Bernhard Voelkl

Supply and demand determine the market value of food providers in wild vervet monkeys

Animals neither negotiate verbally nor conclude binding contracts, but nevertheless regularly exchange goods and services without overt coercion and manage to arrive at agreements over exchange rates. Biological market theory predicts that such exchange rates fluctuate according to the law of supply and demand. Previous studies showed that primates pay more when commodities become scarcer: subordinates groomed dominants longer before being tolerated at food sites in periods of shortage; females groomed mothers longer before obtaining permission to handle their infants when there were fewer newborns and males groomed fertile females longer before obtaining their compliance when fewer such females were present. We further substantiated these results by conducting a 2-step experiment in 2 groups of free-ranging vervet monkeys in the Loskop Dam Nature Reserve, South Africa. We first allowed a single low-ranking female to repeatedly provide food to her entire group by triggering the opening of a container and measured grooming bouts involving this female in the hour after she made the reward available. We then measured the shifts in grooming patterns after we added a second food container that could be opened by another low-ranking female, the second provider. All 4 providers received more grooming, relative to the amount of grooming they provided themselves. As biological market theory predicts, the initial gain of first providers was partially lost again after the introduction of a second provider in both groups. We conclude that grooming was fine-tuned to changes in the value of these females as social partners.

Upwash exploitation and downwash avoidance by flap phasing in ibis formation flight.

Many species travel in highly organized groups. The most quoted function of these configurations is to reduce energy expenditure and enhance locomotor performance of individuals in the assemblage. The distinctive V formation of bird flocks has long intrigued researchers and continues to attract both scientific and popular attention. The well-held belief is that such aggregations give an energetic benefit for those birds that are flying behind and to one side of another bird through using the regions of upwash generated by the wings of the preceding bird, although a definitive account of the aerodynamic implications of these formations has remained elusive. Here we show that individuals of northern bald ibises (Geronticus eremita) flying in a V flock position themselves in aerodynamically optimum positions, in that they agree with theoretical aerodynamic predictions. Furthermore, we demonstrate that birds show wingtip path coherence when flying in V positions, flapping spatially in phase and thus enabling upwash capture to be maximized throughout the entire flap cycle. In contrast, when birds fly immediately behind another bird—in a streamwise position—there is no wingtip path coherence; the wing-beats are in spatial anti-phase. This could potentially reduce the adverse effects of downwash for the following bird. These aerodynamic accomplishments were previously not thought possible for birds because of the complex flight dynamics and sensory feedback that would be required to perform such a feat. We conclude that the intricate mechanisms involved in V formation flight indicate awareness of the spatial wake structures of nearby flock-mates, and remarkable ability either to sense or predict it. We suggest that birds in V formation have phasing strategies to cope with the dynamic wakes produced by flapping wings.

Inferring social structure from temporal data

Social network analysis has become a popular tool for characterising the social structure of populations. Animal social networks can be built either by observing individuals and defining links based on the occurrence of specific types of social interactions, or by linking individuals based on observations of physical proximity or group membership, given a certain behavioural activity. The latter approaches of discovering network structure require splitting the temporal observation stream into discrete events given an appropriate time resolution parameter. This process poses several non-trivial problems which have not received adequate attention so far. Here, using data from a study of passive integrated transponder (PIT)-tagged great tits Parus major, we discuss these problems, demonstrate how the choice of the extraction method and the temporal resolution parameter influence the appearance and properties of the retrieved network and suggest a modus operandi that minimises observer bias due to arbitrary parameter choice. Our results have important implications for all studies of social networks where associations are based on spatio-temporal proximity, and more generally for all studies where we seek to uncover the relationships amongst a population of individuals that are observed through a temporal data stream of appearance records.

The role of social and ecological processes in structuring animal populations: a case study from automated tracking of wild birds

Both social and ecological factors influence population process and structure, with resultant consequences for phenotypic selection on individuals. Understanding the scale and relative contribution of these two factors is thus a central aim in evolutionary ecology. In this study, we develop a framework using null models to identify the social and spatial patterns that contribute to phenotypic structure in a wild population of songbirds. We used automated technologies to track 1053 individuals that formed 73 737 groups from which we inferred a social network. Our framework identified that both social and spatial drivers contributed to assortment in the network. In particular, groups had a more even sex ratio than expected and exhibited a consistent age structure that suggested local association preferences, such as preferential attachment or avoidance. By contrast, recent immigrants were spatially partitioned from locally born individuals, suggesting differential dispersal strategies by phenotype. Our results highlight how different scales of social decision-making, ranging from post-natal dispersal settlement to fission–fusion dynamics, can interact to drive phenotypic structure in animal populations.

Reproducibility Crisis: Are We Ignoring Reaction Norms?

The ‘reproducibility crisis’ in preclinical biomedical research is making headlines at a staggering rate. Spectacular examples of translational failures and poor reproducibility [1,2] have been attributed to various aspects of poor experimental design and conduct, including small sample sizes [3], risks of bias [4,5], selective reporting [6], and publication bias [7]. In a recent review in this journal, Jarvis and Williams [8] drew a more nuanced picture, challenging the increasingly common view that irreproducibility reflects poor research conduct.

Nonlethal predator effects on the turn-over of wild bird flocks

Nonlethal predator effects arise when individuals of a prey species adjust their behaviour due to the presence of predators. Non-lethal predator effects have been shown to affect social group structure and social behaviour as well as individual fitness of the prey. In this experimental study, we used model sparrowhawks to launch attacks on flocks of wild great tits and blue tits whilst monitoring their social dynamics. We show that nonlethal attacks caused instantaneous turn-over and mixing of group composition within foraging flocks. A single experimental ‘attack’ lasting on average less than three seconds, caused the amount of turn-over expected over three hours (2.0–3.8 hours) of undisturbed foraging. This suggests that nonlethal predator effects can greatly alter group composition within populations, with potential implications for social behaviour by increasing the number of potential interaction partners, as well as longer-term consequences for pair formation and emergent effects determined by social structure such as information and disease transmission. We provide the first evidence, to our knowledge, based on in depth monitoring of a social network to comprehensively support the hypothesis that predators influence the social structure of groups, which offers new perspectives on the key drivers of social behaviour in wild populations.

Reproducibility of preclinical animal research improves with heterogeneity of study samples.

Single-laboratory studies conducted under highly standardized conditions are the gold standard in preclinical animal research. Using simulations based on 440 preclinical studies across 13 different interventions in animal models of stroke, myocardial infarction, and breast cancer, we compared the accuracy of effect size estimates between single-laboratory and multi-laboratory study designs. Single-laboratory studies generally failed to predict effect size accurately, and larger sample sizes rendered effect size estimates even less accurate. By contrast, multi-laboratory designs including as few as 2 to 4 laboratories increased coverage probability by up to 42 percentage points without a need for larger sample sizes. These findings demonstrate that within-study standardization is a major cause of poor reproducibility. More representative study samples are required to improve the external validity and reproducibility of preclinical animal research and to prevent wasting animals and resources for inconclusive research.

Effects of Cage Enrichment on Behavior, Welfare and Outcome Variability in Female Mice

The manner in which laboratory rodents are housed is driven by economics (minimal use of space and resources), ergonomics (ease of handling and visibility of animals), hygiene, and standardization (reduction of variation). This has resulted in housing conditions that lack sensory and motor stimulation and restrict the expression of species-typical behavior. In mice, such housing conditions have been associated with indicators of impaired welfare, including abnormal repetitive behavior (stereotypies, compulsive behavior), enhanced anxiety and stress reactivity, and thermal stress. However, due to concerns that more complex environmental conditions might increase variation in experimental results, there has been considerable resistance to the implementation of environmental enrichment beyond the provision of nesting material. Here, using 96 C57BL/6 and SWISS female mice, respectively, we systematically varied environmental enrichment across four levels spanning the range of common enrichment strategies: (1) bedding alone; (2) bedding + nesting material; (3) deeper bedding + nesting material + shelter + increased vertical space; and (4) semi-naturalistic conditions, including weekly changes of enrichment items. We studied how these different forms of environmental enrichment affected measures of animal welfare, including home-cage behavior (time–budget and stereotypic behavior), anxiety (open field behavior, elevated plus-maze behavior), growth (food and water intake, body mass), stress physiology (glucocorticoid metabolites in fecal boluses and adrenal mass), brain function (recurrent perseveration in a two-choice guessing task) and emotional valence (judgment bias). Our results highlight the difficulty in making general recommendations across common strains of mice and for selecting enrichment strategies within specific strains. Overall, the greatest benefit was observed in animals housed with the greatest degree of enrichment. Thus, in the super-enriched housing condition, stereotypic behavior, behavioral measures of anxiety, growth and stress physiology varied in a manner consistent with improved animal welfare compared to the other housing conditions with less enrichment. Similar to other studies, we found no evidence, in the measures assessed here, that environmental enrichment increased variation in experimental results.

Dynamic network measures reveal the impact of cattle markets and alpine summering on the risk of epidemic outbreaks in the Swiss cattle population

BackgroundLivestock herds are interconnected with each other via an intricate network of transports of animals which represents a potential substrate for the spread of epidemic diseases. We analysed four years (2012–2015) of daily bovine transports to assess the risk of disease transmission and identify times and locations where monitoring would be most effective. Specifically, we investigated how the seasonal dynamics of transport networks, driven by the alpine summering and traditional cattle markets, affect the risk of epidemic outbreaks.ResultsWe found strong and consistent seasonal variation in several structural network measures as well as in measures for outbreak risk. Analysis of the consequences of excluding markets, dealers and alpine pastures from the network shows that markets contribute much more to the overall outbreak risk than alpine summering. Static descriptors of monthly transport networks were poor predictors of outbreak risk emanating from individual holdings; a dynamic measure, which takes the temporal structure of the network into account, gave better risk estimates. A stochastic simulation suggests that targeted surveillance based on this dynamic network allows a higher detection rate and smaller outbreak size at detection than compared to other sampling schemes.ConclusionsDynamic measures based on time-stamped data—the outgoing contact chain—can give better risk estimates and could help to improve surveillance schemes. Using this measure we find evidence that even in a country with intense summering practice, markets continue being the prime risk factor for the spread of contagious diseases.