Abstract: Communication in rhinos is primarily mediated by the vocal and olfactory signals as they have relatively poor eyesight. White rhinos are the most social of all the rhinoceros species, they defecate at common dungheaps and the adult bulls use dung and urine to mark their territory. Chemical communication may therefore be particularly important in the social interactions of white rhinos, and its knowledge could be very helpful in their management and conservation. However, no studies have investigated up until now the olfactory discrimination in any rhinoceros species in the wild. We have experimentally studied the reactions of the wild southern white rhinos (Ceratotherium simum) to the dung of familiar and unfamiliar adult females and adult territorial males. We registered the number of sniffing events, the duration of sniffing and the latency of the vigilance posture from the onset of sniffing. The dung of unfamiliar rhinos was sniffed longer than that of familiar rhinos. The rhinos showed a shorter latency of vigilance posture to the familiar dung of males than that of females. For unfamiliar dung, they displayed a shorter latency of vigilance posture to female than male dung. Our results indicate that the rhinos are able to discriminate the familiarity and sex of conspecifics from the smell of their dung. Olfactory cues could therefore play an important role in the social relationships and spatial organization of the southern white rhinoceros.

Abstract: Recognition of information from acoustic signals is crucial in many animals, and individuals are under selection pressure to discriminate between the signals of conspecifics and heterospecifics or males and females. Here, we first report that rhinos use information encoded in their calls to assess conspecifics and individuals of closely related species. The southern (Ceratotherium simum) and critically endangered northern (C. cottoni) white rhinos are the most social out of all the rhinoceros species and use a contact call pant. We found that southern white rhino pant calls provide reliable information about the caller’s sex, age class and social situation. Playback experiments on wild territorial southern white rhinoceros males revealed that they responded more strongly to the pant calls of conspecific females compared to the calls of other territorial males. This suggests that pant calls are more important form of communication between males and females than between territorial males. Territorial southern males also discriminated between female and territorial male calls of northern species and reacted more intensively to the calls of northern than southern males. This might be caused by a novelty effect since both species naturally live in allopatry. We conclude that white rhinos can directly benefit from assessing individuals at long distances using vocal cues especially because their eyesight is poor. Pant calls thus likely play a significant role in their social relationships and spatial organization. In addition, better understanding of vocal communication in white rhinos might be helpful in conservation management particularly because of their low reproduction in captivity.

Abstract: Some domestic animals are thought to be skilled at social communication with humans due to the process of domestication. Horses, being in close relationship with humans, similar to dogs, might be skilled at communication with humans. Previous studies have indicated that they are sensitive to bodily signals and the attentional state of humans; however, there are few studies that investigate communication with humans and responses to the knowledge state of humans. Our first question was whether and how horses send signals to their potentially helpful but ignorant caretakers in a problem-solving situation where a food item was hidden in a bucket that was accessible only to the caretakers. We then examined whether horses alter their behaviours on the basis of the caretakers’ knowledge of where the food was hidden. We found that horses communicated to their caretakers using visual and tactile signals. The signalling behaviour of the horses significantly increased in conditions where the caretakers had not seen the hiding of the food. These results suggest that horses alter their communicative behaviour towards humans in accordance with humans’ knowledge state.

Abstract: Deception, the use of false signals to modify the behaviour of the receiver, occurs in low frequencies even in stable signalling systems. For example, it can be advantageous for subordinate individuals to deceive in competitive situations. We investigated in a three-way choice task whether dogs are able to mislead a human competitor, i.e. if they are capable of tactical deception. During training, dogs experienced the role of their owner, as always being cooperative, and two unfamiliar humans, one acting ‘cooperatively’ by giving food and the other being ‘competitive’ and keeping the food for themselves. During the test, the dog had the options to lead one of these partners to one of the three potential food locations: one contained a favoured food item, the other a non-preferred food item and the third remained empty. After having led one of the partners, the dog always had the possibility of leading its cooperative owner to one of the food locations. Therefore, a dog would have a direct benefit from misleading the competitive partner since it would then get another chance to receive the preferred food from the owner. On the first test day, the dogs led the cooperative partner to the preferred food box more often than expected by chance and more often than the competitive partner. On the second day, they even led the competitive partner less often to the preferred food than expected by chance and more often to the empty box than the cooperative partner. These results show that dogs distinguished between the cooperative and the competitive partner, and indicate the flexibility of dogs to adjust their behaviour and that they are able to use tactical deception.

Abstract: Pair and social bonding has been documented in various taxa, where pair formations are often described as being driven by kinship or sexual motivation. However, pair-bonding between unrelated individuals where sexual motivation is not a factor is not well documented. Many social relationships and pair-bonds between members of a dyad are facilitated by each individual's ability to recognise their partner using cues which are characteristic of that particular individual. The aims of this study were i) to investigate the existence of pair-bonding in domestic donkeys and ii) to determine whether members of a dyad could recognise their companion during a Y-maze recognition test. Subjects were 55 unrelated donkeys (38 gelded males, 15 females) in seven groups of mixed or same sex, comprising 4?14 individuals. Spatial proximity (nearest-neighbour) was observed three times a day over a 22-day period. Using a simulation approach based on observed data to generate randomised nearest-neighbour matrices, the statistical significance of social relationships was estimated. Of these, 42 (79.2%) were involved in significantly (p<0.05) non-random nearest-neighbour relationships, most of which were reciprocal pair relationships. Based on the spatial data, 24 of the donkeys which had shown significant reciprocal nearest-neighbour preferences for one individual (companion) were then used in a Y-maze recognition test in which they were presented with a choice of their companion and either a familiar donkey from the same group or an unfamiliar donkey from a different group. Donkeys? spatial location in the Y-maze demonstrated a preference for their companion versus familiar (one sample Wilcoxon signed rank test, W=239, p=0.002) or unfamiliar donkeys (W=222, p=0.041). These results verify anecdotal evidence from donkey handlers that donkeys often form pair-bonds, and show that reciprocal social preference and recognition are the basis of these. Pair-bond formation and companionship among donkeys have potential implications for their management, husbandry and welfare.