Memory for food caches: not just for retrieval

Many animals use hoarding as a long-term strategy to ensurea food supply at times of shortage. Hoarders employ strategiesthat enhance their ability to relocate caches such as rememberingwhere caches are located. Long-term scatterhoarders, whose cacheshave potentially high pilferage rates, should also hoard ina way to reduce potential cache pilferers' ability to find caches.Previous studies have demonstrated that this could be achievedby hyperdispersing caches to reduce the foraging efficiencyof pilferers. This study investigates whether coal tits (Parusater) indeed place their caches away from existing ones. Inour experiment, birds hoarded food in 3 conditions: when cachesfrom a previous storage session were still present, when cachesfrom a previous storage session were not present anymore becausethe bird had retrieved them, and when caches from a previousstorage session had been removed by the experimenter. We showthat coal tits hoard away from existing caches and that theydo not use cues from extant caches to do this. This evidenceis consistent with the use of memory for the locations of previouscaches when deciding where to place new caches. This findinghas important implications for our understanding of the selectivepressures that have shaped spatial memory in food-hoarding birds.     

Effects of demanding foraging conditions on cache retrival accuracy in food-caching mountain chickadees (Poecile gambeli)

Birds rely, at least in part, on spatial memory for recovering previously hidden caches but accurate cache recovery may be more critical for birds that forage in harsh conditions where the food supply is limited and unpredictable. Failure to find caches in these conditions may potentially result in death from starvation. In order to test this hypothesis we compared the cache recovery behaviour of 24 wild-caught mountain chickadees (Poecile gambeli), half of which were maintained on a limited and unpredictable food supply while the rest were maintained on an ad libitum food supply for 60 days. We then tested their cache retrieval accuracy by allowing birds from both groups to cache seeds in the experimental room and recover them 5 hours later. Our results showed that birds maintained on a limited and unpredictable food supply made significantly fewer visits to non-cache sites when recovering their caches compared to birds maintained on ad libitum food. We found the same difference in performance in two versions of a one-trial associative learning task in which the birds had to rely on memory to find previously encountered hidden food. In a non-spatial memory version of the task, in which the baited feeder was clearly marked, there were no significant differences between the two groups. We therefore concluded that the two groups differed in their efficiency at cache retrieval. We suggest that this difference is more likely to be attributable to a difference in memory (encoding or recall) than to a difference in their motivation to search for hidden food, although the possibility of some motivational differences still exists. Overall, our results suggest that demanding foraging conditions favour more accurate cache retrieval in food-caching birds.     

Landmark learning and visuo-spatial memories in gerbils

The aim of this study is to understand what a rodent (Meriones unguiculatus) learns about the geometrical relations between a goal and nearby visual landmarks and how it uses this information to reach a goal. Gerbils were trained to find sunflower seeds on the floor of a light-tight, black painted room illuminated by a single light bulb hung from the ceiling. The position of the seed on the floor was specified by an array of one or more landmarks. Once training was complete, we recorded where the gerbils searched when landmarks were present but the seed was absent. In such tests, gerbils were confronted either with the array of landmarks to which they were accustomed or with a transformation of this array. Animals searched in the appropriate spot when trained to find seeds placed in a constant direction and at a constant distance from a single cylindrical landmark. Since gerbils look in one spot and not in a circle centred on the landmark, the direction between landmark and goal must be supplied by cues external to the landmark array. Distance, on the other hand, must be measured with respect to the landmark. Tests in which the size of the landmark was altered from that used in training suggest that distance is not learned solely in terms of the apparent size of the landmark as seen from the goal. Gerbils can still reach a goal defined by an array of landmarks when the room light is extinguished during their approach. This ability implies that they have already planned a trajectory to the goal before the room is darkened. In order to compute such a trajectory, their internal representation of landmarks and goal needs to contain information about the distances and bearings between landmarks and goal. For planning trajectories, each landmark of an array can be used separately from the others. Gerbils trained to a goal specified by an array of several landmarks were tested with one or more of the landmarks removed or with the array expanded. They then searched as though they had computed an independent trajectory for each landmark. For instance, gerbils trained with an array of two landmarks were tested with the distance between two landmarks doubled. The animals then searched for seeds in two positions, which were at the correct distance and in the right direction from each landmark.(ABSTRACT TRUNCATED AT 400 WORDS)     

What scatter-hoarding animals have taught us about small-scale navigation

Many animals use cues for small-scale navigation, including beacons, landmarks, compasses and geometric properties. Scatter-hoarding animals are a unique system to study small-scale navigation. They have to remember and relocate many individual spatial locations, be fairly accurate in their searching and have to remember these locations for long stretches of time. In this article, we review what is known about cue use in both scatter-hoarding birds and rodents. We discuss the importance of local versus global cues, the encoding of bearings and geometric rules, the use of external compasses such as the Sun and the influence of the shape of experimental enclosures in relocating caches or hidden food. Scatter-hoarding animals are highly flexible in how and what they encode. There also appear to be differences in what scatter-hoarding birds and rodents encode, as well as what scatter-hoarding animals in general encode compared with other animals. Areas for future research with scatter-hoarding animals are discussed in light of what is currently known.     

Involvement of the sun and the magnetic compass of domestic fowl in its spatial orientation

Domestic chicks are able to find a food goal at different times of day, with the sun as the only consistent visual cue. This suggests that domestic chickens may use the sun as a time-compensated compass, rather than as a beacon. An alternative explanation is that the birds might use the earth's magnetic field. In this study, we investigated the role of the sun compass in a spatial orientation task using a clock-shift procedure. Furthermore, we investigated whether domestic chickens use magnetic compass information when tested under sunny conditions.Ten ISA Brown chicks were housed in outdoor pens. A separate test arena comprised an open-topped, opaque-sided, wooden octagonal maze. Eight goal boxes with food pots were attached one to each of the arena sides. A barrier inside each goal box prevented the birds from seeing the food pot before entering. After habituation, we tested in five daily 5-min trials whether chicks were able to find food in an systematically allocated goal direction. We controlled for the use of olfactory cues and intra-maze cues. No external landmarks were visible. All tests were done under sunny conditions. Circular statistics showed that nine chicks significantly oriented goalwards using the sun as the only consistent visual cue during directional testing. Next, these nine chicks were subjected to a clock-shift procedure to test for the role of sun-compass information. The chicks were housed indoors for 6 days on a light-schedule that was 6 h ahead of the natural light–dark schedule. After clock-shifting, the birds were tested again and all birds except one were disrupted in their goalward orientation. For the second experiment, six birds were re-trained and fitted with a tiny, powerful magnet on the head to disrupt their magnetic sense. The magnets did not affect the chicks’ goalward orientation.In conclusion, although the strongest prediction of the sun-compass hypothesis (significant re-orientation after clock-shifting) was neither confirmed nor refuted, our results suggest that domestic chicks use the sun as a compass rather than as a beacon. These findings suggest that hens housed indoors in large non-cage systems may experience difficulties in orientation if adequate alternative cues are unavailable. Further research should elucidate how hens kept in non-cage systems orient in space in relation to available resources.     

Some psychophysics of the pigeon's use of landmarks

1. Three pigeons (Columba livia) were trained to find hidden food in a sunken well (3.3 cm in diameter) at a constant place within an (160 cm x 160 cm) experimental box (Fig. 1). After learning the location, the animals were tested occasionally with the well and food absent. Landmarks in the experimental box might be transformed on such tests. 2. Changing the height or width of a nearby landmark had no systematic influence on the position of peak search. Translating a nearby landmark, however, led to a shift in peak search position. All three birds then searched most somewhere between the original goal location, as defined by the unmoved landmarks, and the goal location as defined by the shifted landmark. Within a limited range of landmark shift, the peak shift as a function of landmark shift is linear (Fig. 3). 3. To explain the data (Fig. 7), the pigeon records at the location of the goal the algebraic vectors from a number of landmarks to the goal. These vectors have both a direction and a distance component. When searching for the goal again in the experimental box, it computes independently for each landmark a navigation vector. This is arrived at by vector-adding the algebraic vector from the bird's current position to the landmark in question, supplied by perception, to the corresponding landmark-goal vector in its record. The pigeon moves in the direction and distance specified by a weighted average of the independently calculated navigation vectors. For positive vector weights, vector geometry guarantees that the bird would search somewhere between the original goal and the goal according to the shifted landmark. The extent to which it shifts toward the shifted goal reflects the vector weight given to the shifted landmark.     

Use of Landmarks in Three Species of Food-storing Corvids

The use of landmarks by Clark's nutcrackers (Nucifraga columbiana), pinyon jays (Gymnorbinus cyanocepbalus), and Western scrub jays (Apbelocoma coerulescens) was investigated in three experiments. In the first experiment, birds were trained to find a hidden food reward beneath wood chips near a plastic landmark located 20 cm from one edge of a stationary tray. During unrewarded test trials, all species displaced their searching in response to movement of the landmark in both parallel and perpendicular directions; however, searching was displaced significantly more in the parallel direction. In a second experiment, the landmark was removed with the tray stationary or shifted. When the tray was shifted, all species responded by searching relative to the shifted tray. This indicates that the global (room) cues were not being used as salient landmarks. In the third experiment, the tray was placed in different positions for each trial, making global cues unreliable. As in experiment 1, all species displaced their searching in response to parallel and perpendicular movement of the landmark, with movement being similar in both directions. These results indicate no species differences on this landmark-use task.     

Diplochory in western chokecherry: you can��t judge a fruit by its mesocarp

Western chokecherry (Prunus virginiana var. demissa, Rosaceae) is dispersed by frugivorous birds and carnivores, but it has large seeds that are potentially attractive to rodents that could act as seed predators and dispersers. Here, we quantify the benefits of primary dispersal by birds and secondary dispersal by scatter-hoarding rodents. In the fall, avian frugivores (mostly American robins, Turdus migratorius, and cedar waxwings, Bombycilla cedrorum) consumed 87% of the fruit crop and dispersed 67% of the fruit crop away from parent plants. Rodents removed 89% of seeds that simulated bird-dispersed seed rain from transects in riparian zones and 58% from transects in upland habitats. Rodents scatter-hoarded 91.6% of the seeds they removed, burying most in small caches (two to eight seeds) 8?C25?mm deep. About 39% of the seeds in spring caches produced seedlings. Inside rodent-proof exclosures, 52.1% of seeds buried to simulate rodent caches produced seedlings, 29.7% of which were still alive after 1?year. In contrast, only 3.8% of seeds placed on the soil surface, simulating dispersal by avian frugivores, produced seedlings. Seed dispersal by frugivorous birds likely contributes to colonization of unoccupied habitat through long-range dispersal and to escape from distance-dependent seed mortality near the parent plant. Despite seed losses, rodents offer short-range seed dispersal and bury seeds in more favorable sites for germination, improving seedling emergence and establishment. The combined mechanisms of seed dispersal significantly enhanced chokecherry seedling recruitment by providing more dispersal-related benefits than either frugivorous bird or scatter-hoarding rodents could provide alone.     

贮藏点深度、大小及基质含水量对花鼠找寻红松种子的影响

2012年8-10月,在黑龙江省伊春市带岭林业局东方红林场,通过人工围栏控制实验,研究了贮藏点深度、大小及基质含水量对花鼠找寻红松种子的影响。发现：(1) 埋藏深度显著影响花鼠对贮藏点的找寻率,埋藏深度为 1 cm 和 2 cm 的找寻率显著高于 4 cm 和 6 cm。(2) 埋藏点大小对花鼠发现种子也有显著影响,埋藏点越大,花鼠发现贮藏点的比例越高。(3) 高的基质含水量利于花鼠找寻贮藏点。结果表明,围栏条件下食物贮藏点深度、大小及基质含水量的改变能显著影响花鼠对贮藏点的找寻。     

Bolus recovery by gray jays: an experimental analysis

Gray jays (Perisoreus canadensis) typically store food boli in various sites on conifers. In a laboratory setting we determined whether gray jays recover stored boli by means of olfaction, trial-and-error search or spatial memory. Using an artificial tree with 52 possible caching sites, caching and/or recovery trials were performed with five captive gray jays for the following experiments: (1) no extra visual cues on tree; (2) extra visual cues (pine foliage) attached to tree; (3) pungent-smelling food hidden by observer; (4) one bird allowed to cache food but caches recovered by a second bird; (5) one bird allowed to observe another bird cache food and later permitted to recover those caches. Results supported the memory hypothesis, but cache site preferences were apparent for individual birds. To control for this, an additional experiment (6), in which cache site access was limited by the investigators, was conducted with two new birds. These results also indicated that gray jays use spatial memory to recover stored boli.     

Navigational ability in the domestic fowl (Gallus gallus domesticus)

Little is known about the navigational abilities of domestic fowl. The question of how chickens represent and orient in space becomes relevant when they are kept in non-cage systems. Since the sun is known to be the dominant spatial organiser in other diurnal bird species, we started our investigation of the chicken’s spatial abilities by subjecting them to a food-searching task with the sun as the only consistent visual cue. In an additional experiment we tried to rule out the use of auditory cues in finding a food reward.

Eight ISA Brown chicks were housed in outdoor pens. A separate test arena comprised an open-topped, opaque-sided wooden octagon (2 m wide and 1.5 m high). Eight goal boxes with food pots were attached to each of the arena sides; a wooden barrier inside each goal box prevented the birds from seeing the food pot before entering. After habituation we tested during five daily 5 min trials whether the chicks were able to find food in a systematically allocated goal direction. Food residue in every foot pot controlled for the use of olfactory cues and no external landmark cues were visible. Every day each box was unpredictably moved to a randomly assigned side of the arena and the side to face north was also randomly allocated, to prevent the chicks from using cues other than the sun’s position. Circular statistics were used to determine whether birds moved in a non-random direction and if so, if they significantly oriented goalwards. The results showed that seven of the eight birds moved significantly in the goal direction. It seems likely that the chicks used the sun to orient. Due to weather constraints only four chicks received the same treatment on a new location, to rule out the use of auditory cues. Two of these four chicks significantly moved in the goal direction.

The results from our experiments show that domestic chicks use spatial memory to orient towards a hidden goal. Moreover, their orientation is most likely to be based on sun cues opening up the possibility that the sun compass may dominate even in this ancestrally predominantly ground-living forest bird.     

Management of fat reserves and food caches in tufted titmice (Parus bicolor) in relation to unpredictable food supply

In the temperate zone, permanent-resident birds and mammalsthat do not hibernate must survive harsh winter conditions oflow ambient temperature, long nights, and reduced food levels.To understand the energy management strategy of food-hoardingbirds, it has been hypothesized that such birds respond to increasedstarvation risk by increasing the number of their hoards ratherthan by increasing their fat reserves and that they cache earlyin the day and retrieve their caches later to achieve fat reservesnecessary to survive the night We tested these hypotheses byobserving the responses in captivity of a caching bird, thetufted titmouse (Parus bicolor), to the combined influencesof reduced predictability of food and naturally occurring ambienttemperature and photoperiod. When the food supply was unpredictable,birds significantly increased both internal fat reserves atdusk and external food caches. Initially leaner birds tendedto increase their fat reserves to a greater extent and initiallyfatter birds tended to cache more food and to fly significantlyless. Half the birds also increased their dawn and mean dailybody mass. All birds tended to forage, gain body mass, and cachefood at significantly lower rates in the morning and at significantlyhigher rates in the evening. Cache retrieval showed the oppositetrend, with birds retrieving most of their caches in the morning.Our results do not support the hypothesis that caching birdsincrease caching rate but not body mass under an unpredictablefood regime. Instead fat reserves and food caches are both importantcomplementary sources of energy in food-hoarding birds. Energymanagement by wintering birds occurs in response to a numberof biotic and abiotic factors acting simultaneously; thus futuremodels must incorporate independent variables in addition tothe state of the food supply and time of day     

An experimental analysis of cache recovery in Clark's nutcracker

Five hypotheses of cache recovery behaviour in Clark's nutcracker (Nucifraga columbiana) were examined experimentally. Most caches were made in soil within 5 cm of conspicuous large objects. Both seed-caching and non-seed-caching nutcrackers were able to locate caches. Seed-caching nutcrackers relocated caches using large objects as remembered visual cues. Soil microtopography and small (<2 cm diameter) objects may be used as cues to facilitate cache recovery but are not essential. Non-seed-caching nutcrackers located caches by using soil disturbances at cache sites as visual cues and by searching preferentially near objects where caches were concentrated. Success rates of seed-caching nutcrackers ranged from 52 to 78% and those of non-seed-caching nutcrackers ranged from 8 to 12%. Nutcrackers do not use random search or olfactory cues to locate caches.     

The influence of environmental conditions on cache recovery and cache pilferage by yellow pine chipmunks (Tamias amoenus) and deer mice (Peromyscus maniculatus)

I conducted a field experiment in 10 x 10 m enclosures to explorehow seed and soil moisture levels influence the ability ofknowledgeable and naive rodents to find natural caches of Jeffreypine (Pinus jeffreyi) seeds. Subjects were yellow pine chipmunks(Tamias amoenus) and deer mice (Peromyscus maniculatus) searchingfor caches that they had made, caches made by other individualsof the same species, or caches made by individuals of the otherspecies. Subjects that made caches (knowledgeable subjects)relied on spatial memory to find many of their own caches during recovery sessions, and their ability to locate caches was notaffected by water content of seeds or soil. Naive subjectsfound few caches under dry conditions, but under wet conditions,they located as many caches as did the rodents that made them.Naive subjects apparently relied on olfaction to find caches,a sensory modality that works more effectively under moist conditions. Subjects had as much success foraging for caches made by membersof their own species as for caches made by the other species.I present a hypothesis that predicts how foragers could modifypredominately memory-based search to predominately olfactory-basedsearch as the weather changes from dry to wet. When foragersrely on spatial memory, those foragers find only their own caches, but when they can also use olfaction, they pilfer cachesmade by other individuals. Consequently, the nature of competitiveinteractions among members of the seed-caching guild may changeas the weather changes.     

Seasonal diets of insectivorous birds using canopy gaps in a bottomland forest

ABSTRACT.   Little is known about how insectivorous bird diets are influenced by arthropod availability and about how these relationships vary seasonally. We captured birds in forest-canopy gaps and adjacent mature forest during 2001 and 2002 at the Savannah River Site in Barnwell County, South Carolina, and flushed their crops to gather information about arthropods eaten during four periods: spring migration, breeding, postbreeding, and fall migration. Arthropod availability for foliage- and ground-gleaning birds was examined by leaf clipping and pitfall trapping. Coleopterans and Hemipterans were used by foliage- and ground-gleaners more than expected during all periods, whereas arthropods in the orders Araneae and Hymenoptera were used as, or less than, expected based on availability during all periods. Ground-gleaning birds used Homopterans and Lepidopterans in proportions higher than availability during all periods. Arthropod use by birds was consistent from spring through fall migration, with no apparent seasonal shift in diet. Based on concurrent studies, heavily used orders of arthropods were equally abundant or slightly less abundant in canopy gaps than in the surrounding mature forest, but bird species were most frequently detected in gaps. Such results suggest that preferential feeding on arthropods by foliage-gleaning birds in gap habitats reduced arthropod densities or, alternatively, that bird use of gap and forest habitat was not determined by food resources. The abundance of arthropods across the stand may have allowed birds to remain in the densely vegetated gaps where thick cover provides protection from predators.     

Seed hoarding by rodents of the Monte Desert,Argentina

Food storage is an important adaptation of several animal species to the temporally variable or unpredictable food supplies that are typical of desert environments. In the present study, whether Eligmodontia typus and Graomys griseoflavus inhabiting sand dunes in the Monte desert displayed scatter‐hoarding was investigated. Both rodent species prepared surface caches by digging small holes (1–3 cm deep) in the soil. Caches were partially covered with sand and dry leaves, which meant that all of the cache sites at each station could be found. Most caches were found below shrubs, that is, in a sheltered microhabitat rather than near the food source. The mean distance between caches and seed sources was significantly higher than the mean distance from food sources to shrubs. The proportion of caches was significantly higher in shrubs with high safety cover. The adaptive significance of this foraging behaviour could be a competitive tactic for the rapid sequestering of food from a rich and ephemeral source. Such behaviour would reflect predation risk, which constitutes an important foraging cost in deserts, and could also influence plant community dynamics.     

Food hoarding behaviour of black-capped chickadees (Poecile atricapillus ) in relation to forest edges

In temperate forests, small birds avoid the use of forest edges in adverse winter weather suggesting high foraging costs in terms of energetic requirements. Since hoarding species will often retrieve caches during adverse winter weather, they may perceive forest edges, especially exposed ones, as low quality hoarding sites. We tested whether black-capped chickadees ( Poecile atricapillus) inhabiting fragmented forests modify and reduce hoarding activity near forest edges. We also tested whether hoarding behaviour will be most affected in sites with forest edges more exposed to extreme weather. Black-capped chickadees taking food from a feeder 30 m from the nearest forest edges hoarded items mostly towards the forest interior, whereas no preference in hoarding location was observed with birds taking food from a feeder placed >100 m from the edge. Furthermore, birds avoided direct flights towards forest edges and, at sites exposed to prevailing winds, hoarding trips were shorter than at other locations. These results suggest that individuals avoid hoarding near forest edges and there, they lower their investment in terms of hoarding effort. The observed difference in hoarding behaviour was more evident near forest edges delimiting wide unforested areas than in edges delimiting narrower unforested areas. Edge exposure to prevailing winds influenced hoarding behaviour much less. We suggest that hoarding birds may partially overcome the ecological costs of habitat loss and fragmentation due to abiotic edge effects. By hoarding food away from forest edges in good weather, they may use forest interiors as low-cost retrieval sites under adverse weather.     

Spatial orientation in Japanese quails (Coturnix coturnix japonica)

Finding a given location can be based on a variety of strategies, for example on the estimation of spatial relations between landmarks, called spatial orientation. In galliform birds, spatial orientation has been demonstrated convincingly in very young domestic chicks. We wanted to know whether adult Japanese quails (Coturnix coturnix japonica) without food deprivation are also able to use spatial orientation. The quails had to learn the relation of a food location with four conspicuous landmarks which were placed in the corners of a square shaped arena. They were trained to find mealworms in three adjacent food cups in a circle of 20 such cups. The rewarded feeders were located during training between the same two landmarks each of which showed a distinct pattern. When the birds had learned the task, all landmarks were displaced clockwise by 90 degrees. When tested in the new situation, all birds redirected their choices with respect to the landmark shift. In subsequent tests, however, the previously correct position was also chosen. According to our results, quails are using conspicuous landmarks as a first choice for orientation. The orientation towards the previously rewarded location, however, indicates that the neuronal representation of space which is used by the birds also includes more fine grain, less conspicuous cues, which are probably also taken into account in uncertain situations. We also presume that the rare orientation towards never rewarded feeders may be due to a foraging strategy instead of being mistakes.     

The effects of cache loss on choice of cache sites in black-capped chickadees

Food-storing birds lose a great deal of their stored food toother animals. We examined whether blackcapped chickadees (Parusairicapillus) modify their choice of cache sites using informationthat predicts cache loss. In experiment 1, birds learned toavoid caching at spatial locations where cache loss had previouslyoccurred, but they did not avoid caching near local color cuesthat predicted cache loss. Birds did not modify their generaluse of space in the aviary. Birds also learned to reduce searchingfor caches where spatial location predicted cache loss. Experiment2 confirmed the birds’ ability to discriminate among thespatial locations and the local color cues used in experiment1. In experiment 3, learning a food-rewarded approach to potentialcache sites occurred without any change in the choice of sitesfor caching. We discuss how chickadees selectively associatethe choice of cache site with its consequences, even over delaysof several hours between caching and cache recovery.     

Linking alternative food sources to winter habitat selection of herbivores in overbrowsed landscapes

During winter, ungulates in boreal forests must cope with high energetic costs related to locomotion in deep snow and reduced forage abundance and quality. At high density, ungulates face additional constraints, because heavy browsing reduces availability of woody browse, the main source of forage during winter. Under these severe conditions, large herbivores might forage on alternative food sources likely independent of browsing pressure, such as litterfall or windblown trees. We investigated the influence of alternative food sources on winter habitat selection, by studying female white-tailed deer (Odocoileus virginianus) living in 2 landscapes with contrasted browse abundance, recently logged and regenerated landscapes, in a population at high density and on a large island free of predators. We fitted 21 female white-tailed deer with Global Positioning System (GPS) collars and delineated winter home ranges and core areas. We measured snow conditions in different habitat categories and sampled vegetation in the core areas and in the rest of the home ranges to determine how forage abundance, protective cover, and snow conditions influenced habitat selection within the home range. In both landscapes, deer were less likely to use open habitat categories as snow accumulated on the ground. At a finer scale, deer inhabiting the regenerated landscape intensively used areas where balsam fir cover was intermediate with greater balsam fir browse density than in the rest of the home range. In the recently logged landscape, deer were more likely to be found near edges between clear-cuts and balsam fir stands and in areas where windblown balsam fir trees were present; the latter being the most influential variable. Although balsam fir browse was sparse and mainly out of reach in this landscape, deer increased the use of areas where it was present. Our results offer novel insights into the resource selection processes of northern ungulates, as we showed that access to winter forage, such as woody browse and alternative food sources, depends on climatic conditions and stochastic events, such as abundant compacted snow or windthrows. To compensate for these scarce and unpredictable food supplies, deer selected habitat categories, but mostly areas within those habitat categories, where the likelihood of finding browse, litterfall, and windblown trees was greatest. © 2011 The Wildlife Society.