Abstract: Insects, birds, and mammals have been shown capable of encoding spatial information in memory using multiple strategies or frames of reference simultaneously. These strategies include orientation to a goal-specific cue or beacon, to the position of the goal in an array of local landmarks, or to its position in the array of distant landmarks, also known as the global frame of reference. From previous experiments, it appears that birds and mammals that scatter hoard rely primarily on a global frame of reference, but this generalization depends on evidence from only a few species. Here we examined spatial memory in a previously unstudied scatter hoarder, the southern flying squirrel. We dissociated the relative weighting of three potential spatial strategies (beacon, global, or relative array strategy) with three probe tests: transposition of beacon and the rotation or the expansion of the array. The squirrels' choices were consistent with a spatial averaging strategy, where they chose the location dictated by at least two of the three strategies, rather than using a single preferred frame of reference. This adaptive and flexible heuristic has not been previously described in animal orientation studies, yet it may be a common solution to the universal problem of encoding and recalling spatial locations in an ephemeral physical landscape.

Abstract: The effects of the size of the environment on animals' spatial reorientation was investigated. Domestic chicks were trained to find food in a corner of either a small or a large rectangular enclosure. A distinctive panel was located at each of the four corners of the enclosures. After removal of the panels, chicks tested in the small enclosure showed better retention of geometrical information than chicks tested in the large enclosure. In contrast, after changing the enclosure from a rectangular-shaped to a square-shaped one, chicks tested in the large enclosure showed better retention of landmark (panels) information than chicks tested in the small enclosure. No differences in the encoding of the overall arrangement of landmarks were apparent when chicks were tested for generalisation in an enclosure differing from that of training in size together with a transformation (affine transformation) that altered the geometric relations between the target and the shape of the environment. These findings suggest that primacy of geometric or landmark information in reorientation tasks depends on the size of the experimental space, likely reflecting a preferential use of the most reliable source of information available during visual exploration of the environment.

Abstract: A dynamic 3-D virtual environment was constructed for humans as an open-field analogue of Blaisdell and Cook's (2005) pigeon foraging task to determine if humans, like pigeons, were capable of integrating separate spatial maps. Participants used keyboard keys and a mouse to search for a hidden goal in a 4x4 grid of raised cups. During Phase 1 training, a goal was consistently located between two landmarks (Map 1: blue T and red L). During Phase 2 training, a goal was consistently located down and left of a single landmark (Map 2: blue T). Transfer trials were then conducted in which participants were required to make choices in the presence of the red L alone. Cup choices during transfer assessed participants' strategies: association (from Map 1), generalization (from Map 2), or integration (combining Map 1 and 2). During transfer, cup choices increased to a location which suggested an integration strategy and was consistent with results obtained with pigeons. However, additional analyses of the human data suggested participants initially used a generalization strategy followed by a progressive shift in search behavior away from the red L. This shift in search behavior during transfer was responsible for the changes in cup choices across transfer trials and was confirmed by a control condition. These new analyses offer an alternative explanation to the spatial integration account proposed for pigeons.

Abstract: Locating food and refuge is essential for an animal's survival. However, little is known how mammals navigate under natural conditions and cope with given environmental constraints. In a series of six experiments, I investigated landmark-based navigation in free-ranging Columbian ground squirrels (Spermophilus columbianus). Squirrels were trained individually to find a baited platform within an array of nine identical platforms and artificial landmarks set up on their territories. After animals learned the location of the food platform in the array, the position of the latter with respect to local artificial, local natural, and global landmarks was manipulated, and the animal's ability to find the food platform was tested. When only positions of local artificial landmarks were changed, squirrels located food with high accuracy. When the location of the array relative to global landmarks was altered, food-finding accuracy decreased but remained significant. In the absence of known global landmarks, the presence of a familiar route and natural local landmarks resulted in significant but not highly accurate performance. Squirrels likely relied on multiple types of cues when orienting towards a food platform. Local landmarks were used only as a secondary mechanism of navigation, and were not attended to when a familiar route and known global landmarks were present. This study provided insights into landmark use by a wild mammal in a natural situation, and it demonstrated that an array of platforms can be employed to investigate landmark-based navigation under such conditions.

Abstract: Detour behaviour is the ability of an animal to reach a goal stimulus by moving round any interposed obstacle. It has been widely studied and has been proposed as a test of insight learning in several species of mammals, but few data are available in birds. A comparative study in three species of birds, belonging to different eco-ethological niches, allows a better understanding of the cognitive mechanism of such detour behaviour. Young quails (Coturnix sp.), herring gulls (Larus cachinnans) and canaries (Serinus canaria), 1 month old, 10-25 days old and 4-6 months old, respectively, were tested in a detour situation requiring them to abandon a clear view of a biologically interesting object (their own reflection in a mirror) in order to approach that object. Birds were placed in a closed corridor, at one end of which was a barrier through which the object was visible. Four different types of barrier were used: vertical bar, horizontal bar, grid and transparent. Two symmetrical apertures placed midline in the corridor allowed the birds to adopt routes passing around the barrier. After entering the apertures, birds could turn either right or left to re-establish social contact with the object in the absence of any local sensory cues emanating from it. Quails appeared able to solve the task, though their performance depended on the type of barrier used, which appeared to modulate their relative interest in approaching the object or in exploring the surroundings. Young herring gulls also showed excellent abilities to locate spatially the out-of-view object, except when the transparent barrier was used. Canaries, on the other hand, appeared completely unable to solve the detour task, whatever barrier was in use. It is suggested that these species differences can be accounted for in terms of adaptation to a terrestrial or aerial environment.