Why do African elephants (Loxodonta africana) simulate oestrus? An analysis of longitudinal data.

Female African elephants signal oestrus via chemicals in their urine, but they also exhibit characteristic changes to their posture, gait and behaviour when sexually receptive. Free-ranging females visually signal receptivity by holding their heads and tails high, walking with an exaggerated gait, and displaying increased tactile behaviour towards males. Parous females occasionally exhibit these visual signals at times when they are thought not to be cycling and without attracting interest from musth males. Using demographic and behavioural records spanning a continuous 28-year period, we investigated the occurrence of this "simulated" oestrus behaviour. We show that parous females in the Amboseli elephant population do simulate receptive oestrus behaviours, and this false oestrus occurs disproportionately in the presence of naïve female kin who are observed coming into oestrus for the first time. We compare several alternative hypotheses for the occurrence of this simulation: 1) false oestrus has no functional purpose (e.g., it merely results from abnormal hormonal changes); 2) false oestrus increases the reproductive success of the simulating female, by inducing sexual receptivity; and 3) false oestrus increases the inclusive fitness of the simulating female, either by increasing the access of related females to suitable males, or by encouraging appropriate oestrus behaviours from female relatives who are not responding correctly to males. Although the observed data do not fully conform to the predictions of any of these hypotheses, we rule out the first two, and tentatively suggest that parous females most likely exhibit false oestrus behaviours in order to demonstrate to naïve relatives at whom to direct their behaviour.

Primate social cognition: uniquely primate, uniquely social, or just unique?

Primates undoubtedly have impressive abilities in perceiving, recognizing, manipulating, and predicting other individuals, but only great apes seem to recognize the cognitive basis of manipulative and cooperative tactics or the concept of self. None of these abilities is unique to primates. We distinguish (1) a package of quantitative advantages in social sophistication, perhaps based on more efficient memory, in which neocortical enlargement is associated with the challenge of social living; from (2) a qualitative difference in understanding, whose taxonomic distribution--including several distantly related species, including birds--does not point to an evolutionary origin in social challenges and may instead relate to a need to acquire novel ways of dealing with the physical world. The ability of great apes to learn new manual routines by parsing action components may have driven their qualitatively greater social skill, suggesting that strict partition of physical and social cognition is likely to be misleading.

Imitation: what animal imitation tells us about animal cognition.

Imitation of actions is widespread in the animal kingdom, but the mental capacities thereby implied vary greatly according to the adaptive function of copying. Behavioral synchrony in social species has many possible benefits, including minimizing predation risk and using food resources optimally, but can be understood by the simple cognitive mechanism of response facilitation by priming. Imitation can send a social message, either one of short-term meshing or group identity. Where the imitative match is opaque, as in neonatal imitation, the correspondence problem may imply an innate system of behavior matching; but in other cases, no more than priming may be involved, although there are persistent suggestions that great ape imitation implies empathic abilities. Imitation in the service of learning new skills by following another's example can be divided into contextual imitation (when to employ a familiar action, and to what problem) and production imitation (learning of new skills by imitation). Cognitively, the former requires little more than response facilitation, whereas production imitation needs at least the ability to extract the statistical regularities of repeated action and to incorporate the result into hierarchical program construction. Among our close relatives, only the great apes show much evidence of production imitation of actions, along with the ability to selectively imitate the most rational components of what they observe. Copyright © 2010 John Wiley & Sons, Ltd. For further resources related to this article, please visit the WIREs website.

African elephants have expectations about the locations of out-of-sight family members.

Monitoring the location of conspecifics may be important to social mammals. Here, we use an expectancy-violation paradigm to test the ability of African elephants (Loxodonta africana) to keep track of their social companions from olfactory cues. We presented elephants with samples of earth mixed with urine from female conspecifics that were either kin or unrelated to them, and either unexpected or highly predictable at that location. From behavioural measurements of the elephants' reactions, we show that African elephants can recognize up to 17 females and possibly up to 30 family members from cues present in the urine-earth mix, and that they keep track of the location of these individuals in relation to themselves.

Brain evolution: when is a group not a group?

In testing the 'social brain hypothesis' with comparative data, most research has used group size as an index of cognitive challenge. Recent work suggests that this measure is too crude to apply to a wide range of species, and biologists may need to develop other ways of extending these analyses.

Elephants classify human ethnic groups by odor and garment color.

Animals can benefit from classifying predators or other dangers into categories, tailoring their escape strategies to the type and nature of the risk. Studies of alarm vocalizations have revealed various levels of sophistication in classification. In many taxa, reactions to danger are inflexible, but some species can learn the level of threat presented by the local population of a predator or by specific, recognizable individuals. Some species distinguish several species of predator, giving differentiated warning calls and escape reactions; here, we explore an animal's classification of subgroups within a species. We show that elephants distinguish at least two Kenyan ethnic groups and can identify them by olfactory and color cues independently. In the Amboseli ecosystem, Kenya, young Maasai men demonstrate virility by spearing elephants (Loxodonta africana), but Kamba agriculturalists pose little threat. Elephants showed greater fear when they detected the scent of garments previously worn by Maasai than by Kamba men, and they reacted aggressively to the color associated with Maasai. Elephants are therefore able to classify members of a single species into subgroups that pose different degrees of danger.

Sociality, evolution and cognition.

Variations in brain size and proportions can be linked to the cognitive capacities of different animal species, and correlations with ecology may give clues to the evolutionary origins of these specializations. Much recent evidence has implicated the social domain as a major challenge driving increases in problem-solving abilities of mammals. However, the methods of measurement available to researchers are often indirect and sometimes appear to give conflicting answers, and other intellectual challenges may also have been influential in cognitive evolution. While the cause of an evolutionary increase in intelligence may be domain-specific (sociality, for example), and the brain specialization that results may largely implicate a single perceptual system, such as vision, the intelligence shown in consequence can be very 'general-purpose' (as in primates and some avian taxa). Future research needs to get beyond vague ascription of 'greater intelligence' or 'faster learning' towards a precise account of the cognitive mechanisms that underlie particular mental skills in different species; that will allow theory-testing against data from complex, natural situations as well as from the laboratory, on a common metric.

Creative or created: using anecdotes to investigate animal cognition.

In non-human animals, creative behaviour occurs spontaneously only at low frequencies, so is typically missed by standardised observational methods. Experimental approaches have tended to rely overly on paradigms from child development or adult human cognition, which may be inappropriate for species that inhabit very different perceptual worlds and possess quite different motor capacities than humans. The analysis of anecdotes offers a solution to this impasse, provided certain conditions are met. To be reliable, anecdotes must be recorded immediately after observation, and only the records of scientists experienced with the species and the individuals concerned should be used. Even then, interpretation of a single record is always ambiguous, and analysis is feasible only when collation of multiple records shows that a behaviour pattern occurs repeatedly under similar circumstances. This approach has been used successfully to study a number of creative capacities of animals: the distribution, nature and neural correlates of deception across the primate order; the occurrence of teaching in animals; and the neural correlates of several aptitudes--in birds, foraging innovation, and in primates, innovation, social learning and tool-use. Drawing on these approaches, we describe the use of this method to investigate a new problem, the cognition of the African elephant, a species whose sheer size and evolutionary distance from humans renders the conventional methods of comparative psychology of little use. The aim is both to chart the creative cognitive capacities of this species, and to devise appropriate experimental methods to confirm and extend previous findings.