Daniela Hedwig

Genetic Architecture of Tameness in a Rat Model of Animal Domestication

A common feature of domestic animals is tameness—i.e., they tolerate and are unafraid of human presence and handling. To gain insight into the genetic basis of tameness and aggression, we studied an intercross between two lines of rats (Rattus norvegicus) selected over >60 generations for increased tameness and increased aggression against humans, respectively. We measured 45 traits, including tameness and aggression, anxiety-related traits, organ weights, and levels of serum components in >700 rats from an intercross population. Using 201 genetic markers, we identified two significant quantitative trait loci (QTL) for tameness. These loci overlap with QTL for adrenal gland weight and for anxiety-related traits and are part of a five-locus epistatic network influencing tameness. An additional QTL influences the occurrence of white coat spots, but shows no significant effect on tameness. The loci described here are important starting points for finding the genes that cause tameness in these rats and potentially in domestic animals in general.

An experimental study of nettle feeding in captive gorillas.

Mountain gorillas (Gorilla beringei beringei) in Karisoke, Rwanda, feed on the stinging nettle Laportea alatipes by means of elaborate processing skills. Byrne [e.g. Philosophical Transactions of the Royal Society of London, Series B, Biological Sciences 358:529–536, 2003] has claimed that individuals acquire these skills by means of the so‐called program‐level imitation, in which the overall sequence of problem‐solving steps (not the precise actions) is reproduced. In this study we present western lowland gorillas (Gorilla gorilla gorilla) with highly similar nettles. Twelve gorillas in three different groups (including also one nettle‐naïve gorilla) used the same program‐level technique as wild mountain gorillas (with differences mainly on the action level). Chimpanzees, orangutans, and bonobos did not show these program‐level patterns, nor did the gorillas when presented with a plant similar in structural design but lacking stinging defenses. We conclude that although certain aspects (i.e. single actions) of this complex skill may be owing to social learning, at the program level gorilla nettle feeding derives mostly from genetic predispositions and individual learning of plant affordances. Am. J. Primatol. 70:584–593, 2008.

Chimpanzee accumulative stone throwing

The study of the archaeological remains of fossil hominins must rely on reconstructions to elucidate the behaviour that may have resulted in particular stone tools and their accumulation. Comparatively, stone tool use among living primates has illuminated behaviours that are also amenable to archaeological examination, permitting direct observations of the behaviour leading to artefacts and their assemblages to be incorporated. Here, we describe newly discovered stone tool-use behaviour and stone accumulation sites in wild chimpanzees reminiscent of human cairns. In addition to data from 17 mid- to long-term chimpanzee research sites, we sampled a further 34 Pan troglodytes communities. We found four populations in West Africa where chimpanzees habitually bang and throw rocks against trees, or toss them into tree cavities, resulting in conspicuous stone accumulations at these sites. This represents the first record of repeated observations of individual chimpanzees exhibiting stone tool use for a purpose other than extractive foraging at what appear to be targeted trees. The ritualized behavioural display and collection of artefacts at particular locations observed in chimpanzee accumulative stone throwing may have implications for the inferences that can be drawn from archaeological stone assemblages and the origins of ritual sites.

Behavioral variation in gorillas: Evidence of potential cultural traits

The question of whether any species except humans exhibits culture has generated much debate, partially due to the difficulty of providing conclusive evidence from observational studies in the wild. A starting point for demonstrating the existence of culture that has been used for many species including chimpanzees and orangutans is to show that there is geographic variation in the occurrence of particular behavioral traits inferred to be a result of social learning and not ecological or genetic influences. Gorillas live in a wide variety of habitats across Africa and they exhibit flexibility in diet, behavior, and social structure. Here we apply the ‘method of exclusion’ to look for the presence/absence of behaviors that could be considered potential cultural traits in well-habituated groups from five study sites of the two species of gorillas. Of the 41 behaviors considered, 23 met the criteria of potential cultural traits, of which one was foraging related, nine were environment related, seven involved social interactions, five were gestures, and one was communication related. There was a strong positive correlation between behavioral dissimilarity and geographic distance among gorilla study sites. Roughly half of all variation in potential cultural traits was intraspecific differences (i.e. variability among sites within a species) and the other 50% of potential cultural traits were differences between western and eastern gorillas. Further research is needed to investigate if the occurrence of these traits is influenced by social learning. These findings emphasize the importance of investigating cultural traits in African apes and other species to shed light on the origin of human culture.

Acoustic structure and variation in mountain and western gorilla close calls: a syntactic approach

Our understanding of the functioning of a species’ vocal repertoire can be greatly improved by investigating acoustic variation and using objective classification schemes based on acoustic structure. Here we used a syntactic approach to investigate the acoustic structure of the gorilla close distance vocalizations (‘close calls’), which remain as yet little understood. We examined 2130 calls of 10 mountain gorillas (Gorilla beringei beringei) from Bwindi Impenetrable National Park, Uganda, and 5 western lowland gorillas (Gorilla gorilla gorilla) from Bai Hokou, Central African Republic. We segmented calls into units using distinct acoustic features and employed model-based cluster analyses to define the repertoire of unit types. We then examined how unit types were combined into calls. Lastly, we compared unit type use between age–sex classes and the two study groups. We found that the gorilla close calls consist of 5 intergraded acoustic unit types which were flexibly but yet non-randomly concatenated into 159 combinations. Our results are in line with previous quantitative acoustic analyses demonstrating a high degree of acoustic variation in a variety of animal vocal repertoires, particularly close distance vocalizations. Our findings add on to (1) the recent argument that the common practice of describing vocal repertoires as either discrete or graded may be of little value as such distinctions may be driven by human perception and non-quantitative descriptions of vocal repertoires, and (2) recent studies indicating that flexibility in close range social calls can come about through combinatorial systems, which previously have been studied primarily in long distance vocalizations. Furthermore, our study highlights differences in the vocal repertoire of western and mountain gorillas, as expected given differences in environment and social behaviour. Our results offer opportunities for further in-depth studies investigating the function of the gorilla close calls, which will contribute to a more comprehensive understanding of ape vocal communication in general.

Contextual correlates of syntactic variation in mountain and western gorilla close-distance vocalizations: Indications for lexical or phonological syntax?

The core of the generative power of human languages lies in our ability to combine acoustic units under specific rules into structurally complex and semantically rich utterances. While various animal species concatenate acoustic units into structurally elaborate vocal sequences, such compound calls do not appear to be compositional as their information content cannot be derived from the information content of each of its components. As such, animal compound calls are said to constitute a form of phonological syntax, as in the construction of words in human language, whereas evidence for rudimentary forms of lexical syntax, analogous to the construction of sentences out of words, is scarce. In a previous study, we demonstrated that the repertoire of close-distance calls of mountain and western gorillas consists of acoustic units that are either used singularly or non-randomly combined. Here, we investigate whether this syntactic variation provides indications for lexical or phonological syntax. Specifically, we examined the differences between the potential information content of compound calls and their components used singularly through investigating the contexts in which they are used. We found that the gorillas emitted compound calls in contexts similar to some but not all components, but also in a context rarely found for any of their components. As such, the investigated compound calls did not appear to be compositional as their information content cannot be derived from the information content of each of their components. Our results suggest that combining acoustic units into compound vocalizations by gorillas constitutes a form of phonological syntax, which may enable them to increase the number of messages that can be transmitted by an otherwise small repertoire of acoustic units.

Audience effects, but not environmental influences, explain variation in gorilla close distance vocalizations—A test of the acoustic adaptation hypothesis

Close distance vocalizations are an integral part of primate vocal communication. They exhibit large acoustic variation which has been suggested to constitute flexible responses to the highly variable social setting of group living animals. However, a recent study suggested that acoustic variation in close distance calls of baboons may also arise from acoustic adaptations to environmental factors in order to counteract sound degradation. We tested whether the variation in calling rate and acoustic structure of gorilla close distance vocalizations may serve to counteract distorting effects of vegetation during sound propagation. Using focal animal sampling we recorded the vocal behavior of 15 adult individuals living in two groups: one group of western lowland gorillas Gorilla gorilla gorilla and one group of mountain gorillas Gorilla beringei beringei. We considered the distance between the caller and its nearest neighbor as the minimum transmission distance of calls; while vegetation density was quantified through measures of visibility. Our analysis revealed vocal plasticity in gorilla close calls in relation to changes in visibility and nearest neighbor distance. However, the observed changes in fundamental frequency and calling rate are unlikely to counteract degrading effects of vegetation, but rather seem to reflect reactions to variation in spatial and visual separation from other group members, similar to the audience effects demonstrated in a range of other species. We propose that vocal plasticity to counteract distorting environmental effects may not be prevalent across taxa and perhaps confined to species living in heterogeneous habitats with highly variable transmission conditions. Am. J. Primatol. 77:1239–1252, 2015.

Song functions in nonduetting gibbons: Evidence from playback experiments on Javan gibbons (Hylobates moloch)

Territorial, pair-living primates usually perform long-distance calls as duets in which adult males and females coordinate their calls. Previous studies using playback experiments have shown that gibbon duets convey information about the status of the caller (location, familiarity, sex of the caller, and paired status) and gibbons use this information to respond to achieve several nonmutually exclusive functions, including intragroup contact, territorial defense, and pair-bond advertisement and strengthening. However, not all pair-living gibbons duet, and it is unclear whether the same results should be expected in nonduetting species. We conducted song playback experiments (N = 47 trials) to test hypotheses about song functions in nonduetting gibbons on two groups of wild Javan gibbons (Hylobates moloch) in the Gunung Halimun-Salak National Park, Indonesia. Javan gibbons initiated movement toward the speaker more quickly in response to songs broadcast in the center of the territory, stranger songs, and songs of unpaired individuals than to songs at the border, neighbor songs, and songs from paired individuals. These results suggest that Javan gibbons can localize songs, and that Javan gibbon songs transmit information about the identity and paired status of the caller. Our results imply that Javan gibbon solo songs are likely to function for territorial defense and pair-bond advertisement like duets in other primates.