A protocol for assessing bias and robustness of social network metrics using GPS based radio-telemetry data.

BACKGROUND: Social network analysis of animal societies allows scientists to test hypotheses about social evolution, behaviour, and dynamic processes. However, the accuracy of estimated metrics depends on data characteristics like sample proportion, sample size, and frequency. A protocol is needed to assess for bias and robustness of social network metrics estimated for the animal populations especially when a limited number of individuals are monitored. METHODS: We used GPS telemetry datasets of five ungulate species to combine known social network approaches with novel ones into a comprehensive five-step protocol. To quantify the bias and uncertainty in the network metrics obtained from a partial population, we presented novel statistical methods which are particularly suited for autocorrelated data, such as telemetry relocations. The protocol was validated using a sixth species, the fallow deer, with a known population size where ∼ 85 % of the individuals have been directly monitored. RESULTS: Through the protocol, we demonstrated how pre-network data permutations allow researchers to assess non-random aspects of interactions within a population. The protocol assesses bias in global network metrics, obtains confidence intervals, and quantifies uncertainty of global and node-level network metrics based on the number of nodes in the network. We found that global network metrics like density remained robust even with a lowered sample size, while local network metrics like eigenvector centrality were unreliable for four of the species. The fallow deer network showed low uncertainty and bias even at lower sampling proportions, indicating the importance of a thoroughly sampled population while demonstrating the accuracy of our evaluation methods for smaller samples. CONCLUSIONS: The protocol allows researchers to analyse GPS-based radio-telemetry or other data to determine the reliability of social network metrics. The estimates enable the statistical comparison of networks under different conditions, such as analysing daily and seasonal changes in the density of a network. The methods can also guide methodological decisions in animal social network research, such as sampling design and allow more accurate ecological inferences from the available data. The R package aniSNA enables researchers to implement this workflow on their dataset, generating reliable inferences and guiding methodological decisions.

Do human-wildlife interactions predict offspring hiding strategies in peri-urban fallow deer?

Human activities can induce significant behavioural changes in wildlife. Often explored through extractive interactions (e.g. hunting) that can favour certain behavioural traits, the implications of non-extractive ones, such as wildlife feeding, remain understudied. Research shows that people tend to favour bolder individuals within populations despite their dynamics and consequences being unclear. Using fallow deer in a peri-urban environment, we studied whether mothers that show reduced fear of humans and consistently approach them for food adopt weaker anti-predator strategies by selecting less concealed fawning bedsites closer to human hotspots. This would provide the advantage of additional feeding opportunities in comparison with shyer mothers while keeping their fawns close. Our dataset encompassed 281 capture events of 172 fawns from 110 mothers across 4 years. Surprisingly, mothers that regularly accepted food from humans selected more concealed bedsites farther from human hotspots, giving their offspring better protection while also benefitting from additional food during lactation. Our results show behavioural adaptations by a subset of females and, for the first time, link the tendency to approach humans and strategies to protect offspring. Given previous findings that these begging females also deliver heavier fawns at birth, our research further investigates human-wildlife feeding interactions and their behavioural implications.

Does artificial feeding impact neonate growth rates in a large free-ranging mammal?

Variation and disparity in resource access between individuals in an animal population within human-dominated landscapes require attention as artificial selection processes may be at work. Independent, recreational human-wildlife feeding interactions constitute an increasingly prevalent, yet understudied, food resource for birds and mammals living in our cities. However, only a limited number of risk-taking individuals may access it. Using urban fallow deer as our model species, we hypothesized that if these interactions result in positive effects for the engaging individual, e.g. increased milk quality and yield, then this would result in the increased growth rates of their offspring. Alternatively, if these individuals were prioritizing investing time in engagement with humans, resulting in decreased maternal care, then this would result in slower growth rates in their offspring. We found that the offspring of those females that regularly interacted with humans displayed significantly faster growth rates than their risk-adverse counterparts. This advantage for fearless mothers in terms of boosted neonatal growth rates could be mirrored in birds accessing garden feeders, seagulls or pigeons utilizing urban resources, or seals approaching city harbours. Here, we add a new piece to the complex puzzle of how humans are impacting wildlife living within human-dominated landscapes.

Neonate personality affects early-life resource acquisition in a large social mammal.

Although it is widely acknowledged that animal personality plays a key role in ecology, current debate focuses on the exact role of personality in mediating life-history trade-offs. Crucial for our understanding is the relationship between personality and resource acquisition, which is poorly understood, especially during early stages of development. Here we studied how among-individual differences in behavior develop over the first 6 months of life, and their potential association with resource acquisition in a free-ranging population of fallow deer (Dama dama). We related neonate physiological (heart rate) and behavioral (latency to leave at release) anti-predator responses to human handling to the proportion of time fawns spent scanning during their first summer and autumn of life. We then investigated whether there was a trade-off between scanning time and foraging time in these juveniles, and how it developed over their first 6 months of life. We found that neonates with longer latencies at capture (i.e., risk-takers) spent less time scanning their environment, but that this relationship was only present when fawns were 3-6 months old during autumn, and not when fawns were only 1-2 months old during summer. We also found that time spent scanning was negatively related to time spent foraging and that this relationship became stronger over time, as fawns gradually switch from a nutrition rich (milk) to a nutrition poor (grass) diet. Our results highlight a potential mechanistic pathway in which neonate personality may drive differences in early-life resource acquisition of a large social mammal.

Risk-taking neonates do not pay a survival cost in a free-ranging large mammal, the fallow deer (Dama dama).

Recent debate has focused on whether variation in personality primarily reflects variation in resource allocation or resource acquisition of individuals. These two mechanisms predict different relationships between personality and survival. If personality mainly reflects variation in resource allocation, then bold (i.e. risk-taking) individuals are expected to live shorter lives, whereas the opposite pattern is expected with resource acquisition. Here we studied the relationship between neonate personality and early-life survival in 269 juveniles of a population of fallow deer (Dama dama). We found that bolder individuals paid no apparent survival cost. Interestingly, among-individual differences in the physiological response at capture (heart rates, which covary with the behavioural response, i.e. latency to leave) were linked to survival, where individuals with lower heart rates when handled by humans had a higher probability of early-life survival. This suggests that bolder individuals may be of higher state than their shyer counterparts. As the first study linking neonate personality to survival in a free-ranging mammal, we provide novel insights into drivers behind early-life individual variation.

Artificial selection in human-wildlife feeding interactions.

The artificial selection of traits in wildlife populations through hunting and fishing has been well documented. However, despite their rising popularity, the role that artificial selection may play in non-extractive wildlife activities, for example, recreational feeding activities, remains unknown. If only a subset of a population takes advantage of human-wildlife feeding interactions, and if this results in different fitness advantages for these individuals, then artificial selection may be at work. We have tested this hypothesis using a wild fallow deer population living at the edge of a capital city as our model population. In contrast to previous assumptions on the randomness of human-wildlife feeding interactions, we found that a limited non-random portion of an entire population is continuously engaging with people. We found that the willingness to beg for food from humans exists on a continuum of inter-individual repeatable behaviour; which ranges from risk-taking individuals repeatedly seeking and obtaining food, to shyer individuals avoiding human contact and not receiving food at all, despite all individuals having received equal exposure to human presence from birth and coexisting in the same herds together. Bolder individuals obtain significantly more food directly from humans, resulting in early interception of food offerings and preventing other individuals from obtaining supplemental feeding. Those females that beg consistently also produce significantly heavier fawns (300-500 g heavier), which may provide their offspring with a survival advantage. This indicates that these interactions result in disparity in diet and nutrition across the population, impacting associated physiology and reproduction, and may result in artificial selection of the begging behavioural trait. This is the first time that this consistent variation in behaviour and its potential link to artificial selection has been identified in a wildlife population and reveals new potential effects of human-wildlife feeding interactions in other species across both terrestrial and aquatic habitats.

Variations in the Behavior of Pigs During an Open Field and Novel Object Test.

Tail biting is a serious welfare concern in pig production. It not only causes distress for victims, but may occur where pigs are unable to cope, and become biters. An animal's ability to cope with stressful situations may vary between individuals, but the behavioral response could be consistent across different fear eliciting situations. We exposed 75 pigs to open field (OF) and novel object (NO) tests at 14 weeks of age. Within each pen of pigs (n = 16 pens, 55 pigs/pen), 6 pigs were selected for testing using the following criteria: 3 pigs that had severe bite wounds (BITTEN), 1 confirmed biter (BITER), 1 pig which could be easily approached and trained to provide a saliva sample (BOLD) and 1 pig which was extremely evasive, and was unable to be trained to willingly provide a saliva sample (SHY). Given that responses may be consistent in different scenarios, we hypothesized that SHY pigs would display more characteristics of a fear response (i.e., less movement in the open field, more time spent by the door, and longer latency to approach the novel object) than human BOLD pigs. We also hypothesized that BITTEN pigs would behave similarly to SHY and BITERS similarly to BOLD. The BOLD and BITER pigs spent more time exploring (P < 0.05) and less time by the door (P < 0.01) than the BITTEN and SHY pigs. Although there was an overall increase in cortisol level from before to after the tests (P < 0.001), this was only significant for BITTEN (P < 0.001) and SHY (P < 0.05) pigs. Therefore, as hypothesized, for several measures, BOLD, and BITER pigs behaved similarly, and differently to SHY and BITTEN. However, the low sample size potentially meant that for several measures, although numeric differences were in the direction hypothesized, there were no statistical differences. Further work in which a greater number of BITER pigs were included in the sample, may elucidate our hypotheses more clearly, as to whether responses to fear tests in pigs could be associated with the likelihood of being a tail biter, or victim.

Irish pig farmer's perceptions and experiences of tail and ear biting.

Abnormal behaviours such as ear and tail biting of pigs is of significant welfare and economic concern. Currently, pig welfare legislation is under renewed focus by the EU commission and is likely to be enforced more thoroughly. The legislation prohibits routine tail docking and requires adequate enrichment to be provided. In Ireland, tail-docking is still the most utilised control mechanism to combat tail biting, but biting is still widespread even in tail-docked pigs. In addition, as pig farms are almost all fully slatted, bedding type material cannot be provided. Thus, the opinions, and practices of farmers in countries like Ireland, which may need to make significant adaptations to typical pig management systems soon, need to be considered and addressed. We carried out a survey of pig farmers during 2015 in order to gain a greater understanding of the extent of biting on Irish farms, perception on the most important preventive measures, current enrichment use and actions following outbreaks. Fifty-eight farmers from 21 Counties responded with an average herd size of 710 ± 597 sows (range 90-3000 sows). Only two farms had experienced no biting in the last year. Of the farms that had experienced tail biting (88%), 86% had also experienced ear biting. The most common concerns relating to biting were condemnation and reduced productivity of bitten pigs with both receiving an average score of 4 (most serious). Ear biting occurred most commonly in the 2nd stage (approximately 47-81 days from weaning) weaner and tail biting in the finishing stage. The most important preventive measures were felt to be taking care of animal health, restricting density, maintaining an even quality of feed/content and maintaining good air movement. Sixty-five percent of respondents added additional enrichment following an outbreak. Chains were the most common form of enrichment currently used (83%). Those not using chains favoured wood, toys and rope (17%). Identification of the most effective and accessible control and prevention measures both for the animals and for the farming community is thus essential. Improved understanding of the concerns and practices of producers, which this survey contributes to, is a first step towards this aim.

The effects of environmental and visitor variables on the behavior of free-ranging ring-tailed lemurs (Lemur catta) in captivity.

The effect of the zoo environment on captive animals is an increasingly studied area of zoo research, with visitor effects and exhibit design recognized as two of the factors that can contribute to animal welfare in captivity. It is known that in some situations, visitors may be stressful to zoo-housed primates, and this may be compounded by environmental factors such as the weather, the time of day, and zoo husbandry routines. Exhibit design and proximity of the public are also known to influence behavioral response of primates to visitors; however, there is minimal research on free-ranging zoo animals, even though they are potentially subjected to intense interactions with visitors. The current study explores the effect of the zoo environment, several visitor variables and specific animal-visitor interactions on the behavior of free-ranging ring-tailed lemurs (Lemur catta) at Fota Wildlife Park, Ireland. Data were obtained through scan samples collected over 18 months (n = 12,263) and analyzed using a range of statistical tests, including general estimating equations (GEE). Results demonstrate that the free-ranging lemurs' behavior at Fota Wildlife Park is affected by season, weather and time of day. Similarities in feeding behavior exist between the free-ranging group and lemurs in the wild when resources are plentiful. Visitor variables had a limited effect on lemur behavior and behavioral diversity level. Lemurs rarely reacted to visitors when specific interactions were considered. Generally, the results indicate that the ring-tailed lemurs in this study have adapted well to the zoo environment and habituated to visitors.