Ivermectin decreases parasite load, testosterone, and potentially antler length in a group of captive red deer males (Cervus elaphus).

Secondary sex traits (SSTs) can favour males in intra-sexual competition, allowing females to reliably assess their quality. They can also be connected to other aspects of fitness, such as resistance to parasites and pathogens, as parasites have negative effects on the development of SSTs. Antlers are one of the most recognizable examples of SSTs whose development is regulated by testosterone and reflects the actual condition of the bearer. Elevated testosterone can exaggerate the size of SSTs while impairing the function of the immune system ("The Immunocompetence Handicap Hypothesis") posing a trade-off between antler development and immune function. In this study, we experimentally manipulated the parasite load in captive red deer (Cervus elaphus) males with Ivermectin during antler development for two consecutive years. Expecting an inverse proportionality between parasite load and antler size, we hypothesized the treated deer to have larger antlers than the untreated ones. Our results showed that, following the immunocompetence handicap hypothesis, parasite load was positively associated with testosterone levels. However, the application of Ivermectin suppressed the parasite load of the treated animals but did not lead to the development of larger antlers. Instead, it significantly suppressed the concentration of testosterone in the treated animals, whilst the animals that had higher testosterone also had the highest parasite load. Our findings show that Ivermectin can potentially decrease the levels of testosterone and, consequently, antler size. These findings have important implications for the management of captive populations, especially in contexts where the development of large trophies is desired.

The effects of challenge or social buffering on cortisol, testosterone, and antler growth in captive red deer (Cervus elaphus) males.

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.

Not just for males: Flehmen as a tool for detection of reproductive status and individual recognition across sexes in four African equid species.

Flehmen is frequently explained as part of male sexual behaviour, but it can also be associated with overmarking behaviour and thus individual recognition. We tested three explanatory hypotheses of flehmen behaviour: to detect sexual status of a female, to decide whether to overmark an individual, and to improve individual recognition. Additionally, we examined interspecific flehmen differences in the African equids. We observed 130 individuals of all 4 species among 15 groups in 5 zoos. We recorded 4445 eliminations: 142 were accompanied by flehmen and 1648 were inspected by another animal and followed by flehmen (n = 147 cases). As males of all age categories flehmened more often than females we conclude that flehmen serves to detect reproductive status of a female. However, this is not an exclusive explanation as animals of all sex and age categories flehmened when inspecting an elimination. Flehmen was not the predictor of overmarking. Nevertheless, we suggest that foals could use it for individual recognition. We found large interspecific differences with the highest rate of flehmen in African wild ass and least in mountain zebra. Thus, while the main function of flehmen is to detect female reproductive status, inter-individual and inter-species differences also play a role.

Why wait to mark? Possible reasons behind latency from olfactory exploration to overmarking in four African equid species.

Whereas most studies on overmarking in mammals analysed the rate of overmarking, that those investigate time between exploration of an olfactory stimulus and the response to it remain less common, with inconsistent results. We examined the latency in time between elimination by the sender and sniffing by the receiver, and from sniffing and overmarking, in four captive African equid species to explore differences among species, and among age and sex classes. We investigated these latency time periods in light of three potential hypotheses explaining overmarking behaviour in equids: social bonds, group cohesion, and intrasexual competition. Analysing 1684 events of sniffing and 719 of overmarking among 130 individuals, we found that (i) the time from elimination to overmarking was shorter among female friends and in parent-offspring dyads, proving support to the social bond hypothesis; (ii) intraspecific differences in time periods do not reflect the social organisation of species, thus not supporting the group cohesion hypothesis; (iii) males were more attracted to elimination of conspecifics than females, and female's eliminations were inspected longer, in line with the sexual competition hypothesis and/or reproductive behaviour. In addition, we found that the younger foals came to sniff eliminations faster than older ones, and in larger groups foals devoted longer time to sniffing the elimination before overmarking. We concluded that examination of the elimination could be driven by motivations other than the decision to overmark. Whereas overmarking serves to express bonds to a familiar individual, the latency of overmarking reflects more reproductive interests.

Further behavioural parameters support reciprocity and milk theft as explanations for giraffe allonursing.

Suckling of a non-filial calf, or allosuckling, is an extreme case of allomaternal care in mammals. There have been many hypotheses postulated in an attempt to explain this behaviour; however, the supporting evidence differs, together with the conclusions drawn from the investigated variables. Previously, suckling bout rejection was analysed, and the milk theft and reciprocity hypotheses were both determined as the most appropriate explanation of allosuckling in giraffe. In this study, seven hypotheses were tested using different behavioural parameters, namely suckling bout frequency, suckling bout duration, and time spent suckling. It is well-documented that these parameters are associated with various aspects in ungulate biology; for example, suckling rejection typically reflects milk intake and parent-offspring conflict, whereas the suckling bout duration and frequency is associated with social behaviours (affiliation, bonding, social stress). In total, 22 nursing females and 47 suckling calves were observed, in four Czech zoos during a five-year period. The correlation of the observed parameters between the reciprocal female-calf dyads was found to be in line with the reciprocity hypothesis. In addition, non-filial calves tried to steal the milk from non-maternal females, supporting the milk theft hypothesis. Thus, the results support both the reciprocity and milk-theft hypotheses as the most plausible explanation of allosuckling in giraffe, and illustrates the importance of using suckling bout duration and frequency, and the time spent suckling, as behavioural parameters that may aid in explaining the extremity of maternal investment, such as allosuckling.

Does social facilitation affect suckling behaviour in zebras?

Many recent studies show that synchronisation of various behaviours are advantageous for group-living mammals. However, studies on synchronisation of suckling behaviour inmonotocous species are nearly absent. We observed 49 individual foals of three zebra species in Dvur Králové Zoo, and recorded a total of 5 890 suckling bouts, almost one fifth (1 036) of which were synchronised (defined as at least two bouts overlapping at the same time). We found that synchronised suckling bouts lasted longer and were less likely to be terminated by the mother than non-synchronised ones. This is in line with social facilitation theory. In addition the occurrence of synchronised suckling bout differed interspecifically. In plains (Equus quagga) and mountain zebra (E. zebra) the probability of synchronised bouts increased with increasing numbers of foals, whereas the opposite result was found in Grevy's zebra (E. grevyi). We suggest that the interspecific differences may reflect differences in social organisation of respective species as suckling synchronisation rate was higher in species that form nurseries in the wild.

To beat or not to beat: Behavioral plasticity during the antler growth period affects cortisol but not testosterone concentrations in red deer (Cervus elaphus) males.

Out of rut, male red deer (Cervus elaphus) associate themselves in bachelor groups where animals compete for rank position via agonistic interactions. In a previous study on red deer, males were recognized either as "Non-Fighters" (NF, low frequency of attacks) or "Fighters" (F, high frequency of attacks). This study, therefore, aims to verify the consistency of the inter-individual differences in fighting attitude across different social contexts and investigate whether they could be considered an individual characteristic. Behavioral consistency was presumed across three different sampling seasons, assuming that NF would have lower cortisol (C) and testosterone (T) concentrations than the F males. In 2015 the males were kept in one large group and labelled NF and F. In 2016, the herd was divided into two subgroups ("NF" and "F") based on the frequency of attacks. Finally, in 2017, the males were divided into two randomly composed subgroups. Data about agonistic behavior and concentration of C and T were collected during each season. In 2015 the individuals differed only for the fighting attitude. After the division, the frequency of the attacks always increased, being consistently lower in NF than in F. Unexpectedly, a slight increase in the concentration of C was detected in the NF in 2016, compared to the F who experienced no difference neither in 2015 nor 2017. No significant differences were found in T. We concluded that, even though the males had shown behavioral plasticity, their diversified interaction-prone attitude had been maintained despite the modifications of the social environment.

Effect of social organisation on interspecific differences in overmarking behaviour of foals in African equids.

Overmarking remains an unstudied topic in juvenile mammals. We have previously documented a very high rate of overmarking by foals in four captive African equid species: mountain zebra (Equus zebra), plains zebra (Equus quagga), Grévy's zebra (Equus grevyi), and African wild ass (Equus africanus). African equids vary interspecifically in their social organisation. Since differences in social organisation affect many mammalian behaviours, in this study we investigated interspecific differences in overmarking behaviour of foals, analysing only cases where elimination of any other individual was explored by a foal. We hypothesised that the pattern of overmarking by foals should reflect either differences in social organisation of the species or phylogenetic relations among them. We found that in all species very young foals explored mostly maternal eliminations, and this preference declined with increasing age of the foal and reflected the social organisation of the species; the highest overmarking rate was in species with high intragroup aggression (mountain zebra) and lowest in species with low intragroup aggression and which form crèches (African wild ass). Similarly, the rate of overmarking of the mother, as opposed to other herdmates, was associated with social organisation of the respective species. Thus, we found interspecific differences in overmarking by foals, which were associated with variability in social organisation. Since we also revealed differences between African wild ass and zebra behaviour in early stages of ontogeny, we cannot refute the effect of phylogeny on overmarking behaviour. Additionally, our results supported the identity sharing hypothesis as an explanation of overmarking.

Test of four hypotheses to explain the function of overmarking in foals of four equid species.

Overmarking occurs when one individual places its scent mark directly on top of the scent mark of another individual. Although it is almost ubiquitous among terrestrial mammals, we know little about the function of overmarking. In addition, almost all studies on mammalian overmarking behaviour dealt with adult individuals. Reports on this behaviour in juveniles are extremely rare, yet may elucidate the function of this behaviour. We tested four mutually non-exclusive hypotheses which might explain this behaviour in juveniles: (1) conceal the individual's scent identity, (2) announcement of association with other group members, especially the mother-i.e., sharing identity with the mother, (3) to prevent the next conception of the mother, i.e., parent-offspring conflict, and (4) an early expression of male sexual behaviour. We observed 43 foals (out of 108 individuals) from all African equid species (Equus africanus, E. grevyi, E. quagga, E. zebra) in five zoos. In total, we recorded 3340 eliminations; 260 of these events were overmarked by 38 individual foals representing all species. This represents one of the highest rates of overmarking ever recorded by mammalian juveniles. Foals of all species except African wild ass overmarked the mother more often than another herdmate: with male foals overmarked at a higher rate than female foals. Mothers preferred to overmark foals, but not exclusively their own foal. Our results provide support for the hypotheses that overmarking serves to share identity between foal and mother, and that it is an early expression of male sexual behaviour.

Interbirth intervals are associated with age of the mother, but not with infant mortality in Indian rhinoceroses.

Rhinoceroses are among the most endangered mammals in the world. Despite a recent increase in numbers in most wild populations, poaching or political instability may exterminate large populations very quickly. Therefore, captive or ex situ rhinoceros populations can play an important role in their conservation. Previous studies identified infant mortality and interbirth intervals among the main parameters affecting the viability and survival of rhinoceros populations. In our study, we tested the recently suggested prediction that in captive Indian rhinoceros Rhinoceros unicornis, longer interbirth intervals may result in higher infant mortality. We also examined the factors that are the main predictors of infant mortality and interbith intervals using the studbook data on Indian rhinoceros born in zoos worldwide as well as data from Dudhwa National Park, India, where rhinoceroses were successfully reintroduced. We found no association between interbirth intervals and infant mortality. In both populations, the main predictor of infant mortality was mother's parity, with higher mortality in calves born to primiparous mothers. In addition, we found that the interbirth intervals were shorter in zoos than in Dudhwa and that they increased with increase in age of the mother, which was the only factor affecting interbirth interval in both populations. Our results show that the same factors affect both parameters in both populations and thus illustrate that the reproduction and infant survival of Indian rhinoceros in zoos reflect the natural pattern. Furthermore, we suggest that in captivity, the interbirth intervals could be slightly prolonged to approach the situation in the wild.

First records of preorbital gland opening in rare wild barasingha (Rucervus duvaucelii) in social contexts may help to explain this phenomenon in cervids.

The opening of the preorbital gland in deer serves as a visual communication and has been linked to a wide variety of behavioural situations. As we reported recently, all previous long-term studies on preorbital gland opening were carried out on only one species, whereas case reports on six other rarely studied species have shown associations with different behaviours, thus greatly increasing our overall understanding of the real function of this gland in animal visual communication. Here, we report for the first time preorbital gland opening in the barasingha (Rucervus duvaucelii) in social contexts as observed in a wild population in Kanha National Park, India. We observed this behaviour in two different contexts: agonistic and sexual. Moreover, our record of preorbital gland opening during copulation is the first one amongst cervids. Our findings of preorbital gland opening in both contexts in wild barasingha indicate that the gland was opened only when the individual was highly excited. We suggest that preorbital gland opening may be an important behavioural indicator of an individual involved in a serious intraspecific interaction, and thus a useful tool with which to distinguish between playful and serious behaviours, especially in agonistic and sexual situations.

Pre-orbital gland opening during aggressive interactions in rusa deer (Rusa timorensis).

The opening of the preorbital gland in cervids has a visual meaning and is frequently associated with agonistic and/or stress related situations. Apart from in red deer, this behaviour has scarcely been studied and the range of situations when it may occur remains unclear. In this study we report the unusual case of preorbital gland opening in rusa deer, Rusa timorensis, associated to direct aggressive agonistic interaction (biting/kicking) between two adult hinds. This case observed in Tierpark Berlin (Germany) is the first one ever recorded in female-female interactions in cervids. Preorbital gland opening was also studied in 116 social interactions in Plzen Zoo (Czech Republic). Preorbital gland opening by the dominant adult male was twice observed with relation to alert behaviour, which is also rare. In order to contextualise our observations we summarise the current knowledge about the behaviour associated with preorbital gland opening in R. timorensis and in cervids in general.

Laterality of suckling behaviour in three zebra species.

Although side preference while suckling is an easily characterised lateralised behaviour, few studies have been conducted. We observed laterality in suckling behaviour in three captive zebra species to test two hypotheses: laterality affected by the foal (motor laterality) and laterality affected by the mother. In total we observed 35 foals of Grevy's, plains, and mountain zebra in two zoos and recorded 5128 successful suckling bouts and 9095 unsuccessful suckling attempts. At the population level the only factor affecting side preference of suckling bouts and attempts was the identity of the individual foal. Ten foals showed individual preferences: seven foals preferred suckling from the left side of the mother, three preferred suckling from the right side of the mother. The individual preferences increased with increasing age of the foal. Only one foal was refused more often from the opposite side than the preferred side used for suckling whereas three other foals were refused from the preferred side. Foals that preferred suckling either from left or right side were refused by the mare more often than foals which showed non-preference. Thus lateral preferences in suckling behaviour of zebra foals seem to be in line with the motor laterality hypotheses.

Concurrent lactation and pregnancy: pregnant domestic horse mares do not increase mother-offspring conflict during intensive lactation.

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.

A case of adoption and allonursing in captive plains zebra (Equus burchellii).

Although allonursing (allowing non-filial offspring to suckle) can be a costly behaviour, it has been reported for many mammals including ungulate species. However, such behaviour is very rare in equids. This is the first report on adoption and allonursing in captive plains zebra (Equus burchellii), recorded in the Dvur Králové Zoo, Czech Republic. We observed a case of adoption of an orphaned foal by a lactating mare, who then regularly nursed two foals (filial and non-filial). The allonursing mare rejected more suckling attempts, terminated suckling bouts more often, and had a shorter suckling bout duration than other mares. When nursing both foals at the same time, the suckling bout lasted for less time than when nursing a single foal, regardless of whether it was filial or non-filial. The allonursing mare apparently did not discriminate between the filial and non-filial foal.

Personality in free-ranging Hanuman langur (Semnopithecus entellus) males: subjective ratings and recorded behavior.

The authors obtained behavioral observations and personality ratings for 27 free-ranging Hanuman langur males. Subjects were rated using a questionnaire based on the human Five-Factor Model (FFM). Behavioral observations were taken over 5 months using an ethogram that included 50 behaviors. Principal Component Analysis (PCA) of ratings revealed Agreeableness_(R), Confidence_(R), and Extraversion_(R) components. Each personality dimension was associated with a unique set of observed behaviors. PCA of 36 behavioral indices revealed Dominance_(B), Involvement_(B), and Activity_(B) components. Bivariate correlations showed that Agreeableness_(R) was negatively correlated with Dominance_(B); Confidence_(R) was positively correlated with Dominance_(B) and Involvement_(B) but negatively correlated with Activity_(B); and Extraversion_(R) was positively correlated with Activity_(B). Dominance rank was positively correlated with Confidence_(R) and Dominance_(B) but negatively correlated with Agreeableness_(R) and Activity_(B). These results highlight the comparability of behavioral coding and personality ratings and suggest that some aspects of personality structure were present in the common ancestor of Old World monkeys.