Animal Behaviour

To make adaptive foraging decisions, naive individuals need to gather information about the local prey community. Besides sampling prey personally, the young could gather information about prey profitability by observing the foraging behaviour of other individuals, and parental provisioning provides the first opportunity to acquire this social information. Still, previous research on vertical transmission of prey preferences from parents has provided mixed results that are often confounded with other information sources, such as siblings and peers. It is also not known whether information from parents can change potential innate biases against certain prey types, such as avoidance of warningly coloured insects. Here, we tested whether social information acquired by offspring during parental provisioning influences the development of prey preferences in a generalist predator, the Great Tit (Parus major). We brought 15 great tit broods and their parents into captivity at late nestling stage (14 days old) and divided them into three social information treatments where parents were provided with either brown, red or yellow palatable maggots to feed to their dependent young for 8 days. Once foraging independently from parents, we conducted a preference test where juveniles were offered the full array of coloured maggots. Regardless of palatable exposure to typical warning-coloured maggots (i.e. red and yellow), juveniles consistently preferred yellow over red, and preferred brown maggots the most (i.e. lacking warning coloration). This supports the existence of innate biases against typical warning colours, and that social information from parents is unlikely to override these, at least when alternative prey is easily available.

Approaching conspecifics is a fundamental component of social interactions. Such decisions are typically investigated through the study of social relationships (e.g., kinship, dominance, affiliation) and their impact. In the framework, the role of social affordances, i.e., interaction opportunities arising from the immediate social environment, remains poorly understood. We investigated how context-dependent factors shape approach decisions in wild crested macaques (Macaca nigra) in North Sulawesi, Indonesia. From September 2024 to May 2025, we collected data on 27 adults (20 females, 7 males) using focal observations and an approach-specific protocol recording neighbours identities and behaviours at the initiation of close-proximity approaches (<1 body length). We first compared data collected during approaches with focal data, then conducted within-approach analyses contrasting approached partners and bystanders. We further tested the effects of behavioural state, subgroup size and composition, including infant presence. Bayesian models revealed that affiliative, neutral, and grooming behaviours were more frequent in approach contexts than in general observations. Within approaches, individuals avoided congeners already engaged in grooming and approached individuals in neutral or affiliative states (e.g., lip-smacking, embracing). Macaques globally approached more isolated individuals or dyads. When multiple subgroups were present (i.e., multiple choices), the likelihood of being approached increased with subgroup size and was higher in subgroups containing infants. These findings indicate that macaques use behavioural cues and local social density as affordance signals, adjusting approach decisions to maximize interaction opportunities and navigate complex social environments.

Growing evidence of animals combining discrete, meaningful calls into sequences--a feature once thought unique to linguistic syntax--has presented the opportunity to investigate the evolutionary origins of syntactic communication. The arbitrary assignment of meaning to words marks an important step in human language evolution, and a necessary precursor to generating further meaning through sentences. Studying how other animals that produce call sequences learn the meaning of these signals could help shed light on how referentiality and semantic combinatoriality evolved. Given the presence of meaningful call sequences has only recently been revealed in several non-human animals, ontogenetic studies of the comprehension of these vocalisations are, to date, non-existent. Western Australian magpies (Gymnorhina tibicen dorsalis) combine discrete calls into a diverse array of call sequences. Recent evidence shows these sequences are socially learned, but the developmental stage at which fledglings respond correctly to them remains unstudied. We performed playbacks of a discrete alarm call and call sequence to fledglings over the course of their first 18 weeks out of the nest, identifying when they differentiate between the low-level disturbance associated with the discrete call and the high-grade aerial threat associated with the sequence. Fledglings showed immediate vigilance to both vocalisations but exhibited significantly greater vigilance and upward scanning following the sequence. Critically, fledglings showed this response to the sequence from the first week of testing, with no effect of age on the response to either vocalisation. These findings suggest that comprehension precedes production of sequences in magpies and that sequence meanings are either learned rapidly or have an innate basis. While further investigation is essential, this study offers the first empirical insight into the ontogenetic emergence of combinatorial comprehension in a non-human animal.

Human-influenced environments pose challenges but also provide wildlife with anthropogenic resources. Individuals vary widely in their ability to exploit such resources, often as a function of behavioural type. However, we lack a clear understanding of how variation in behavioural traits influences stages of resource exploitation required to use anthropogenic resources. Using fully automated foraging puzzles, we examined how boldness and sociability influenced three aspects of resource exploitation - discovery, problem-solving, and overall performance - in two wild populations of California ground squirrels (Otospermophilus beecheyi). Bolder individuals discovered the resource earlier, solved the task faster and achieved higher performance, indicating that boldness promotes efficient exploitation of anthropogenic resources across multiple stages in the process. Greater sociability and more opportunities to observe conspecifics solving the task led to faster problem-solving, consistent with evidence for observational learning. Squirrels in the recreational-use population - with regular exposure to humans and anthropogenic food - were faster to discover the resource than those in a trail-use population - where human exposure was transient and no anthropogenic food available. Problem-solving latency and performance were consistent between populations. Our findings highlight how individual variation in behavioural traits drives performance in novel ecological contexts, providing a mechanistic understanding of behavioural plasticity in human-influenced environments.

As foraging increases exposure to predators, individuals must allocate time to vigilance in addition to food acquisition. Although those behaviors are assumed to be mutually exclusive, this is often hard to disentangle, given that multiple sensory modalities are often applied. However, when foraging, animals often have one sensory modality more impaired (e.g. vision), than others (e.g. auditory); thus, a level of compromise on vigilance capabilities is assumed while foraging. Here, we test the idea that when a food item requires manipulation, thus reducing vigilance capabilities, social foraging should promote shared costs of vigilance. We conducted video recordings to investigate whether there is a trade-off between foraging and vigilance behavior in red-legged seriemas (Cariama cristata) when foraging in pairs. We studied the social foraging behavioral patterns while seriemas were feeding on macauba palm (Acrocomia aculeata) fruit, which requires a series of throwing maneuvers to crack the hard outer shell. Our results showed that within a pair recording one individual was observed to spend more time vigilant than their partner, while the other individual spent more time throwing fruit than their partner. However, we found no evidence for synchronization or coordination of vigilance and foraging between individuals within a feeding bout. These results provide insight into the foraging ecology of red-legged seriemas and the trade-off between foraging and vigilance in social contexts.

Behaviour innovation plays a pivotal role in a species adaptability to dynamic environments. Investigating innovative behaviour and its underlying mechanism is therefore crucial for elucidating the development of cognitive flexibility across animals. Animal personality--which shapes how individuals perceive and engage with their surroundings--could offer insights into individual variation in this process. This study used a three-step foraging puzzle to evaluate the innovation capacity in 28 rosy-faced lovebirds (Agapornis roseicollis), specifically examining their capacity to recombine individually innovated component behaviours into integrated, more sophisticated techniques. We found that nearly half of the individuals spontaneously innovated multiple component behaviours to solve novel puzzles. Crucially, when challenged with a more sophisticated task, they recombined these behaviours into functionally dependent sequences without prior social demonstration. We further identified sex, persistence, and asocial learning capacity as key predictors of innovative problem-solving performance, with females, persistent individuals, and superior asocial learners excelled at problem-solving. Our findings demonstrate that behavioural innovation is not a static event, but a dynamic process--modulated by physical, cognitive, and personality variables--in which behaviours are flexibly transferred and recombined into increasingly complex forms to enable rapid individual adaptation.

For many group living species, vocal signals are a vital form of communication needed to coordinate social behaviours. Investigating these processes first requires a detailed description of the species vocal repertoire. White-nosed coatis (Nasua narica), which forage and move together in forest habitats, are thought to rely on vocalisations to coordinate movements and maintain group cohesion. However, quantitative studies of their vocal repertoire are lacking. We examined the vocal repertoire of white-nosed coatis from wild populations in Panama and Arizona, USA, to gain a more comprehensive view of their calling behaviours. By combining traditional acoustic analyses with an unsupervised approach based on spectrogram structure, we characterised the diversity of calls in this species and described the temporal and structural features of their vocalisations. We identified 19 call types, with some of these calls emitted in multi-syllable call sequences or in fast succession. We found variability in call rates among group members, which may be driven by differences in social status within the group. In addition, our results indicate that white-nosed coatis likely have individually-recognisable vocalisations. This study provides a foundational description of the white-nosed coati vocal repertoire, laying the groundwork for future research on vocal communication in this species.

Chorus howls play a central role in wolves' (Canis lupus) social communication, shaping both intra- and inter-pack interactions. While synchrony has been shown to influence perceived threat from outgroups in birds and humans, whether the synchrony of an outside pack's chorus howl plays a role in wolf pack behaviour remains unclear. This study investigated behavioural responses of 19 zoo-housed wolf packs to playbacks of synchronous and asynchronous chorus howls from unfamiliar packs, in comparison to a dove control call. We hypothesized that packs would show stronger intra-pack cohesion and greater spatial withdrawal in response to more synchronous, and thus cohesive - and potentially more formidable - sounding vocalisations. Alternatively, asynchronous howls might elicit stronger responses due to their ambiguity and the "Beau Geste" effect (where asynchronous signals make it harder to infer the number of callers). Synchronous howls resulted in more distance from the sound source and less activity compared with asynchronous howls. This could be interpreted as a more cautious response towards a perceived threat. However, since there was no effect on pack-directed behaviour and no difference between the synchronous and control sound, lower interest or motivation to engage with the stimulus cannot be excluded. In contrast, wolves spent more time near the sound-source after hearing asynchronous howls, approached it more, and oriented towards it for longer compared to synchronous playbacks, a pattern consistent with threat-investigation. In how far these reactions result from a more difficult localisation of synchronous howls or more ambiguity in pack size in the asynchronous playbacks, as well as how this translates to wild wolf behaviour, remains to be investigated.

Recent evidence of extensive call sequence use in non-human primates has led to the theory that syntax evolved to mitigate the constraints of their genetically fixed repertoires, before vocal production learning later emerged in humans. However, evidence of similarly extensive sequence repertoires in an open-ended vocal production learner--the Western Australian magpie (Gymnorhina tibicen dorsalis)--offers a unique opportunity to explore potential alternative pathways to syntactic communication. Our previous work revealed fledgling magpies learn group-specific repertoires of structured call sequences from their social contacts, with more sociable individuals acquiring larger repertoires earlier in development. Notably however, the individual vocal segments that combine to form their calls and call sequences were shared across groups and emerged as early as the first week post-fledging--suggesting the underlying vocal elements may not be learned. Here we utilised acoustic neighbourhood-based dimensionality reduction to compare clustering patterns of vocal segments across magpie fledgling developmental stages, and between fledglings and adults. We found no evidence of acoustic development over time, and no significant distinction between fledgling and adult productions of the same vocal segments. The same coarticulatory effects--where a vocal element is produced differently when combined with another--and geographic variation established previously in adults were supported in fledglings too. These findings support that the vocal building blocks underpinning magpie call sequences are innate, suggesting usage learning better explains how fledglings learn to combine calls. In a species capable of open-ended production learning, this suggests learning to combine existing signals may be more adaptive than productively learning new ones. Rather than evolving solely to compensate for genetically fixed repertoires, syntax may have evolved as a flexible, convergent solution to the various challenges of expanding communicative capacity--whether due to genetic constraints, cognitive limitations or the cost of establishing new meaning in novel signals.

In over a thousand avian species, pairs coordinate vocalizations to form duets. Songbirds may use calls (simple, non-learned vocalizations) and songs (complex, learned vocalizations) to answer their partners. In Galapagos Yellow Warblers (Setophaga petechia aureola), song answers in duets appear to have evolved recently in addition to older call answers. Here, we explored the structure and functions of duets in an observational study to understand why song answers in duets may have evolved. We first described the temporal structure of duets initiated by either sex. We then tested whether the distance between pair members, countersinging with neighbours, and timing relative to aggressive encounters predicted answering probability for each sex. Finally, we tested whether the behavioral context of call and song answers differed. We found that most duets were initiated by males, and call answers were almost exclusively given by females. Female answers were also more tightly coordinated than those of males. Female answering probability decreased with increasing distance between pair members, whereas male answering probability did not. In addition, females were less likely to answer with songs when farther from their mate. Females answering probability also increased slightly when their mates initiation was preceded by a neighbors song. Our results show some evidence in support of the joint resource defence hypothesis for female answers. On the other hand, the significance of song in female answers may be related to some other functions that we did not test here, such as signaling quality or commitment to the pair bond. Lay summaryWho turns a song into a duet? In Galapagos Yellow Warblers, it is usually the female: she answers her mates song with a tightly timed call or song. Male replies were rarer and less coordinated. While male replies didnt seem to have a function, female replies were used in territorial disputes with neighbours and may indicate joint ownership of a territory. Female song replies, on the other hand, may signal commitment to the pair bond.

Zoological gardens strive to prioritize excellent animal care and adhere to the highest standards of animal welfare, ensuring that the conditions of animals` social and physical environment are as close as possible to those in the wild. This study investigates the effects of group split and relocation to the new enclosure on the behaviour of western lowland gorillas (Gorilla gorilla gorilla) (N = 7) living in Prague Zoo, Czech Republic. We conducted over 289 hours of behavioural observations focusing on daily activities, social interactions, and behaviours that serve as potential stress and welfare indicators. The group split led to establishment of two groups in two separate enclosures; the old enclosure consisted of a "bachelor group" (i.e. three males) and a new enclosure consisted of a mixed-sex group (i.e. three females and one juvenile male). The behavioural comparisons across different study periods were conducted using linear mixed models (LMMs). The changes led to an increase in time spent moving, feeding, being in social proximity, and higher rates of approaches among the gorillas, as well as to a decrease in rates of self-directed and "undesirable" behaviors. Our findings indicate that the gorillas effectively adapted to the changes, most likely by relying on social support, to navigate new conditions. This study contributes to our understanding of how socio-cognitively complex species cope with necessary alterations in captive animal care programs. Furthermore, these observations may inform strategies to enhance the welfare of zoo-housed animals and to improve their captive care.

Collective movement requires coordination between individuals, yet how this emerges during early interactions remains poorly understood. We investigated how partner familiarity influences coordination, leader-follower dynamics, and learning in homing pigeon pairs navigating from novel sites. Birds were released repeatedly with either familiar or unfamiliar partners, followed by solo releases to assess learning. By quantifying bidirectional information flow, we found familiarity influenced information-transfer dynamics during the first release: familiar pairs exhibited more asymmetric information transfer, likely reflecting established leader-follower relationships, whereas unfamiliar pairs showed more symmetric exchange. These differences disappeared after one release. Conversely, familiarity had little effect on cohesion or navigational performance. There was some evidence for an influence on learning: birds from familiar pairings had higher homing efficiency on a subsequent solo release. Finally, across partnerships, followership was more predictable than leadership with respect to individual identity and flight speed, indicating stable variation in individuals' tendency to follow rather than lead. This suggests that a shift in emphasis from leadership to followership might enhance our understanding of collective decision-making dynamics. Our results demonstrate how flight partners rapidly coordinate, producing limited downstream effects on navigation and learning, with implications for many animals that travel in fission-fusion transitory collectives.

Song evolves rapidly in songbirds, as has been proven for a substantial number of songbird species by demonstrating acoustic differences between current songs and "historical" songs recorded 20 or 30 years previously. In two species of songbirds, white-crowned sparrows and savannah sparrows, it has been further shown that evolutionary changes over such time spans are sufficient to affect the response of receivers, with territorial males responding more aggressively to current songs than to historical ones. These two species, however, have especially low population variability in song, with most males in any population singing the same, single song type; this background of song uniformity makes it especially easy to discern temporal changes. Here we examine response to temporal change in song in a third sparrow species, the song sparrow, in which population variation in song is much higher: males sing 5-13 song types each, with low song-sharing between males, so that hundreds of distinct song types occur in a local population. We find evidence that temporal change has occurred in the songs of our study population, in that 24 song current song types share more introductory phrases with other current song types than do 24 historical song types recorded 27-29 years earlier. Nevertheless, in a song playback experiment, current males showed no difference in response to current and historical songs. The results are in accord with the hypothesis that high levels of population variability in song make temporal changes in song difficult to discern.

Classic foraging models and discounting tasks may oversimplify the decision-makers environment, resulting in a discrepancy between predicted and observed behaviour. In delay discounting tasks, animals typically steeply devalue larger-later (LL) outcomes, choosing smaller-sooner (SS) rewards after short delays. This steep discounting appears to be irreconcilable with natural foraging behaviour, where animals frequently endure long delays when travelling to find food, handling tough items, and storing food for future use. This apparent mismatch in behaviour has led to questions regarding the ecological validity of laboratory discounting tasks. Here, we developed a rich dynamic optimisation model to identify the conditions under which animals should choose LL or SS outcomes. In our model, a food-storing animal encounters food items differing in energy content and handling time and must decide which items to eat immediately and which to cache, so that it has enough stored food to survive winter. We simulated a range of environments, including laboratory conditions where foragers face negligible predation risk when searching for food and have a high probability of finding food, compared to more natural conditions where searching for food is risky, and food is harder to find. In line with preference reversals seen in the discounting literature, our model predicts that LL items should be rejected more often when the handling time for such items is increased, whereas SS items should be rejected more often when the handling time for both items is increased. Importantly, our model only predicts rejection of food items under parameter values that reflect laboratory conditions, supporting the notion that the steep devaluation of rewards seen in animals may be driven by the artificiality of traditional discounting tasks.

Inequity aversion, the sensitivity to inequitable outcomes or processes, has been widely studied in nonhuman primates since 2003. Yet, findings remain debated as some could be explained by alternative mechanisms such as frustration or loss aversion, so-called individual contrasts. Previous work on nonhuman animals has restricted the definition of inequity aversion to the social domain. Here, we propose distinguishing between two different forms: a socially driven form, which depends on comparison with another individual, and an individually based form, caused by a discrepancy between ones own effort and the expected outcome. Using a task-based methodology, we manipulated both effort and the amount of reward to test for the existence of those different forms and distinguish them from individual contrasts. We presented seven Tonkean macaques (Macaca tonkeana) and three brown capuchins (Sapajus apella) with low and high effort tasks associated with low and high reward quantities. To test for the presence of individually based inequity aversion, subjects were tested alone in an individual phase. In some of the trials, they were rewarded less than they deserved for the task. To test for socially based inequity aversion, two individuals performed the same task in a social phase but, in some trials, one received a higher-value reward than the other. We recorded the latency to engage in the task as an indicator of reluctance. In the individual phase, macaques were slower to engage with inequitably rewarded tasks, a pattern consistent with an individually based expectation of equity. By contrast, capuchins were faster in this context, suggesting that their responses were more likely driven by individual contrast effects. In a context of social inequity, both species slowed engagement in tasks with inequitable rewards, suggesting an aversion to socially based inequity. These results demonstrate that such a methodology provides a means to study non-social components of inequity aversion. Future studies, conducted on larger and more diverse samples, with rigorous motivational controls and explicit tests looking at the understanding of the link between task and reward, are needed to confirm whether nonhuman primates can display inequity aversion independently of social comparison.

One challenge of reintroduction programs is ensuring that captive individuals are prepared for life in the wild, including having the ability to forage on wild resources. Foraging proficiency can be especially challenging to foster in species with a wide dietary breadth or reliance on ephemerally available resources. A relatively unexplored obstacle to release readiness may be an animals hesitancy to approach or consume unfamiliar foods, i.e. their neophobia and dietary conservatism. In this study, we examined the responses of a conservation breeding population of the extinct-in-the-wild alal[a] (Corvus hawaiiensis), a dietary generalist, to novel and familiar native and non-native fruits. Birds were similarly willing to contact both novel and familiar fruits, suggesting little aversion to the presence and appearance of novel food items. However, birds showed a clear preference for contacting non-native fruits over native fruits, regardless of familiarity. Additionally, while birds were willing to contact novel items, they were less likely to consume novel native fruits, suggesting that dietary conservatism may limit incorporation of these resources in the wild. Finally, juvenile birds were less likely to approach the testing setup and contact items, suggesting that neophobia may constrain their exploration of novel environments and resources. We discuss the implications of alal[a] preferences for non-native foods and reluctance to consume novel native items for pre-release preparations and the goal of wild species recovery.

Non-human primates are crucial animal models in biomedical research including neurosciences. Legal frameworks ensure that participation of monkeys in biomedical research follows an ethical assessment, driven by specific guidelines for ensuring animal well-being. Those guidelines promote continued refinement of practices to improve animal wellbeing, yet quantitative data to inform such refinements is limited. In this context, we investigated the impact of modifications to the housing that created either navigational complexity or visual access to neighbouring animals on the behaviour of rhesus macaques. Then we quantified the value of different enrichment programmes based on manipulable objects. We showed that making navigation within a modular housing more complex, or adding transparent separations between housing units is associated with significant reduction of severe aggression, stereotypical behaviours and a significant increase of appeasing behaviours. We also observed a positive effect of manipulable objects with an increase in the expression of appeasing behaviours and a decrease in abnormal or stereotypical behaviours. Finally, our study revealed significant inter-individual variability that was not explained by sex, age, social network size or social status. Overall, our results point to the importance and benefits of environmental changes and enrichment protocols, and of the critical need for personalised enrichment programmes.

Task allocation in eusocial insects has long been studied under the framework of division of labor, implying a relatively rigid association between individuals and tasks. However, most eusocial species lack morphological specialization, and workers regularly switch tasks as colony demands change. This raises a fundamental question: do tasks shape the behavioral profiles of workers, or does individual behavioral variation cut across task boundaries? We addressed this in a controlled laboratory study of Aphaenogaster senilis ants, comparing the behavioral profiles of four task groups (scouts, recruits, nurses, and necrophores) spatially segregated by their location within the colony setup and subsequently tested individually in four ecologically relevant contexts. This multivariate profiling, still rarely applied in ants, revealed that some tasks impose clear behavioral specialization (scouting, brood care), whereas others do not (recruitment, necrophoresis). Critically, this specialization appears in foraging-related tasks, whereas sociality does not: it varies considerably among workers, even within a single task group. Behavioral specialization, therefore, exists, but not across every dimension of behavior, and it is not a fixed property of the task. These results suggest that workers may differ in their readiness to shift roles depending on the task at hand, and this variation may in turn shape how colonies adapt to environmental change. More broadly, our results speak to a question central to collective behavior research well beyond ants: how individual variability translates into functional structure at the group level.

Risk-taking behaviour permeates daily decision-making across many taxa and has significant impact on fitness outcomes. Previous research finds that individuals display consistent tendencies in their propensity to take risks, often labelled as a "Boldness-Shyness" personality trait. In this experimental study, we investigated repeatability in risk-taking behaviour in 241 wild vervet monkeys by measuring their responses to capture threat (via human-initiated darting procedure) and predation threat (via two predator models). We further examined the influence of socio-demographic, group identity and hormonal factors on risk-taking behaviour. We found that multiple behavioural responses, including the likelihood of approaching a food source under threat, were consistent across contexts. Risk-taking behaviour was influenced by age, sex, dominance rank, and hormonal profiles: juveniles, males, and higher-ranking individuals were more likely to approach food under (perceived) predation risk. Additionally, we observed significant among-group differences, suggesting that individuals within the same social group exhibit similar risk sensitivities, potentially due to social facilitation or shared environmental effects. The findings contribute to our understanding of animal personality and the ecological and social drivers underlying variation in risk-related behaviours.

The vast majority of birds are diurnal and concentrate their vocal activity during daylight hours. However, some diurnal birds can also be vocally active at night, although the functions of this phenomenon remain poorly understood. We conducted playback experiments in the Warta Landscape Park (central Poland) to determine whether nocturnal singing by diurnal birds serves breeding-related functions by analysing responses to playback of songs from unfamiliar conspecific males. The sedge warbler (Acrocephalus schoenobaenus) was selected as the focal species because it exhibits relatively high levels of nocturnal song activity, while nine additional diurnal species detected near focal sedge warbler territories were included to explore whether responsiveness to nocturnal conspecific song extended across a broader taxonomic range. Playback experiments were conducted during the early and late stages of the breeding season and during the early and late parts of the nautical night. Out of 10 species tested, three responded vocally: sedge warbler, Savi s warbler (Locustella luscinioides), and common snipe (Gallinago gallinago). Sedge warblers did not modify song rate and song duration but increased flight activity after nocturnal playback. Savi s warblers and common snipes produced more vocalisations after playback than before in May, a pattern consistent with territorial defence function. General nocturnal vocal activity was higher at the beginning of the season, suggesting that birds motivation to establish territories and form pairs can extend into the night, providing additional benefits. Moreover, the probability of singing by the sedge warbler was higher in the latter part of the night. Our study demonstrates that nocturnal stimulation of foreign male playback of diurnal birds can elicit vocal responses from conspecifics, suggesting that nocturnal singing can occur in the absence of obvious artificial light pollution, but in the case of some species and environmental conditions, it may contribute to nocturnal social communication, especially in the early breeding season.