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Hamilton, W. D. (1971). Geometry for the selfish herd. J. Theor. Biol., 31(2), 295–311.
Abstract: This paper presents an antithesis to the view that gregarious behaviour is evolved through benefits to the population or species. Following Galton (1871) and Williams (1964) gregarious behaviour is considered as a form of cover-seeking in which each animal tries to reduce its chance of being caught by a predator.
It is easy to see how pruning of marginal individuals can maintain centripetal instincts in already gregarious species; some evidence that marginal pruning actually occurs is summarized. Besides this, simply defined models are used to show that even in non-gregarious species selection is likely to favour individuals who stay close to others.
Although not universal or unipotent, cover-seeking is a widespread and important element in animal aggregation, as the literature shows. Neglect of the idea has probably followed from a general disbelief that evolution can be dysgenic for a species. Nevertheless, selection theory provides no support for such disbelief in the case of species with outbreeding or unsubdivided populations.
The model for two dimensions involves a complex problem in geometrical probability which has relevance also in metallurgy and communication science. Some empirical data on this, gathered from random number plots, is presented as of possible heuristic value.
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Clutton-Brock, T. H., Greenwood, P. J., & Powell, R. P. (1976). Ranks and relationships in Highland ponies and Highland Cows. Z. Tierpsychol., 41(2), 202–216.
Abstract: Recent studies of primates have questioned the importance of dominance hierarchies in groups living under natural conditions. In a herd of Highland ponies and one of Highland cattle grazing under free-range conditions on the Isle of Rhum (Inner Hebrides) well defined hierarchies were present. The provision of food produced a marked increase in the frequency of agonistic interactions but had no effect on the rank systems of the two herds. While rank was clearly important in affecting the distribution of agonistic interactions, it was poorly related to behaviour in non-agonistic situations.
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Skov-Rackette, S. I., Miller, N. Y., & Shettleworth, S. J. (2006). What-where-when memory in pigeons. J Exp Psychol Anim Behav Process, 32(4), 345–358.
Abstract: The authors report a novel approach to testing episodic-like memory for single events. Pigeons were trained in separate sessions to match the identity of a sample on a touch screen, to match its location, and to report on the length of the retention interval. When these 3 tasks were mixed randomly within sessions, birds were more than 80% correct on each task. However, performance on 2 different tests in succession after each sample was not consistent with an integrated memory for sample location, time, and identity. Experiment 2 tested binding of location and identity memories in 2 different ways. The results were again consistent with independent feature memories. Implications for tests of episodic-like memory are discussed.
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Range, F., Bugnyar, T., Schlogl, C., & Kotrschal, K. (2006). Individual and sex differences in learning abilities of ravens. Behav. Process., 73(1), 100–106.
Abstract: Behavioral and physiological characteristics of individuals within the same species have been found to be stable across time and contexts. In this study, we investigated individual differences in learning abilities and object and social manipulation to test for consistency within individuals across different tasks. Individual ravens (Corvus corax) were tested in simple color and position discrimination tasks to establish their learning abilities. We found that males were significantly better in the acquisition of the first discrimination task and the object manipulation task, but not in any of the other tasks. Furthermore, faster learners engaged less often in manipulations of conspecifics and exploration of objects to get access to food. No relationship between object and social manipulation and reversal training were found. Our results suggest that individual differences in regard to the acquisition of new tasks may be related to personalities or at least object manipulation in ravens.
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Poti, P. (2005). Chimpanzees' constructional praxis (Pan paniscus, P. troglodytes). Primates, 46(2), 103–113.
Abstract: This study investigated chimpanzees' spontaneous spatial constructions with objects and especially their ability to repeat inter-object spatial relations, which is basic to understanding spatial relations at a higher level than perception or recognition. Subjects were six chimpanzees-four chimpanzees and two bonobos-aged 6-21 years, all raised in a human environment from an early age. Only minor species differences, but considerable individual differences were found. The effect of different object samples was assessed through a comparison with a previous study. A common overall chimpanzee pattern was also found. Chimpanzees repeated different types of inter-object spatial relations such as insertion (I), or vertical (V), or next-to (H) relations. However chimpanzees repeated I or V relations with more advanced procedures than when repeating H relations. Moreover, chimpanzees never repeated combined HV relations. Compared with children, chimpanzees showed a specific difficulty in repeating H relations. Repeating H relations is crucial for representing and understanding multiple reciprocal spatial relations between detached elements and for coordinating independent positions in space. Therefore, the chimpanzees' difficulty indicates a fundamental difference in constructive space in comparison to humans. The findings are discussed in relation to issues of spatial cognition and tool use.
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Fiset, S., & Leblanc, V. (2007). Invisible displacement understanding in domestic dogs (Canis familiaris): the role of visual cues in search behavior. Anim. Cogn., 10(2), 211–224.
Abstract: Recently, (Collier-Baker E, Davis JM, Suddendorf T (2004) J Comp Psychol 118:421-433) suggested that domestic dogs do not understand invisible displacements. In the present study, we further investigated the hypothesis that the search behavior of domestic dogs in invisible displacements is guided by various visual cues inherent to the task rather than by mental representation of an object's past trajectory. Specifically, we examined the role of the experimenter as a function of the final position of the displacement device in the search behavior of domestic dogs. Visible and invisible displacement problems were administered to dogs (N = 11) under two conditions. In the Visible-experimenter condition, the experimenter was visible whereas in the Concealed-experimenter condition, the experimenter was visibly occluded behind a large rigid barrier. Our data supported the conclusion that dogs do not understand invisible displacements but primarily search as a function of the final position of the displacement device and, to a lesser extent, the position of the experimenter.
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Gibbs, S. E. B., Lea, S. E. G., & Jacobs, L. F. (2007). Flexible use of spatial cues in the southern flying squirrel (Glaucomys volans). Anim. Cogn., 10(2), 203–209.
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.
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Chiesa, A. D., Pecchia, T., Tommasi, L., & Vallortigara, G. (2006). Multiple landmarks, the encoding of environmental geometry and the spatial logics of a dual brain. Anim. Cogn., 9(4), 281–293.
Abstract: A series of place learning experiments was carried out in young chicks (Gallus gallus) in order to investigate how the geometry of a landmark array and that of a walled enclosure compete when disoriented animals could rely on both of them to re-orient towards the centre of the enclosure. A square-shaped array (four wooden sticks) was placed in the middle of a square-shaped enclosure, the two structures being concentric. Chicks were trained to ground-scratch to search for food hidden in the centre of the enclosure (and the array). To check for effects of array degradation, one, two, three or all landmarks were removed during test trials. Chicks concentrated their searching activity in the central area of the enclosure, but their accuracy was inversely contingent on the number of landmarks removed; moreover, the landmarks still present within the enclosure appeared to influence the shape of the searching patterns. The reduction in the number of landmarks affected the searching strategy of chicks, suggesting that they had focussed mainly on local cues when landmarks were present within the enclosure. When all the landmarks were removed, chicks searched over a larger area, suggesting an absolute encoding of distances from the local cues and less reliance on the relationships provided by the geometry of the enclosure. Under conditions of monocular vision, chicks tended to rely on different strategies to localize the centre on the basis of the eye (and thus the hemisphere) in use, the left hemisphere attending to details of the environment and the right hemisphere attending to the global shape.
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Wallace, D. G., Hamilton, D. A., & Whishaw, I. Q. (2006). Movement characteristics support a role for dead reckoning in organizing exploratory behavior. Anim. Cogn., 9(3), 219–228.
Abstract: Rat exploration is an organized series of trips. Each exploratory trip involves an outward tour from the refuge followed by a return to the refuge. A tour consists of a sequence of progressions with variable direction and speed concatenated by stops, whereas the return consists of a single direct progression. We have argued that processing self-movement information generated on the tour allows a rat to plot the return to the refuge. This claim has been supported by observing consistent differences between tour and return segments independent of ambient cue availability; however, this distinction was based on differences in movement characteristics derived from multiple progressions and stops on the tour and the single progression on the return. The present study examines movement characteristics of the tour and return progressions under novel-dark and light conditions. Three novel characteristics of progressions were identified: (1) linear speeds and path curvature of exploratory trips are negatively correlated, (2) tour progression maximum linear speed and temporal pacing varies as a function of travel distance, and (3) return progression movement characteristics are qualitatively different from tour progressions of comparable length. These observations support a role for dead reckoning in organizing exploratory behavior.
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Sturz, B. R., Bodily, K. D., & Katz, J. S. (2006). Evidence against integration of spatial maps in humans. Anim. Cogn., 9(3), 207–217.
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.
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