Abstract: Behavioural innovations have become central to our thinking about how animals adjust to changing environments. It is now well established that animals vary in their ability to innovate, but understanding why remains a challenge. This is because innovations are rare, so studying innovation requires alternative experimental assays that create opportunities for animals to express their ability to invent new behaviours, or use pre-existing ones in new contexts. Problem solving of extractive foraging tasks has been put forward as a suitable experimental assay. We review the rapidly expanding literature on problem solving of extractive foraging tasks in order to better understand to what extent the processes underpinning problem solving, and the factors influencing problem solving, are in line with those predicted, and found, to underpin and influence innovation in the wild. Our aim is to determine whether problem solving can be used as an experimental proxy of innovation. We find that in most respects, problem solving is determined by the same underpinning mechanisms, and is influenced by the same factors, as those predicted to underpin, and to influence, innovation. We conclude that problem solving is a valid experimental assay for studying innovation, propose a conceptual model of problem solving in which motor diversity plays a more central role than has been considered to date, and provide recommendations for future research using problem solving to investigate innovation. This article is part of a Special Issue entitled: Cognition in the wild.

Abstract: The horse is a common domestic animal whose anatomy has been studied since the XVI century. However, a modern neuroanatomy of this species does not exist and most of the data utilized in textbooks and reviews derive from single specimens or relatively old literature. Here, we report information on the brain of Equus caballus obtained by sampling 131 horses, including brain weight (as a whole and subdivided into its constituents), encephalization quotient (EQ), and cerebellar quotient (CQ), and comparisons with what is known about other relevant species. The mean weight of the fresh brains in our experimental series was 598.63 g (SEM ± 7.65), with a mean body weight of 514.12 kg (SEM ± 15.42). The EQ was 0.78 and the CQ was 0.841. The data we obtained indicate that the horse possesses a large, convoluted brain, with a weight similar to that of other hoofed species of like mass. However, the shape of the brain, the noteworthy folding of the neocortex, and the peculiar longitudinal distribution of the gyri suggest an evolutionary specificity at least partially separate from that of the Cetartiodactyla (even-toed mammals and cetaceans) with whom Perissodactyla (odd-toed mammals) are often grouped.

Abstract: Viele Pferdebesitzer bevorzugen aus Angst vor aggressiven Interaktionen und Verletzungsgefahr der Tiere untereinander die Einzelhaltung, obwohl von Tierschutzorganisationen die Gruppenhaltung für Pferde empfohlen wird. In dieser Studie beobachteten wir während des alltäglichen Soziallebens als auch bei der Eingliederung von neuen Gruppenmitgliedern das Sozialverhalten, insbesondere das Aggressionsverhalten, von elf Gruppen domestizierter Pferde (Equus caballus) verschiedener Größe und Zusammensetzung. Während des alltäglichen Soziallebens hatten die Gruppe und der Paddock-Typ (Gras / kein Gras) keinen Einfluss auf die Verhaltensweisen, wohingegen die Paddockgröße unter 10000 m2 einen signifikanten Einfluss auf die submissiven Verhaltensweisen (GzLM; n=56; t=-2.061, P=0.044) und einen nicht signifikanten Einfluss auf die aggressiven Verhaltensweisen (GzLM; n=56; t=-1.782, P=0.081) hatte. Allerdings verringerten sich sowohl die aggressiven als auch die submissiven Verhaltensweisen mit steigendem Platzangebot bis zu 10000 m2 (Spearman rank Korrelation; n=56; aggressive Verhaltensweisen: r = -0.313, P = 0.019; submissive Verhaltensweisen: r = -0.328, P = 0.014). Während den Eingliederungen reduzierten sich die Aggressionen pro Stunde mit der Vergrößerung des Platzangebotes (Spearman rank Korrelation; n=28; r=-0.402, P=0.034). Dies zeigte sich noch deutlicher, wenn Beobachtungen mit einem Platzangebot von über 10000 m2 ausgeschlos- sen wurden (Spearman rank Korrelation; n=23; r=-0.549, P=0.007). Während des alltäglichen Soziallebens näherte sich der Aggressionslevel der Nulllinie an, wenn das Platzangebot pro Pferd mehr als 331 m2 betrug. Deshalb empfehlen wir zur Reduzierung des Aggressionslevels und des Verletzungsrisikos von sozial gehaltenen Pferdegruppen ein Platzangebot von mindestens 331 m2 pro Pferd.
[Even though animal welfare organisations propose group housing for horse welfare, many owners stable horses individually for fear of aggressive interactions and injury risks. In the present study we observed social behaviour, and especially aggressiveness, in eleven domestic horse groups (Equus caballus) of different size and composition, in basic social situations and when new group members were introduced. During basic social situations, the group and the type of paddock (grass / no grass) had no effect on any of the behaviours, where- as the enclosure size below 10,000 m2 had a significant effect on submissive behaviour (GzLM; n=56; t=-2.061, P=0.044) and an insignificant effect on aggressive behaviour (GzLM; n=56; t=-1.782, P=0.081). However, aggressive and submissive behaviour dimi- nished with the increase of enclosure sizes up to 10,000 m2 (Spearman rank correlation; n = 56; aggressive behaviour: r = -0.313, P=0.019; submissive behaviour: r=-0.328, P=0.014). During introductions, aggression levels per hour decreased with any increase of enclosure size (Spearman rank correlation; n=28; r=-0.402, P=0.034) and even more when enclosure sizes above 10,000 m2 were excluded (Spearman rank correlation; n=23; r=-0.549, P=0.007). During basic social situations the aggression level approached zero when the space allowance was more than 331 m2 per horse. We therefore recommend keeping horse groups in an enclosure with at least 331 m2 per horse to reduce aggression and injuries.]

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.