel stressor.

Elevated plasma corticosterone during stressful events is linked to rapid changes in behavior in vertebrates and can mediate learning and memory consolidation. We tested the importance of acute corticosterone elevation in aversive learning of a novel stressor by wild male eastern fence lizards (Sceloporus undulatus). We found that inhibiting corticosterone elevation (using metyrapone, a corticosterone synthesis blocker) during an encounter with a novel attacker impaired immediate escape responses and limited learning and recall during future encounters. In the wild and in outdoor enclosures, lizards whose acute corticosterone response was blocked by an earlier metyrapone injection did not alter their escape behavior during repeated encounters with the attacker. Control-injected (unblocked) lizards, however, progressively increased flight initiation distance and decreased hiding duration during subsequent encounters. Aversive responses were also initially higher for control lizards exposed to a higher intensity first attack. Further, we demonstrate a role of corticosterone elevation in recollection, since unblocked lizards had heightened antipredator responses 24-28 h later. Exogenously restoring corticosterone levels in metyrapone-injected lizards maintained aversive behaviors and learning at control (unblocked) levels. We suggest that the corticosterone mediation of antipredator behaviors and aversive learning is a critical and general mechanism for the behavioral flexibility of vertebrate prey.

Predators and the breeding bird: behavioral and reproductive flexibility under the risk of predation.

A growing body of work suggests that breeding birds have a significant capacity to assess and respond, over ecological time, to changes in the risk of predation to both themselves and their eggs or nestlings. This review investigates the nature of this flexibility in the face of predation from both behavioural and reproductive perspectives, and also explores several directions for future research. Most available work addresses different aspects of nest predation. A substantial change in breeding location is perhaps the best documented response to nest predation, but such changes are not always observed and not necessarily the best strategy. Changes in nesting microhabitat (to more concealed locations) following predation are known to occur. Surprisingly little work addresses the proactive avoidance of areas with many nest predators, but such avoidance is probably widespread. Individual birds could conceivably adopt anti-predator strategies based on the nest predators actually present in an area, but such effects have yet to be demonstrated. In fact, the ways in which birds assess the risk of nest predation is unclear. Nest defence in birds has historically received much attention, but little is known about how it interacts with other aspects of decision-making by parents. Other studies concentrate on predation risk to adults. Some findings suggest that risk to adults themselves influences territory location, especially relative to raptor nests. An almost completely unexplored area concerns the sorts of social protection from predators that might exist during the breeding season. Flocking typical of the non-breeding season appears unusual while breeding, but a mated pair may sometimes act as a "flock of two". Opportunistic heterospecific sociality may exist, with heterospecific protector species associations more prevalent than currently appreciated. The dynamics of singing during the breeding season may also respond to variation in predation risk, but empirical research on this subject is limited. Furthermore, a few theoretical and empirical studies suggest that changes in predation risk also influence the behaviour of lekking males. The major influence of predators on avian life histories is undoubtedly expressed at a broad phylogenetic scale, but several studies hint at much flexibility on an ecological time scale. Some species may forgo breeding completely if the risk of nest predation is too high, and a few studies document smaller clutch sizes in response to an increase in nest predation. Recent evidence suggests that a female may produce smaller eggs rather than smaller clutches following an increase in nest predation risk. Such an increase may also influence decisions about intraspecific brood parasitism. There are no clear examples of changes in clutch/egg size with changes in risk experienced by adults, but parental responses to predators have clear consequences for offspring fitness. Changes in risk to adults may also influence body mass changes across the breeding season, although research here is sparse. The topics highlighted herein are all in need more empirical attention, and more experimental field work whenever feasible.

History and future of comparative analyses in sleep research.

The comparative methods of evolutionary biology are a useful tool for investigating the functions of sleep. These techniques can help determine whether experimental results, derived from a single or few species, apply broadly across a specified group of animals. In this way, comparative analysis is a powerful complement to experimentation. The variation in the time mammalian species spend asleep has been most amenable for use with this approach, given the large number of mammals for which sleep data exist. Here, it is assumed that interspecific variation in the time spent asleep reflects underlying differences in the need for sleep. If true, then significant predictors of sleep times should provide insight into the function of sleep. Many such analyses have sought the evolutionary determinants of mammalian sleep by relating the time spent in the two basic states of sleep, rapid eye movement (REM) and non-REM sleep, to constitutive variables thought to be functionally related to sleep. However, the early analyses had several methodological problems, and recent re-analyses have overturned some widely accepted relationships, such as the idea that species with higher metabolic rates engage in more sleep. These more recent studies also provide evolutionarily broad support for a neurophysiological role for REM sleep. Furthermore, results from comparative analyses suggest that animals are particularly vulnerable to predation during REM sleep, a finding that lends further support to the notion that REM sleep must serve an important function. Here, we review the methodology and results of quantitative comparative studies of sleep. We highlight important developments in our understanding of the evolutionary determinants of sleep and emphasize relationships that address prevailing hypotheses for the functions of sleep. Lastly, we outline a possible future for comparative analyses, focusing on work in non-mammalian groups, the use of more physiologically meaningful variables, and electrophysiological sleep studies conducted in the wild.

Acute corticosterone elevation enhances antipredator behaviors in male tree lizard morphs.

In response to stressful events, most vertebrates rapidly elevate plasma glucocorticoid levels. Corticosterone release stimulates physiological and behavioral responses that can promote survival while suppressing behaviors that are not crucial to immediate survival. Corticosterone also has preparatory effects for subsequent stressors. Using male tree lizards (Urosaurus ornatus), we tested our prediction that elevated corticosterone is important for mediating and enhancing antipredator behaviors. Male tree lizards express developmentally fixed polymorphisms that are mediated by early organizational actions of steroid hormones, and thus we also tested the hypothesis that morph-specific differences in antipredator behaviors of adults are independent of circulating corticosterone levels. Plasma corticosterone levels were elevated exogenously for 12-16 h using non-invasive dermal patches, and we then compared the behavioral responses of these corticosterone-patched males to control-patched males during a simulated encounter with a caged predator (collared lizard, Crotaphytus nebrius) in outdoor enclosures. Elevating corticosterone did not alter the antipredator behavioral repertoire of each male morph, but did enhance their responses during the predator encounter: all corticosterone-patched males responded more quickly, hid longer, and displayed more toward the predator than control-patched males. With the corticosterone patch, the non-territorial and wary orange morph was still behaviorally the most wary morph, responding more quickly and hiding longer than either the bolder orange-blue or mottled morphs. Smaller males were generally warier than larger males, regardless of the endocrine treatment or color morph type. In sum, elevated circulating corticosterone enhances antipredator responses for all male tree lizard morphs, without altering morph-specific or size-specific differences in their behavioral responses.

Phylogenetics and the correlates of mammalian sleep: a reappraisal.

The correlates of mammalian sleep have been investigated previously in at least eight comparative studies in an effort to illuminate the functions of sleep. However, all of these univariate analyses treated each species, or taxonomic Family, as a statistically independent unit, which is invalid due to the phylogenetic relationships among species. Here, we reassess these influential correlates of mammalian sleep using the formal phylogenetic framework of independent contrasts. After controlling for phylogeny using this procedure, the interpretation of many of the correlates changed. For instance, and contrary to previous studies, we found interspecific support for a neurophysiological role for rapid-eye-movement sleep, such as memory consolidation. Also in contrast to previous studies, we did not find comparative support for an energy conservation function for slow-wave sleep. Thus, the incorporation of a phylogenetic control into comparative analyses of sleep yields meaningful differences that affect our understanding of why we sleep.

Predator-induced plasticity in sleep architecture in wild-caught Norway rats (Rattus norvegicus).

Sleep is a prominent behaviour in the lives of animals, but the unresponsiveness that characterizes sleep makes it dangerous. Mammalian sleep is composed of two neurophysiological states: slow wave sleep (SWS) and rapid-eye-movement (REM) sleep. Given that the intensity of stimuli required to induce an arousal to wakefulness is highest during deep SWS or REM sleep, mammals may be most vulnerable during these states. If true, then animals should selectively reduce deep SWS and REM sleep following an increase in the risk of predation. To test this prediction, we simulated a predatory encounter with 10 wild-caught Norway rats (Rattus norvegicus), which are perhaps more likely to exhibit natural anti-predator responses than laboratory strains. Immediately following the encounter, rats spent more time awake and less time in SWS and REM sleep. The reduction of SWS was due to the shorter duration of SWS episodes, whereas the reduction of REM sleep was due to a lower number of REM sleep episodes. The onset of SWS and REM sleep was delayed post-encounter by about 20 and 100 min, respectively. The reduction of REM sleep was disproportionately large during the first quarter of the sleep phase, and slow wave activity (SWA) (0.5-4.5 Hz power density) was lower during the first 10 min of SWS post-encounter. An increase in SWA and REM sleep was observed later in the sleep phase, which may reflect sleep homeostasis. These results suggest that aspects of sleep architecture can be adjusted to the prevailing risk of predation.

Use of prey hotspots by an avian predator: purposeful unpredictability?

The use of space by predators in relation to their prey is a poorly understood aspect of predator-prey interactions. Classic theory suggests that predators should focus their efforts on areas of abundant prey, that is, prey hotspots, whereas game-theoretical models of predator and prey movement suggest that the distribution of predators should match that of their prey's resources. If, however, prey are spatially anchored to one location and these prey have particularly strong antipredator responses that make them difficult to capture with frequent attacks, then predators may be forced to adopt alternative movement strategies to hunt behaviorally responsive prey. We examined the movement patterns of bird-eating sharp-shinned hawks (Accipiter striatus) in an attempt to shed light on hotspot use by predators. Our results suggest that these hawks do not focus on prey hotspots such as bird feeders but instead maintain much spatial and temporal unpredictability in their movements. Hawks seldom revisited the same area, and the few frequently used areas were revisited in a manner consistent with unpredictable returns, giving prey little additional information about risk.

The predatory behavior of wintering Accipiter hawks: temporal patterns in activity of predators and prey.

Studies focused on how prey trade-off predation and starvation risk are prevalent in behavioral ecology. However, our current understanding of these trade-offs is limited in one key respect: we know little about the behavior of predators. In this study, we provide some of the first detailed information on temporal patterns in the daily hunting behavior of bird-eating Accipiter hawks and relate that to their prey. During the winters of 1999-2004, twenty-one sharp-shinned hawks (A. striatus) and ten Cooper's hawks (A. cooperii) were intensively radio tracked in rural and urban habitats in western Indiana, USA. Cooper's hawks left roost before sunrise and usually returned to roost around sunset, while sharp-shinned hawks left roost at sunrise or later and returned to roost well before sunset. An overall measure of Cooper's-hawk-induced risk (a composite variable of attack rate and activity patterns) generally reflected the timing of prey activity, with peaks occurring around sunrise and sunset. In contrast, risk induced by the smaller sharp-shinned hawk did not strongly reflect the activity of their prey. Specifically, an early morning peak in prey activity did not correspond to a period with intense hawk activity. The lack of early morning hunting by sharp-shinned hawks may reflect the high risk of owl-induced predation experienced by these hawks. The net effect of this intraguild predation may be to "free" small birds from much hawk-induced predation risk prior to sunrise. This realization presents an alternative to energetics as an explanation for the early morning peak in small bird activity during the winter.