Eötvös Loránd University, Budapest

Biology Doctorate School, Head: Anna Erdei, DSc

Ethology Doctorate Program, Head: Ádám Miklósi, DSc

Dog (Canis familiaris) – robot interaction

Experimental research on ’ socio-communicative behaviour

Doctoral thesis

Anna Gergely

Supervisors: Ádám Miklósi, DSc József Topál, DSc

ELTE Department of Ethology 1117 Budapest, Pázmány Péter sétány 1/c

2014

Contents

1. . Introduction ...... 5 5 1.1 Social interaction ...... 5 1.2 Cooperative interactions between members of different species ...... 5 1.3 Animal-robot interaction as a special case of social interaction ...... 7 1.3.1 Analysis of the honeybee dance communication system ...... 8 1.3.2 Analysis of different channels of animal communication by means of robots ...... 9 1.3.3 Analysis of mating behaviour in birds ...... 10 1.3.4 Analysis of collective behaviour ...... 10 1.3.5 -robot interaction experiments ...... 11 1.4 The separation of behaviour from the body ...... 13 1.4.1 The general concept of the Unidentified Moving Object (UMO) ...... 15 1.5 Aims and questions ...... 16 1.5.1 Practical applications ...... 18 2. Experiments ...... 19 2.1 Experiment 1: The emergence of social interaction between dog and an Unidentified Moving Object (UMO) ...... 19 2.1.1 Materials and Methods ...... 20 2.1.2 Results ...... 25 2.1.3 Discussion ...... 32 2.2 Experiment 2: Dogs are able to adjust their social behaviour in accordance with their inanimate partners’ different capabilities...... 35 2.2.1 Materials and Methods ...... 36 2.2.2 Results ...... 42 2.2.3 Discussion ...... 44 2.3 Experiment 3: Dogs rapidly develop socially competent behaviour while interacting with a contingently responding Unidentified Moving Object (UMO) ...... 46 2.3.1 Materials and Methods ...... 48 2.3.2 Results ...... 53 2.3.3 Discussion ...... 57

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2.4 Experiment 4: Dogs are willing to follow the preference of their inanimate partner in a food choice task ...... 59 2.4.1 Materials and Methods ...... 60 2.4.2 Results ...... 65 2.4.3 Discussion ...... 68 2.5 Experiment 5: Dogs are able to generalise directional acoustic signals to different contexts and tasks ...... 70 2.5.1 Materials and Methods ...... 72 2.5.2 Results ...... 79 2.5.3 Discussion ...... 82 3. General discussion ...... 85 3.1 Outlook ...... 92 4. Acknowledgements ...... 93 5. References ...... 94 6. Summary ...... 103 7. Összefoglaló ...... 105 8. Appendix ...... 107

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„At bottom, robotics is about us. It is the discipline of emulating our lives,

of wondering how we work.”

Rod Grupen (2008)

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1. Introduction

1.1. Social interaction

Behaviour ecology defines social behaviour as interactions between individuals of the same species that has fitness consequences (Székely et al 2010), and which, at the functional level, is organised for achieving different goals such as finding a suitable mate, evading predators, cooperating in the acquisition of food etc. In contrast to many traits that are passively selected by the environment, social behaviour relies complex mechanisms where animals create a selective environment for themselves by interacting with each other. Accordingly, features of social behaviour and social traits have evolved specifically to contribute to the survival of the individual if group living provides some selective advantage (Székely et al 2010). In general, social interactions between individuals can be categorised as competitive or cooperative. Competition refers to interactions among two or more individuals in which the fitness of one is lowered by the presence of another (Begon et al 2006). Individuals compete for resources, territory, mate etc. required for growth, survival and reproduction in order to increase their fitness. In contrast, cooperation is defined as interactions with benefit for all participants involved (i.e. which increase the reproductive success of the participants) (Noë 2006). The so-called kin-selection theory provides a solution to the problem of cooperative behaviour between relatives and helps to understand the evolution of social behaviour (e.g. West et al 2002). Individuals predicted to behave less competitively and more cooperatively toward their relatives, because they share a relatively high proportion of their genes. Consequently, by helping kin (i.e. relative), individuals are helping copies of their own genes (Hamilton 1964). At the same time, this theory did not solve the riddle of cooperation among unrelated individuals from the same or from different species. The latter issue referred to as the central theoretical problem of sociobiology (Wilson 1975) and has been studied by a large number of researchers.

1.2. Cooperative interactions between members of different species

Because of the functional similarities in the life of different species one may expect that a range of social behaviours reflect some commonalities (matching competencies) based on

5 ancient homologies or convergent evolutionary processes. Given that group living or limited co-existence may also confer some advantages in the case of different species social behaviour could also emerge in heterospecific contexts, both developmentally and on an evolutionary time scale (e.g. interspecific communication, see also Kostan 2002, Miklósi & Gácsi 2012). One well known example for this is the collaboration between honey guide birds (Indicator indicator) and African tribal people in order to find honey by locating beehives in the forest (Isack & Reyer 1989). In another case Bshary et al (2006) show that the grouper (Plectropomus pessuliferus) and the giant moray eel (Gymnothorax javanicus) hunt cooperatively, probably, because they have complementary behavioural skills, and the two partners, belonging to different species, are able to coordinate their actions at the behavioural level, that is, the grouper uses a specific visual signal to lure the moray eel on a hunting trip (Bshary et al 2006). The evolution of such interspecific social interaction has long been a topic in the field of behavioural ecology, evolutionary biology and ethology, as well as different cognitive prerequisites required for various forms of social behaviour to take place in humans and non- humans alike. Three paths have been reviewed in the study of cooperation between non- related individuals from the same or from different species (for a review see Dugatkin 2002). Trivers (1971) argued that one path to cooperative behaviour, among humans and non-human animals, is reciprocity which most likely evolves when the minor cost paid by the helper individual is made up for when the other individual restores the favour in the future. Another possibility is mutualism which occurs in “harsh” environment where the cost of not being cooperative is immediate and the benefit of cooperation outweighs cheating (i.e. by-product mutualism, e.g. Dugatkin 2002). The third and probably the most controversial one is trait- group selection where natural selection operates at two levels: within groups and between groups. Group selection models showed that cooperation is favoured when the response to between-group selection outweighs the response to within-group selection (see Sober & Wilson 1998 for a review). In addition to examine fitness consequences, the study of cognitive abilities involved in social interactions is also essential. For example effective cooperative hunting requires skills of communication (e.g. initializing the hunt) and behavioural synchronisation (e.g. Clutton- Brock 1977, 1996) that relies on cognitive abilities like role differentiation and coordinated movements (Boesch & Boesch 1989). Effective communication for initializing the joint hunt via signals appears more difficult to achieve between heterospecific interactants of sharply different behavioural patterns. Based on an observational study Pryor and co-workers (1990)

6 described cooperative fish