Biology subjects

McElligott, A. G.

Publications and source records attributed to McElligott, A. G..

14 recordsLinked to original sources

bioRxiv2026

Personality and dominance shape core area use and territory-related signalling in a feral ungulate

Space use strategies determine how animals acquire resources and encounter competitors, which ultimately influence their survival. Instead of utilising their entire range uniformly, animals typically concentrate activity in locations where resources are most valuable, and these 'core areas' provide key insights into the behavioural strategies underlying space use. Although competitive ability is often expected to influence core area use, substantial individual-level variation remains unexplained. Consistent behavioural differences among individuals, or personality, may contribute to this variation, but their role in shaping spatial behaviour under natural conditions is poorly understood. To address this gap, we observed adult males (n=24) from a free-ranging, semi-urban population of Asian water buffalo (Bubalus bubalis). We quantified individual core area size and territory-related signalling behaviour, and assessed personality from >9,300 minutes of observations. Most males (79%) shared overlapping core areas with one to five other males but with clear dominance relationships. Social engagement, reflecting a social personality, was positively associated with core area size in subordinates, but not in dominant males. Social engagement was also positively associated with territory-related signalling frequency. By contrast, a non-social personality labelled as activity-maintenance did not influence with core area size or signalling behaviour. We also found that dominant males exhibited lower signalling frequencies than subordinates. Overall, we show that personality and dominance are associated with distinct aspects of spatial behaviour. These findings demonstrate that individual spatial strategies emerge from the interplay between behavioural phenotypes and social lives, highlighting the spatial-social interface as a source of variation in animal space use.

animal behavior and cognition↗

bioRxiv2026

Drivers of space use in a large semi-urban feral ungulate under seasonally fluctuating resource conditions

Resource scarcity prompts animals to adjust their space use in ways that enhance their survival. In wild herbivores, seasonal habitat shifts are well studied; however, little is known about how large herbivores navigate human-dominated landscapes under fluctuating resource conditions. In Hong Kong, feral water buffalo (Bubalus bubalis; henceforth buffalo) experience declines in body condition score during the dry season. While buffalo expand spatial ranges in the dry season, whether this expansion reflects access to improved ecological conditions, and how key anthropogenic factors (like distance to roads and human density) further influence space use, remains understudied. We observed six buffalo herds (n=92 known individuals) across one wet (July-September 2023) and one dry (January-March 2024) season. We recorded herd locations and extracted remotely sensed habitat and environmental data. We used Normalized Difference Vegetation Index (NDVI) as a proxy of vegetation productivity and included elevation, distance to roads, and human density as additional predictors. We hypothesized that dry-season space use would shift toward higher vegetation productivity, and that anthropogenic factors would have weak influence on space use due to the adaption of buffalo in human-dominated landscapes. We found that buffalo used areas with higher vegetation productivity and higher elevation in the dry season than in the wet. Distance to roads and human density had no detectable effect. Our findings reveal that space use by a large herbivore like buffalo in human-dominated landscapes is strategic and resource driven, and that these seasonal shifts may have important implications for local biodiversity and human-animal interactions.

zoology↗

bioRxiv2025

Domestic goats can follow the direction of human voices to solve a hidden-object task

The capacity for animals to produce and comprehend vocalisations that provide referential acoustic cues to their eliciting cause (e.g. predator-specific alarm calls and food calls) is a highly adaptive means of maximising the benefits of group living, and has been widely studied in diverse species. However, an underexplored dimension of referentiality is the ability to process the direction in which a vocalisation is emitted as a cue towards its referent. Here, we replicated an experimental design previously applied to dogs, chimpanzees and human infants to investigate whether domestic goats (Capra hircus) can use human voices as a directional cue in a hidden-object task. Twenty-nine goats from a UK sanctuary participated in three experimental conditions. In each condition, goats were individually presented with a human experimenter obscured by a barrier, and two identical containers, one of which was baited with food. In the reward directed speech condition, the experimenter vocalised excitedly towards the baited container while sitting closer to the un-baited container, and then the goat was able to select which container to explore. While substantial inter-individual variation existed, on average, subjects chose the baited container at above chance level across the four trials. Two control conditions explored alternative explanations for this result: a "no speech" condition, in which the experimenter was silent but remained in the same location as in the test condition, and a "non-reward directed speech" condition, in which the experimenter directed their voice away from both containers. Subjects showed no evidence of choosing the baited container at above chance level in either of these control conditions. We conclude that goats, like dogs, but not chimpanzees, are capable of attending to the directional cues provided by human voices, and discuss the possible role of domestication in the taxonomic distribution of this ability.

animal behavior and cognition↗

bioRxiv2025

Spatial proximity and dyadic social relationships affect ungulate behavioral synchrony

Collective group decisions are important for the survival and reproduction of social mammals, with inter-individual interactions often driving group-level emergent behavior. Activity synchronization is an important collective behavior, with differences in nutritional requirements leading to foraging asynchrony. Individual variation between animals (such as sex or social relationships) are predicted to affect ungulate synchronization and spatial proximity, with between-sex differences consequently influencing sexual segregation evolution in ungulates. Although investigated independently, the relative roles of sex, sociality and proximity in synchronization are rarely investigated concurrently, especially in regards to affiliative relationships. Asynchrony influences fission-fusion dynamics and social segregation, but little is known how short-term changes in synchrony affects fission. Using a mixed-sex group of feral cattle (Bos taurus), we evaluated the supporting evidence for several predictions arising from the current understanding of synchronization in ungulates. We investigated if sex and social relationships (dominance and affiliation) affected foraging, behavioral synchrony and proximity. We also investigated whether group synchrony affected short-term changes in group size (fission events). We found that same-sex dyads were more likely to be synchronized than mixed-sex dyads, but differences in dominance and affiliation did not affect dyadic synchrony. Focal animals were more synchronized with closest neighbors than with another randomly selected conspecific. Reduction in group size was more likely when group synchrony was lower, highlighting the importance of asynchrony in temporary movement decisions. Inter-individual differences can explain variation in collective behavior, with synchronization being biased towards certain individuals by favoring animals in close spatial proximity and those of the same-sex. LAY SUMMARYIn ungulates, differences in energetic requirements lead to variation in activity, resulting in social and sexual segregation. However, sex, social relationships and spatial proximity are rarely investigated concurrently in relation to synchrony. We investigated synchronization and fission in a feral ungulate relative to individual differences in social relationship and proximity. Sex and proximity affected synchrony and fission events were more likely when synchrony was lower, highlighting the underlying processes in the evolution of sexual segregation.

zoology↗

bioRxiv2025

Reliability and precision of thermal imaging measurements to study animal behaviour and welfare

The use of infrared thermal imaging has become increasingly popular in animal behaviour, health, and welfare research over the last decade. Yet, there is a lack of consensus regarding how this technique should be best applied when measuring peripheral temperatures in animals, including which regions of interest to favour. This fundamental issue necessitates checking the reliability and precision of thermal imaging data when taking repeated measurements, both over short and relatively long time windows. Using goats (Capra hircus) as a model, we investigated two subcategories of reliability, short-term repeatability (measurements taken in the same session) and reproducibility (over multiple sessions), as well as the precision of surface temperatures in two facial regions. We collected data from 20 goats over five measurement sessions over consecutive days. During each session, five frames were collected from approximately one-minute-long videos. From each video, we extracted the mean, maximum, and minimum surface temperatures from the left eye, right eye, and nose tip. To calculate repeatability, we compared temperature variation attributed to differences between goats against total variation in surface temperatures measured in a single session. We defined precision as the temperature deviation within which the mean temperature measured from one to five thermal images was expected to fall in relation to the mean of five image replicates 95% of the time. Reproducibility was investigated by comparing variation attributed to differences in temperature between measurement sessions against total variation in surface temperatures. Our results revealed that repeatability and precision of mean and maximum temperatures across five repeated measurements were high for all facial regions, with between 93.50% and 99.81% of total temperature variation attributable to the individual goat tested. Conversely, minimum temperatures were more variable, less repeatable, and less precise. For reproducibility, measurement sessions accounted for a high proportion of variation in nasal temperatures (74.61-85.85%), and a lower, but substantial proportion of eye temperature variation (49.59-67.01%). We conclude that mean and maximum thermal measures show promise for quantifying nasal and eye temperatures in the short term. However, surface temperature measured across several days was not readily comparable, highlighting the importance of considering ambient conditions in thermal imaging research. Overall, this study provides valuable insights into the appropriate use of thermal imaging in goats and, more broadly, animal behaviour and welfare research.

animal behavior and cognition↗

bioRxiv2025

Adaptive behavioural strategies to seasonal challenges by a semi-urban feral ungulate

Seasonality, or temporal shifts between wet and dry seasons, profoundly affects the behavioural ecology of animals, particularly large herbivores, in (sub-) tropical climatic conditions. Behavioural strategies are crucial for overcoming challenges associated with seasonality. Group-level strategies to seasonality in the wild have received considerable attention, yet little is known about how large herbivores in human-dominated landscapes respond to seasonal challenges. Moreover, focusing solely on groups can obscure how individual animals of different sexes, ages, and personalities respond. We investigated a semi-urban feral population of a large ungulate, the water buffalo (Bubalus bubalis) (N=71) during a wet and a dry season. Individual body condition was non-invasively assessed to evaluate whether seasonality impacts physiology. To identify behavioural strategies, we collected data on feeding (grazing and browsing) and space use (core area and home range). Further, using a subset (n=30), we investigated the associations between personality and space use in the dry season. Body condition scores were highest during the wet season and then declined during the dry season, indicating an impact of seasonality on physiology. Older individuals were more affected than younger ones. Browsing increased during the dry compared to the wet season. While there was no change in core area use, home ranges expanded in the dry compared to the wet season. Regardless of season, females utilised home ranges larger than males. Females exhibiting higher social tension personalities had smaller core areas and home ranges than those with lower social tension. We show that buffalo employ behavioural strategies to overcome challenges associated with seasonality, and these strategies are contingent on intrinsic factors: sex, age, and personality. Our findings offer insights into the ecological principles -habitat use, resource utilisation, and competition- that govern the behavioural ecology of herbivores, and have implications for conservation and welfare in increasingly human-dominated and climate-sensitive landscapes.

zoology↗

bioRxiv2025

Sexual dimorphism and phenotypic plasticity in a subtropical feral bovid Bos taurus (Artiodactyla: Bovidae).

Phenotypes reflect how organisms adapt to their environments. Hong Kong (HK) feral cattle, a crossbreed of Bos taurus taurus and Bos taurus indicus, present an opportunity to study these adaptations in one of the very few global cattle populations not directly controlled by humans. These cattle are free-ranging since their release from farms in the 1970s. HK has a subtropical climate, characterized by high humidity and temperatures during the wet season, and scarce precipitation during the dry season. We studied seasonal coat colour changes in HK feral cattle, and sexual dimorphism in body size and horn length. We provide the first evidence of seasonal changes in coat colour in cattle, with paler coats being more common in the wet season, while darker coats prevailed in the dry season. These seasonal changes were influenced by temperature, humidity, solar radiation, and body condition. We found that males were larger and had longer horns than females. Our results show a male-biased sex dimorphism in the HK feral cattle. Additionally, our findings suggest that thermoregulation costs drive colouration in these cattle. The phenotypic plasticity we demonstrate in these subtropical feral cattle improves our knowledge of the adaptations of ungulates to their habitat.

zoology↗

bioRxiv2024

Personality homophily drives female friendships in a feral ungulate

Friendships, exhibited by both humans and non-human animals, have considerable adaptive benefits. In humans, similarity or homophily in personality is considered a proximate mechanism driving friendships, yet little is known about the behavioural decision rules underlying animal friendships. Some empirical research suggests that animal friendships can be driven by personality homophily. However, these studies are restricted to non-human primates, limiting our understanding of the mechanisms of friendships. We investigated a feral population of water buffalo (Bubalus bubalis) to determine whether homophily in personality drives friendships in this non-model social species in free-ranging environmental conditions. We conducted observations on females (n=30) and assessed their friendships and personalities. Close spatial proximity served as a behavioural indicator of friendship, validated by affiliative body contact. An objective bottom-up method revealed three personality traits - social tension, vigilance, and general dominance. We found that females with comparatively lower personality differences in social tension and general dominance traits exhibited significantly higher close spatial associations. We did not find an effect of kinship on close spatial associations. Our findings show that friendships in buffalo can form based on personality homophily, a decision rule attributed predominantly to primates. We discuss these findings in light of buffalo socioecology but emphasise their implications in the broader evolutionary context of animal personalities and friendships.

animal behavior and cognition↗

bioRxiv2023

Do Goats Recognise Humans Cross-Modally?

Recognition plays a key role in the social lives of gregarious species, enabling animals to distinguish among social partners and tailor their behaviour accordingly. As domesticated animals regularly interact with humans, as well as members of their own species, we might expect mechanisms used to discriminate between conspecifics to also apply to humans. Given that goats can combine visual and vocal cues to recognize one another, we investigated whether this cross-modal recognition extends to discriminating among familiar humans. We presented 28 goats with facial photographs of familiar people and two repeated playbacks of a voice, either congruent (from the same person) or incongruent with that photograph (from a different person). When cues were incongruent, violating their expectations, we expected goats to respond faster and for longer after playbacks and show increases in physiological arousal (increased heart rate and/or decreased heart rate variability). We found the increase in latency that goats took to respond as the playback series progressed was greater when the face and voice were incongruent. As differences were not as predicted and only observed in one response measured, our evidence is tentative, but the variation in latency to look between congruency conditions suggests goat cross-modal recognition extends to humans. If this is the case, not only would this further demonstrate the flexibility of complex recognition systems to discriminate among members of a very different species, but indicates goats can produce mental representations for familiar people, a key component of individual recognition.

animal behavior and cognition↗

bioRxiv2023

Goat Discrimination of Emotional Valence in the Human Voice

Reading another animals emotional state can enable receivers to anticipate their behavioural motivations, which is important in guiding interactions with that individual. For species living closely alongside humans, the emotional cues that people express can be almost as informative as those of conspecifics. Goats can discriminate differences in emotional valence present in another goats calls, and we investigated whether this ability extends to human speech. We presented goats with a habituation-dishabituation-rehabituation paradigm, where they experienced multiple playbacks of a familiar or unfamiliar human voice conveying a single emotional valence (e.g., angry; habituation phase), before the valence of the voice changed (e.g., happy; dishabituation phase) and then reversed again in-line with the habituation phase (e.g., angry; rehabituation phase). Over the habituation phase, goat behavioural responses decreased, showing evidence of having habituated to the playback stimuli presented. Following a change in emotional valence (dishabituation phase), although goats were overall less likely to respond, those that did looked for longer, suggesting they had perceived the shift in emotional content conveyed in human voice playbacks. We found no changes in physiological arousal (heart rate or heart rate variability) with shifts in playback valence. Goats, as a domesticated species, may have developed a sensitivity to our cues over their long association with humans, but the differences in individual behaviour towards the playback paradigm could highlight a role for learning and individual experience in particular on interspecific emotional communication.

animal behavior and cognition↗

bioRxiv2023

Sex and dominance status affect allogrooming in free-ranging feral cattle

Social interactions are fundamental properties of gregarious species, helping to establish dominance hierarchies and maintain social bonds within groups, thus having significant effects on fitness. Cattle (Bos taurus) are social ungulates which engage in affiliative and agonistic relationships with other individuals. Although there are approximately 1.5 billion cattle on the planet, the opportunity to research cattle behaviour in free-ranging groups is rare, as there are few feral populations worldwide. Cattle engage in positive social behaviours such as allogrooming, where one individual licks the body of another. The relationship between affiliative behaviours and other individual characteristics (such as sex and dominance status) are frequently studied in other gregarious species, but are largely undetermined in cattle. To investigate the relationships between sex, dominance status and allogrooming, we observed a mixed-sex feral cattle herd in Hong Kong, recording dominance interactions and allogrooming events. We found that dominant females received more allogrooming than subordinate females, but subordinate females did not perform more than dominant ones. Males performed allogrooming more towards females than other males, but females groomed both sexes equally. Sex affected dominance position, with males obtaining higher status than females, but not all females were subordinate to males. These preferential allogrooming patterns improve our knowledge of sex-specific interactions, and help us to understand the dynamics of agonistic and affiliative behaviours in multi-male, multi-female ungulate groups. Studying a free-ranging feral population provides us with a unique insight into ungulate behavioural patterns and the evolution of cattle social behaviours.

animal behavior and cognition↗

bioRxiv2022

Delayed benefits for fallow bucks: more fights decrease same day mating success, but may increase matings the next day

Dominance hierarchies help to reduce unnecessary fights and associated costs during the mating season. Fallow deer (Dama dama) typically have high levels of male-male competition and strong reproductive skew. Nevertheless, how male dominance and daily fight rates affect mating success remains unknown. We used a two-year dataset from a large population of tagged fallow deer (620-689 individuals), to calculate male dominance ranks based on their agonistic interactions prior to the mating season ( prerut), in order to then examine how rank is related to fight rates and mating success during the mating season ( rut). Overall, higher-ranked males fought at least twice a day on a higher proportion of days during the rut and secured more matings. Males engaging in more than 10 fights per day were less likely to secure a mating that same day, and those males exceeding 15 fights per day secured no matings at all. Nevertheless, males with the highest numbers of fights (i.e. 15-21 fights per day) on a given day had higher mating success on subsequent days. Although higher-ranked males secured most matings during the rut, their fight rates decreased towards the end. We propose that engaging in more fights negatively affects daily individual mating success, but may benefit mating success on subsequent days, and potentially increase long-term fitness benefits. Additionally, engaging in more fights as the rut progresses probably allows lower-ranked males to secure some matings before the availability of oestrous females ends for almost a year. SIGNIFICANCE STATEMENTFighting carries a risk of injury and high energetic costs. Male fallow deer establish dominance hierarchies, that help reduce unnecessary fights among individuals of different competitive abilities. However, whether high-ranked males fight more or less is yet unknown. By calculating social ranks of fallow bucks before the start of their mating period, we show that males of higher social status do fight and mate more during the mating season (rut). Furthermore, by investigating how investment in fights affects individual mating success that same day and the next day, we find that males that fight more cause a decrease in their immediate daily mating success, but can potentially increase their chances of mating in subsequent days. Thus, to fight more may allow males to climb the hierarchical social ladder, hence increasing longer-term fitness benefits associated to higher ranks.

animal behavior and cognition↗

bioRxiv2022

Goat kid recognition of their mothers' calls is not impacted by changes in source-filter parameters.

Features varying more between than within individuals are usually considered as potential cues for individual recognition. According to the source-filter theory of vocal production, the fundamental frequency of mammals vocalisations depends on the characteristics of the vocal folds, while formants are determined by the characteristics of the vocal tract. Goat mothers and their kids (Capra hircus) display mutual recognition, and both source-related parameters (F0) and filter-related ones (formants) have been shown to be individualised in their vocalisations. Here, we aimed to identify if these parameters (source-related parameters (F0) and/or filter vocal parameters) are used by goat kids to recognise their mothers vocalisations. To this aim, we used an algorithm to modify either F0 or formants of the calls of goat mothers to different degrees (within or exceeding the range of natural intra-individual variability), and we played back these modified calls to their kids. We did not observe any difference in the kid reactions to the modified maternal vocalisations and to the natural calls. We suggest that either: (i) fundamental frequency and formants are not involved in maternal recognition in goats; (ii) goat kids have a tolerance for variation when recognising their mothers calls that exceeds the shifts we performed; (iii) goat maternal recognition is based on other vocal features than those tested here, or (iv) goat kid maternal recognition is based on a combination of different features and might be more flexible than previously thought, such that when one main feature is modified, kids focus on other features.

animal behavior and cognition↗

bioRxiv2022

Welfare concerns for mounted load carrying by working donkeys in Pakistan

Working donkeys (Equus asinus) are vital to peoples livelihoods. They are essential for carrying goods, however globally, overloading is one of the primary welfare concerns of working donkeys. We studied mounted load carrying by donkeys and associated factors in Pakistan. A cross-sectional study of donkey owners (n = 332) was conducted, and interviews were undertaken based on a questionnaire. Owners estimated that the median weight of their donkeys was 110kg (interquartile range (IQR) 100-120kg), and that they carried a median mounted load of 81.5kg (IQR 63-99kg). We found that 87.4% of donkeys carried a load above 50% of their bodyweight ratio (BWR), the median BWR carried was 77.1% (IQR 54.5-90.7%), and 25.3% of donkeys carried above 90% BWR. Donkeys that were loaded at more than 50% BWR were more likely to sit, compared to donkeys loaded with less weight (p=0.01). Donkeys working in peri-urban and urban areas were more likely to carry a greater BWR than donkeys working in rural areas (P<0.001), as were those carrying construction materials or bricks, compared to agricultural materials (p=0.004). Age (p=0.03) and breed (p=0.01) were also associated with carrying a higher weight. Overloading based on current recommendations (50% BWR) was common, with the majority (87.4%) of donkeys reported to carry more than the recommended 50% limit. This survey provides evidence of on-the-ground working practices and factors associated with mounted load carrying, which is critical for developing evidence-based recommendations for loading, in order to improve the welfare of working donkeys.

scientific communication and education↗