Abstract: The four questions that Niko Tinbergen identified for behavioural biology ? evolution, function, development and causation ? are all important and should be studied in their own right. Recently, there has been a debate as to whether these four questions should be investigated separately or whether they should be integrated. Integration of the four questions has been attempted in novel research disciplines such as cognitive ecology, evolutionary psychology and neuroecology. Euan Macphail and I have criticised these integrative approaches, suggesting that they are fundamentally flawed as they confound function and mechanism. Investigating the function or evolutionary history of a behaviour or cognitive system is important and entirely legitimate. However, such investigations cannot provide us with answers to questions about the mechanisms underlying behaviour or cognition. At most, functional or evolutionary considerations can provide clues that may be useful for a causal analysis of the underlying mechanisms. However, these clues can be misleading and are often wrong, as is illustrated with examples from song learning and food storing in birds. After summarising the main issues in the neuroecology debate, I discuss some misunderstandings that were apparent in the responses to our critique, as well as some recent relevant data. Recent results do not support the neuroecological approach. Finally, I suggest that the way forward is a cautious and critical use of functional and evolutionary clues in the study of the mechanisms of behaviour.

Abstract: Th e applicability of the concept of dominance was investigated in a captive herd of  Icelandic
horses and  ponies of diff erent breeds. Eight out of  behaviours possibly related
to dominance occurred frequently enough to be investigated in detail. For these eight agonistic
behaviours the coverage, the unidirectionality in the exchange, and the degree of
transitivity (Landau`s linearity index) were calculated. Four off ensive behaviours, together
with avoidance, were suitable for further analysis with regard to dominance. Th e patterns
of asymmetries with which these behaviours were exchanged were suffi ciently similar as to
justify the application of the dominance concept and to construct a (nearly) linear dominance
hierarchy. Th e rank order of the castrated stallions was completely linear, the hierarchy
of the mares was almost completely linear. Th e results suggest that off ensive and defensive
aggressive behaviours should be treated separately and that the concept of dominance
is applicable. However, ritualized formal dominance signals between adult horses appear to
be (almost) absent. Th e rank positions of the individuals were correlated with age and residency
in the herd but not with height. Middle ranking horses tended to be more frequently
in the close vicinity of another horse than high ranking or low ranking horses. Over and
above this correlation at the individual level, it was found that pairs of horses close in rank
to each other were more often also spatially close to each other. Being in oestrus did not infl
uence the dominance relationships between mares. For castrated stallions the rank positions
were correlated with the age at which they were castrated. Th is suggests that in male
horses experience prior to neutering infl uences the behaviour afterwards.

Abstract: ABSTRACT: BACKGROUND: Researchers have developed a variety of techniques for the visual presentation of quantitative data. These techniques can help to reveal trends and regularities that would be difficult to see if the data were left in raw form. Such techniques can be of great help in exploratory data analysis, making apparent the organization of data sets, developing new hypotheses, and in selecting effects to be tested by statistical analysis. Researchers studying social interaction in groups of animals and humans, however, have few tools to present their raw data visually, and it can be especially difficult to perceive patterns in these data. In this paper I introduce a new graphical method for the visual display of interaction records in human and animal groups, and I illustrate this method using data taken on chickens forming dominance hierarchies. RESULTS: This new method presents data in a way that can help researchers immediately to see patterns and connections in long, detailed records of interaction. I show a variety of ways in which this new technique can be used: (1) to explore trends in the formation of both group social structures and individual relationships; (2) to compare interaction records across groups of real animals and between real animals and computer-simulated animal interactions; (3) to search for and discover new types of small-scale interaction sequences; and (4) to examine how interaction patterns in larger groups might emerge from those in component subgroups. In addition, I discuss how this method can be modified and extended for visualizing a variety of different kinds of social interaction in both humans and animals. CONCLUSION: This method can help researchers develop new insights into the structure and organization of social interaction. Such insights can make it easier for researchers to explain behavioural processes, to select aspects of data for statistical analysis, to design further studies, and to formulate appropriate mathematical models and computer simulations.

Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.