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Abstract We equipped 17 captive red deer males (Cervus elaphus) with GPS collars to measure inter-individual distances throughout the 5-months of the antler growth period. We expected some individuals to associate regularly with others while others would not. We predicted that males aggregating with others within a socially stable environment (Associates) would benefit from a form of “social buffering” and would likely have lowered cortisol (C) and testosterone (T) concentrations. Males only irregularly joining social groupings would experience elevated levels of aggression; according to the “Challenge hypothesis”, their T and C concentrations should increase. Interacting with a higher proportion of Associates did indeed reduce C concentrations. Conversely, avoiding Associates and challenging other males stimulated the T secretion. Admittedly, males avoiding regular proximity to others tended to develop the largest antlers. They probably benefited from frequent successful agonistic threats to conspecifics, resulting in elevated T concentrations. Regular association with tolerant, conspecifics and “social buffering” did not seem sufficient for producing larger antlers despite reducing C concentrations. Alternative social strategies were adopted within the same group of individuals and showed how the trade-off between these strategies could have an essential impact on C and T concentrations.

Abstract It is well known that gibbons emit a pattern of vocalizations, which is specific for species and sex. A previous study showed, however, that immature southern yellow-cheeked gibbon (Nomascus gabriellae) males produce only female-like great calls from 2.3 to 5.3 years of age in co-singing interactions with their mothers. To date, nothing is known about how the vocal repertoire of a male changes from the female-like call (great call) to the male call (staccato notes and multi-modulation phrase) during vocal ontogeny. The goal of this study was to describe the transition from the female-like great call to the male call and the ontogeny of the male call. We predicted that the transition from the female-like great call to the male-specific call and the development of the male call is a normal part of the aging proces. If this is the case, the following phenomena will occur: (a) female vocalization should no longer be produced with the mature form of the multi-modulation phrase and (b) all stages of the male vocalization should occur gradually as the young male ages. Young males regularly emit both female-like great calls and male-specific calls between the ages of 5.6 to 7.1 years. Once the young males reached 7.1 years of age, they emitted male calls exclusively, and they continued to do so until the end of the observation period (at 8.11 years of age). It was confirmed that the young males emitted only female-like great calls during periods when they produced non-mature forms of a multi-modulation phrase (Fm0,1—none or one frequency modulation in second notes). Furhermore, the decrease in the number of female-like great calls was attributed to the development of the mature form of the multi-modulation phrase (Fm2—two or more frequency modulation in second notes), which developed with age. We also confirmed that the multi-modulation phrase developed gradually, while the development of the staccato notes occurred in leaps. A multi-modulation phrase developed as the initial part of the male-specific call. It was evolved from a simpler to a more complex form as the maximum frequency and age of the young males increased. Staccato notes subsequently developed in certain young males. Possible explanations for such vocal ontogeny in young males are discussed in this work.

Abstract Physical condition is important for the ability to resist various parasites and diseases as well as in escaping predators thus contributing to reproductive success, over-winter survival and possible declines in wildlife populations. However, in-depth research on trends in body condition is rare because decades-long datasets are not available for a majority of species. We analysed the long-term dataset of offspring covering 34 years, male parents (40 years) and female parents (42 years) to find out whether the decline of Tengmalm’s owl population in western Finland is attributable to either decreased adult and/or juvenile body condition in interaction with changing weather conditions and density estimates of main foods. We found that body condition of parent owl males and females declined throughout the 40-year study period whereas the body condition of owlets at the fledging stage very slightly increased. The body condition of parent owls increased with augmenting depth of snow cover in late winter (January to March), and that of offspring improved with increasing precipitation in late spring (May to June). We conclude that the decreasing trend of body condition of parent owl males and females is important factor probably inducing reduced adult survival and reduced reproduction success thus contributing to the long-term decline of the Tengmalm’s owl study population. The very slightly increasing trend of body condition of offspring is obviously not able to compensate the overall decline of Tengmalm’s owl population, because the number of offspring in turn simultaneously decreased considerably in the long-term. The ongoing climate change appeared to work in opposite ways in this case because declining depth of snow cover will make the situation worse but increased precipitation will improve. We suggest that the main reasons for long-term decline of body condition of parent owls are interactive or additive effects of reduced food resources and increased overall predation risk due to habitat degradation (loss and fragmentation of mature and old-growth forests due to clear-felling) subsequently leading to decline of Tengmalm’s owl study population.

Abstract Recent wildlife population declines are usually attributed to multiple sources such as global climate change and habitat loss and degradation inducing decreased food supply. However, interactive effects of fluctuations in abundance of main foods and weather conditions on population densities and reproductive success have been studied rarely. We analysed long-term (1973–2018) data on Tengmalm’s owl (Aegolius funereus) and the influence of prey abundance and weather on breeding densities and reproductive success in western Finland. We found that fledgling production per breeding attempt declined and laying date of the owl population delayed during the period between 1973 and 2018. The breeding density of the owl population decreased with increasing temperature in winter (October–March), fledgling production increased with increasing temperature and precipitation in spring (April–June), whereas the initiation of egg-laying was delayed with increasing depth of snow cover in late winter (January–March). The decreasing trend of fledgling production, which was mainly due to starvation of offspring, was an important factor contributing to the long-term decline of the Tengmalm’s owl study population. Milder and more humid spring and early summer temperatures due to global warming were not able to compensate for lowered offspring production of owls. The main reason for low productivity is probably loss and degradation of mature and old-growth forests due to clear-felling which results in loss of coverage of prime habitat for main (bank voles) and alternative foods (small birds) of owls inducing lack of food, and refuges against predators of Tengmalm’s owls. This interpretation was also supported by the delayed start of egg-laying during the study period although ambient temperatures increased prior to and during the egg-laying period.

Abstract Social deprivation early in life affects further individual development and leads to irreversible behavioural alterations later in life. Although the syndrome is well-studied in vertebrates including humans, its presence in invertebrates has been described only in eusocial insects and cockroaches. Here we show the first evidence of social deprivation in subsocial decapod crustaceans, based on analysis of video-recorded agonistic encounters of juvenile red swamp crayfish (Procambarus clarkii, Girard). In comparison with maternally incubated juveniles, isolated crayfish had altered repertoires, numbers and frequency of agonistic interactions similar to those described in vertebrates. Our results support the view on the syndrome of social deprivation as a ubiquitous trait in species with developed maternal care across diverse taxa.

It was shown earlier that dogs, when selecting between two dishes with snacks placed in front of them, left and right, prefer to turn either clockwise or counterclockwise or randomly in either direction. This preference (or non-preference) is individually consistent in all trials but it is biased in favor of north if they choose between dishes positioned north and east or north and west, a phenomenon denoted as "pull of the north". Here, we replicated these experiments indoors, in magnetic coils, under natural magnetic field and under magnetic field shifted 90° clockwise. We demonstrate that "pull of the north" was present also in an environment without any outdoor cues and that the magnetic (and not topographic) north exerted the effect. The detailed analysis shows that the phenomenon involves also "repulsion of the south". The clockwise turning preference in the right-preferring dogs is more pronounced in the S-W combination, while the counterclockwise turning preference in the left-preferring dogs is pronounced in the S-E combination. In this way, south-placed dishes are less frequently chosen than would be expected, while the north-placed dishes are apparently more preferred. Turning preference did not correlate with the motoric paw laterality (Kong test). Given that the choice of a dish is visually guided, we postulate that the turning preference was determined by the dominant eye, so that a dominant right eye resulted in clockwise, and a dominant left eye in counterclockwise turning. Assuming further that magnetoreception in canines is based on the radical-pair mechanism, a "conflict of interests" may be expected, if the dominant eye guides turning away from north, yet the contralateral eye "sees the north", which generally acts attractive, provoking body alignment along the north-south axis.

Despite anecdotal reports of the astonishing homing abilities in dogs, their homing strategies are not fully understood. We equipped 27 hunting dogs with GPS collars and action cams, let them freely roam in forested areas, and analyzed components of homing in over 600 trials. When returning to the owner (homewards), dogs either followed their outbound track (‘tracking’) or used a novel route (‘scouting’). The inbound track during scouting started mostly with a short (about 20 m) run along the north-south geomagnetic axis, irrespective of the actual direction homewards. Performing such a ‘compass run’ significantly increased homing efficiency. We propose that this run is instrumental for bringing the mental map into register with the magnetic compass and to establish the heading of the animal.

The period of early ontogeny constitutes a time when the physical immaturity of an organism is highly susceptible to external stimuli. Thus, early development plays a major role in shaping later adult behavior. The aim of the study was to check whether stimulating puppies at this early stage in life with sound would improve their responsiveness towards unfamiliar noises during the selection process of the police behavioral test for puppies. The cohort comprised 37 puppies from the litters of three mothers. At the commencement of the experiment the dogs were aged 16 days, rising to the age of 32 days at its close. The mothers and litters of the treatment group were either exposed to radio broadcasts, (see below; three litters totaling 19 puppies), while the control group was not exposed to any radio programs (eight litters totaling 18 puppies). All three mothers had previously experienced both auditory circumstances, as described herein. Ordinary radio broadcasts were played to the puppies in the treatment group three times a day for 20 minute periods, always during feeding time. The cohort was subjected to the so-called Puppy Test, i.e. analysis of the potential of each animal, once the dogs had reached the age of 7 weeks. Such tests included exposure to a sudden noise caused by a shovel (100 dB), noise when alone in a room, and response to loud distracting stimuli (the latter two at 70 dB). Said tasks were rated by the same analyst on a scale of 0-5 points; the better the response of the dog, the higher the score given. The differences between the treatment and control groups were analyzed via Mixed Models (PROC MIXED) in SAS. The animals comprising the treatment group responded with a higher score to the sudden noise caused by the shovel than the control dogs (P

Litter size is one of the most reliable state-dependent life-history traits that indicate parental investment in polytocous (litter-bearing) mammals. The tendency to optimize litter size typically increases with decreasing availability of resources during the period of parental investment. To determine whether this tactic is also influenced by resource limitations prior to reproduction, we examined the effect of experimental, pre-breeding food restriction on the optimization of parental investment in lactating mice. First, we investigated the optimization of litter size in 65 experimental and 72 control families (mothers and their dependent offspring). Further, we evaluated pre-weaning offspring mortality, and the relationships between maternal and offspring condition (body weight), as well as offspring mortality, in 24 experimental and 19 control families with litter reduction (the death of one or more offspring). Assuming that pre-breeding food restriction would signal unpredictable food availability, we hypothesized that the optimization of parental investment would be more effective in the experimental rather than in the control mice. In comparison to the controls, the experimental mice produced larger litters and had a more selective (size-dependent) offspring mortality and thus lower litter reduction (the proportion of offspring deaths). Selective litter reduction helped the experimental mothers to maintain their own optimum condition, thereby improving the condition and, indirectly, the survival of their remaining offspring. Hence, pre-breeding resource limitations may have facilitated the mice to optimize their inclusive fitness. On the other hand, in the control females, the absence of environmental cues indicating a risky environment led to "maternal optimism" (overemphasizing good conditions at the time of breeding), which resulted in the production of litters of super-optimal size and consequently higher reproductive costs during lactation, including higher offspring mortality. Our study therefore provides the first evidence that pre-breeding food restriction promotes the optimization of parental investment, including offspring number and developmental success.

Knowledge about vocal ontogeny and vocal plasticity during ontogeny in primate species is central to understanding the evolution of human speech. Vocalizations in gibbons (Hominoidea) are very interesting and contain complex species- and sex-specific patterns. However, ontogeny of gibbon songs is little studied. Here, we document regular production and ontogenetic changes of female-specific "great call" in 4 immature (2 juvenile-c.a. 3 years old; and 2 adolescent-c.a. 5 years old) males of southern yellow-cheeked gibbon (N. gabriellae) over nine months. None of the males produced fully developed adult-like "great call" and little ontogenetic changes to "great calls" occurred. "Great calls" of sons were shorter, started higher and ended lower than those of their mothers. Regular production of twitter part of great call likely appears around 4th year as it was observed in adolescent but not in juvenile males.

[This corrects the article DOI: 10.1371/journal.pone.0173985.].

Animal home ranges typically characterized by their size, shape and a given time interval can be affected by many different biotic and abiotic factors. However, despite the fact that many studies have addressed home ranges, our knowledge of the factors influencing the size of area occupied by different animals is, in many cases, still quite poor, especially among raptors. Using radio-telemetry (VHF; 2.1 g tail-mounted tags) we studied movements of 20 Tengmalm's owl (Aegolius funereus) males during the breeding season in a mountain area of Central Europe (the Czech Republic, the Ore Mountains: 50° 40' N, 13° 35' E) between years 2006-2010, determined their average hunting home range size and explored what factors affected the size of home range utilised. The mean breeding home range size calculated according to 95% fixed kernel density estimator was 190.7 ± 65.7 ha (± SD) with a median value of 187.1 ha. Home range size was affected by prey abundance, presence or absence of polygyny, the number of fledglings, and weather conditions. Home range size increased with decreasing prey abundance. Polygynously mated males had overall larger home range than those mated monogamously, and individuals with more fledged young possessed larger home range compared to those with fewer raised fledglings. Finally, we found that home ranges recorded during harsh weather (nights with strong wind speed and/or heavy rain) were smaller in size than those registered during better weather. Overall, the results provide novel insights into what factors may influence home range size and emphasize the prey abundance as a key factor for breeding dynamics in Tengmalm's owl.

In altricial birds, the nestling period is an important part of the breeding phase because the juveniles may spend quite a long time in the nest, with associated high energy costs for the parents. The length of the nestling period can be variable and its duration may be influenced by both biotic and abiotic factors; however, studies of this have mostly been undertaken on passerine birds. We studied individual duration of nestling period of 98 Tengmalm's owl chicks (Aegolius funereus) at 27 nests during five breeding seasons using a camera and chip system and radio-telemetry. We found the nestlings stayed in the nest box for 27 - 38 days from hatching (mean ± SD, 32.4 ± 2.2 days). The individual duration of nestling period was negatively related to wing length, but no formally significant effect was found for body weight, sex, prey availability and/or weather conditions. The fledging sequence of individual nestlings was primarily related to hatching order; no relationship with wing length and/or other factors was found in this case. We suggest the length of wing is the most important measure of body condition and individual quality in Tengmalm's owl young determining the duration of the nestling period. Other differences from passerines (e.g., the lack of effect of weather or prey availability on nestling period) are considered likely to be due to different life-history traits, in particular different food habits and nesting sites and greater risk of nest predation among passerines.

Begging behaviour of nestlings has been intensively studied for several decades as a key component of parent-offspring conflict. There are essentially two main theories to account for intensity of food solicitation among offspring: that intensity of begging is related to some form of scramble competition between nest mates or that it offers honest signalling of need to parents. The vast majority of studies which have addressed begging behaviour have been based on observations of, and experiments on, nestlings and have not considered begging behaviour, during the post-fledging period. Begging vocalizations in this post-fledging phase of dependence have rarely been studied, despite the importance of vocalizations as a communication method between offspring and parents, particularly for nocturnal species. We radiotracked 39 fledglings of the Tengmalm's owl (Aegolius funereus) in two years with different availability of prey: 2010 (n = 29 fledglings) and 2011 (n = 10 fledglings) and made 1320 nightly localizations in which we recorded presence or absence of begging calls. Within years, the most important measures related to the probability of vocalization were body condition at fledging, time of night, number of surviving siblings, age and weather conditions. Begging intensity increased with age in both years; however, in the year with low prey availability fledglings vocalized significantly more often. The main factor causing these differences between years was probably the different availability of prey, affecting breeding success, post-fledging behaviour, and thus also both short- and long-term needs of offspring. We believe that our results suggest honest signalling of their fledgling's need.

The aim of the study was to determine whether cortisol and secretory Immunoglobulin A (sIgA) could be used as an indicator of acute stress in both young and adult dogs. Seventeen German shepherd puppies were exposed to the Puppy test (challenge test) at the age of seven weeks. This test has been routinely used to assess the future working ability of potential police dogs. In addition, ten adult females were subjected to 4 minutes of defense training under stressful conditions. Saliva was collected from the puppies and adult females before testing and 20 minutes after the start of testing, using a cotton swab held for 1-2 minutes in each dog's mouth. Cortisol concentrations increased after the test compared to the control sample both in puppies and the adult females. However adult females showed a significant decrease in sIgA after defense training while puppies showed a tendency of increase in sIgA. We propose that salivary cortisol could be used as an indicator of stress in puppies during early ontogeny. It is not yet clear whether sIgA could be used as a useful indicator of short-term stress in dogs.

The behaviour of the nestlings of nocturnal cavity-nesting species has relatively rarely been studied in detail because of problems connected with use of the technical devices required to provide long-term monitoring of individuals. However, long-term observation of nestling behaviour is crucial in order to identify different types of behaviour which may be caused by sibling competition at the end of nesting period. We studied behaviour of 43 Tengmalm's owl (Aegolius funereus) nestlings at 14 nests using a camera and a chip system. The nestlings perched at the nest box entrance from an average age of 28 days from hatching (range 24-34 days) until fledging, spending around 2 hours per day here in total, in periods ranging from a few seconds to 147 min (7.6±10.9 min, mean ± SD). We found that individual duration of perching at the nest box entrance was significantly influenced by nestlings' age and wing length and that the duration of perching at the nest box entrance significantly decreased with time of night. However, during daylight hours, time of day had no effect on either probability or duration of nestlings' perching. We suggest daylight perching at the nest box entrance results from nestlings' preparation for fledging, while individuals perching here during the night may gain an advantageous position for obtaining food from the parents; another possibility at all times of day is that nestlings can reaffirm their social dominance status by monopolizing the nest box entrance.

Allosuckling is a situation when a female nurses a non-filial offspring. It was described in various ungulate species; however for camels this is the first description of this behaviour. The aim of the study was to assess the occurrence of allosuckling in captive camels (Camelus bactrianus) and to test whether it can be explained as a 'milk-theft' (opportunistic behaviour of calves) or alternatively as an altruistic behaviour of females. During 2005 and 2007, nine camel females and ten calves in four zoological gardens in the Czech Republic were observed. In total, 373 sucking bouts were recorded, from which 32 were non-filial (the calf sucked from the non-maternal female). Allosuckling regularly appeared in captive camel herds. As predicted for the milk-theft explanation, the non-filial calves sucked more often in the lateral position and even did not suck in the antiparallel position at all. The non-filial calves preferably joined the filial calf when sucking but in five cases (15.6% of non-filial sucking bouts) the calves sucked from non-maternal dam without the presence of filial calf. We then expected the differences in terminations of sucking bouts by females but did not find any difference in sucking terminations for filial and non-filial calves. As the calves were getting older, the incidence of allosucking increased. This was probably because skills of the calf to outwit the non-maternal dam increased and/or the older calves might be more motivated for allosucking due to the weaning process. Finally, duration of a sucking bout was shorter with non-filial than filial calves. The results of the study support the hypothesis of 'milk theft', being mostly performed by calves behaving as opportunistic parasites, but we cannot reject certain level of altruism from the allonursing females or their increased degree of tolerance to non-filial calves.

Fledgling behaviour and movement patterns throughout the post-fledging dependence period (PFDP), especially in relation to changing environmental conditions, have been rarely studied, despite the fact that this period is recognized as of crucial significance in terms of high mortality of juveniles. The PFDP can extend over quite a protracted period, particularly in birds of prey, and a knowledge of the movement patterns of individuals is fundamental for understanding mechanisms underlying survival, habitat use and dispersion. We radiotracked 39 fledglings of the Tengmalm's owl (Aegolius funereus) in two years with different availability of prey: 2010 (n = 29) and 2011 (n = 10) and obtained 1455 daily locations. Fledglings reached independence on average in 45 days after fledging in 2010 (n = 22) and 57 days in 2011 (n = 6). Within years, the most important measures influencing the distance moved from the nest box were age of fledglings and number of surviving siblings present. Individual home range size and duration of PFDP in particular were dependent on maximal number of siblings seen outside the nest box. In the season with low prey availability fledglings were observed at greater distances from the nest box than in the year with higher prey availability (mean distance: 350 m in 2010 and 650 m in 2011) and occupied larger home ranges (mean: 30.3 ha in 2010 and 57.7 ha in 2011). The main factor causing these differences between years was probably the different availability of prey in these two years, affecting breeding success and post-fledging survivorship of the Tengmalm's owls.

Antlers as a potential model for bone growth and development have become an object of rising interest. To elucidate processes explaining how antler growth is regulated, in vitro cultures have been established. However, until now, there has been no standard method to cultivate antler cells and in vitro results are often opposite to those reported in vivo. In addition, many factors which are often not taken into account under in vitro conditions may play an important role in the development of antler cells. In this study we investigated the effects of the antler growth stage, the male individuality, passaged versus primary cultures and the effect of foetal calf serum concentrations on proliferative potential of mixed antler cell cultures in vitro, derived from regenerating antlers of red deer males (Cervus elaphus). The proliferation potential of antler cells was measured by incorporation of (3)H thymidine. Our results demonstrate that there is no significant effect of the antler growth stage, whereas male individuality and all other examined factors significantly affected antler cell proliferation. Furthermore, our results suggest that primary cultures may better represent in vivo conditions and processes occurring in regenerating antlers. In conclusion, before all main factors affecting antler cell proliferation in vitro will be satisfactorily investigated, results of in vitro studies focused on hormonal regulation of antler growth should be taken with extreme caution.

Earlier studies have shown variation among experimental attempts to establish whether human monozygotic twins that are genetically identical also have identical individual scents. In none of the cases were the dogs able to distinguish all the individual scents of monozygotic twins living in the same environment if the scents were presented to them separately. Ten specially trained police German Shepherd dogs of three Czech Republic Police Regional Headquarters were used for scent identification in our study. The dogs were supposed to match scents of two monozygotic pairs (5 and 7 years old) and two dizygotic twin pairs (8 and 13 years old). Scents were collected on cotton squares stored in glass jars. Dog handlers were blind to the experiment details. In each trial (line-up), one scent was used as a starting scent and the dog was then sent to determine if any of the 7 presented glass jars contained a matching scent. Scents of children of similar ages were used as distractors. In the matching procedure, the dogs matched correctly the scent of one twin with the other, as well as two scents collected from every single identical and non-identical twin to prove their efficacy and likewise, the presence of the matching twin scent in any given glass jar. All dogs in all trials distinguished correctly the scents of identical as well as non-identical twins. All dogs similarly matched positively two scents collected from the same individuals. Our findings indicated that specially trained German Shepherd dogs are able to distinguish individual scents of identical twins despite the fact that they live in the same environment, eat the same food and even if the scents are not presented to them simultaneously.

Lactation is the most energy demanding part of parental care in mammals, so parent-offspring conflict arises over milk provided by the mother. In some species females commonly become pregnant shortly after parturition of previous young. This further intensifies mother-offspring conflict due to concurrent pregnancy and lactation. In equids it has been well established that pregnant females wean their foals earlier than non-pregnant ones. Intensified mother-offspring conflict was presumed to associate with pregnancy also during the period of intensive lactation, i.e., before the weaning process starts. We investigated the effect of pregnancy on suckling behaviour characteristics as indicators of mother-offspring conflict in domestic horses. Contrary to expectation, here we provide evidence of a decreased mother-offspring conflict related to pregnancy in lactating females during first two trimesters of pregnancy. Pregnant mares provided longer suckling bouts and did not reject or terminate suckling of their foals more often than non-pregnant mares. Our results suggest that pregnant mares cope with parallel investment into a nursed foal and a foetus through enhancing nursing behaviour in early stages of pregnancy before the initially low requirements of the foetus increase. They compensate their suckling foal with the perspective of its early weaning due to ongoing pregnancy.

Social dominance is widely known to facilitate access to food resources in many animal species such as deer. However, research has paid little attention to dominance in ad libitum access to food because it was thought not to result in any benefit for dominant individuals. In this study we assessed if, even under ad libitum conditions, social rank may allow dominant hinds to consume the preferred components of food. Forty-four red deer hinds (Cervus elaphus) were allowed to consume ad libitum meal consisting of pellets of sunflower, lucerne and orange, and seeds of cereals, corn, cotton, and carob tree. The meal was placed only in one feeder, which reduced accessibility to a few individuals simultaneously. During seven days, feeding behavior (order of access, time to first feeding bout, total time spent feeding, and time per feeding bout) were assessed during the first hour. The relative abundance of each meal component was assessed at times 0, 1 and 5 h, as well as its nutritional composition. Social rank was positively related to the amount of time spent feeding during the 1(st) h (P = 0.048). Selection indices were positively correlated with energy (P = 0.018 during the 1(st) h and P = 0.047 from 1(st) to 5(th)) and fat (only during the 1(st) h; P = 0.036), but also negatively with certain minerals. Thus, dominant hinds could select high energy meal components for longer time under an ad libitum but restricted food access setting. Selection indices showed a higher selectivity when food availability was higher (1(st) hour respect to 1(st) to 5(th)). Finally, high and low ranking hinds had longer time per feeding bout than mid ones (P = 0.011), suggesting complex behavioral feeding tactics of low ranking social ungulates.