Social networks of reintroduced Przewalski's horses in the Great Gobi B Strictly Protected Area (Mongolia).

Analyzing social networks is a powerful tool for understanding the ecology of social species. While most studies focus on the role of each group member, few compare groups with different characteristics. The current population of Przewalski's horses Equus ferus przewalskii at the Great Gobi B Strictly Protected Area (Mongolia) includes groups of wild-born and captive-bred individuals with different experiences with the area (acclimatizing, long-term reintroduced, and wild-born), therefore serving as an ideal natural behavioral lab. We filmed 11 groups for 141.5 hours in summer 2018 (July), late spring 2019 (May, June), and autumn 2019 (September, October). Affiliative and agonistic interactions were recorded, and social networks were created. We tested the influence of origin, experience, season, sex, age, relative time belonging to the group, relatedness, and dominance rank on different network indices at the individual and group levels. We found that groups with greater experience in the area are generally better connected than members of the newly formed groups. However, these strong networks were created by wild-born individuals with very low interaction rates. On the contrary, inexperienced groups composed of captive-bred individuals displayed many interactions but created weak social networks. The results show a trend of behavioral transition from acclimatizing through long-term reintroduced to wild-born groups, supporting that the newly formed groups of released Przewalski's horses need time to display the typical social behavior patterns of wild-born individuals. Long-term monitoring of released Przewalski's horses is recommended to promote the success of this reintroduction program.

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.

Influence of weather on the behaviour of reintroduced Przewalski's horses in the Great Gobi B Strictly Protected Area (Mongolia): implications for conservation.

BACKGROUND: Reintroduction is a common technique for re-establishing threatened species. However, the adaptation to novel habitats with distinct conditions poses a risk of failure. Weather conditions affect the behaviour of animals, and thus, their adaptation to new conditions and survival. Reintroduced Przewalski's horses living in Mongolia's continental arid climate with extreme temperature and precipitation variability, serve as an ideal model species for studying the behavioural response of selected groups to these harsh conditions. METHODS: The research was conducted in The Great Gobi B Strictly Protected Area, Mongolia. In summer 2018, three groups were recorded (Azaa, Tsetsen and Mares18) involving 29 individuals. In Spring 2019, 4 groups were recorded (Azaa, Tsetsen, Hustai1 and Mares19) involving 34 individuals. In Autumn 2019, 4 groups were recorded (Azaa, Tsetsen, Hustai2 and Tanan) involving 35 individuals. Thirteen weather variables were recorded in 10-min intervals, together with the percentage representation of selected behavioural categories (feeding, locomotion, resting, and social). The effect of weather on behaviour was analysed through GLMM. Influence of the group-history factors (recently reintroduced, long-term reintroduced and wild-born) was also analysed. RESULTS: Feeding significantly increased with cloudy and windy conditions and was more frequent in autumn than spring and summer. Locomotion was positively explained by temperature and cloudiness and was higher in summer than spring and autumn. Resting behaviour decreased with altitude and cloudiness, and the dispersion of the group was lower when resting. Increased social interactions were observed with higher temperatures and were more frequent in summer compared to spring and autumn. Differences were found in the display of the behaviours among the selected harems, showing interesting patterns when grouping them according to their origin and experience. CONCLUSIONS: Weather patterns seem to influence the behaviour of Przewalski's horse. These results might assist in further management plans for the species, especially in the view of intensifying climate change and alteration of weather patterns. As previously suggested, after approximately 1 year, horses adapt to novel conditions and display the typical behavioural pattern of wild-born Przewalski's horses.

Physiological and Transcriptomic Response of Grey Poplar (Populus ×canescens Aiton Sm.) to Cadmium Stress.

(1) Background: Populus ×canescens (Aiton) Sm. is a fast-growing woody plant belonging to the family Salicaceae. Two poplar genotypes characterized by unique phenotypic traits (TP11 and TP20) were chosen to be characterized and tested for a physiological and transcriptomic response to Cd stress. (2) Methods: A comparative analysis of the effects of exposure to high cadmium (Cd) concentrations (10 µM and 100 µM) of TP11 and TP20 was performed. (3) Results: Neither of the tested Cd concentration negatively affected plant growth; however, the chlorophyll content significantly decreased. The potassium (K) content was higher in the shoots than in the roots. The magnesium concentrations were only slightly affected by Cd treatment. The zinc content in the shoots of TP20 was lower than that in the shoots of TP11. Cd accumulation was higher in the roots than in the shoots. After 10 days of exposure, 10 µM Cd resulted in comparable amounts of Cd in the roots and shoots of TP20. The most significant change in transcript amount was observed in endochitinase 2, 12-oxophytodienoate reductase 1 and phi classglutathione S-transferase. (4) Conclusions: Our study provided new insights for effective assessing the ability of different poplar genotypes to tolerate Cd stress and underlying Cd tolerance.

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.

Daddy, daddy cool: stallion-foal relationships in a socially-natural herd of Exmoor ponies.

Different forms of direct paternal investment have been described in mammals. One such species where paternal care was noticed, but remains poorly understood, is the horse (Equus caballus), where the male keeps a long-term relationship with several females and offspring. Therefore, the aim of this study was to analyse the interactions between the harem stallion and his foals. Two herds of Exmoor ponies living under semi-feral conditions in two separate reserves within Czech Republic were studied, each during one of the two consecutive breeding seasons (in 2016 and 2017). Both herds consisted of a stallion, 14 mares and their offspring (12 and 10 foals). The behaviour of all group members was recorded focusing on the stallion-foal interactions. The results show that the stallion receives more friendly interactions, snapping and playful behaviour from foals compared with adult mares. Furthermore, the stallion is more tolerant than mares, and actively plays with his offspring. There is no statistical difference in the sex of the foals in the stallion-foal interactions; however, male foals are more active than female foals in interacting with their father. The probability of a certain behaviour occurring between adult and foals changed over time. The stallion's presence might, therefore, be crucial for the physical and psychological development of the foals, especially the colts. These results may bring new insight into the common management of domestic horses, where stallions are usually kept separately and are not allowed to form natural groups.

Different maternal investment strategies for male and female calves in a polygynous mammal.

Maternal effects occur when the phenotype of the mother influences that of the young to the detriment of her survival, growth or fitness. The investment of the mother can be affected by maternal body condition and/or experience. Trivers-Willard Hypothesis (TWH) and Local Resource Competition Hypothesis (LRCH) are the main hypotheses used to explain bias in birth sex-ratios in mammals, as well as for sex-biased maternal investment. Both hypotheses suggest that a different amount of investment must be expected according to the sex of the young. However, recent studies suggest that these differences are not in quantity but in the strategies: mechanisms and objectives may differ for each sex. We studied how maternal characteristics (age, body mass, body condition, and dominance status) influence relevant aspects of the birth and early growth of the calf (birth date, birth body mass, body mass at weaning, and body condition at weaning) separately for each sex; and how that investment is mediated by milk production and composition (lactose, fat, and protein). One hundred eighty-eight newborns from 75 captive red deer hinds aged from 2 to 19 years were analyzed. The main differential investment observed was related to birth date: when producing a female, hinds give birth earlier in the season only if they have a good body condition; however, when gestating a male it is the older hinds those which deliver earlier. Subsequently, milk production and composition are correlated with birth body mass in female calves, but to weaning body mass in males. Thus, only hind body mass affects the weaning body mass of female calves, compared with age and hind body mass in males. These results suggest that while TWH fits the maternal investment strategy found for male calves, it is LRCH which correlates with the maternal investment patterns observed for females.

The Many Axes of Deer Lactation.

In undomesticated animals information about the production and composition of milk over time is still scarce. In general, for most mammals it is known that milk composition changes across lactation, is different for male and female offspring, and even that marsupials, such as kangaroos, can simultaneously produce milk of different compositions for young of different ages. Such parallel milk production of differing compositions has not yet been studied in single-offspring placental mammals, but may help to explain behavioural processes like allosuckling (feeding the young of other adults) and lateralized suckling preferences. In this study we analysed the production and composition of milk in red deer throughout the lactation period and now confirm for the first time that there are axial differences present. The front teats, which are the favoured suckling positions of the deer's offspring, produce milk with a greater protein-to-fat ratio. Also, from the beginning of lactation the yield is greater on the left side, the side preferred by calves in all of the studied species, both at population and individual level. The links between milk production and calf behaviour in deer deserve further study.

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.

Lateralized suckling in domestic horses (Equus caballus).

Brain lateralization enables preferential processing of certain stimuli and more effective utilization of these stimuli in either the left or the right cerebral hemisphere. Horses show both motor and sensory lateralization patterns. Our aim was to determine whether a lateralized response could be detected in foals during the naturally side-biased behaviour, suckling. The foals' preferred suckling side could be the effect of either visual or motor lateralization. In the case of a visual lateralized response, foals are expected to suck more often from the mother's right side, so potential danger can be detected by the better adapted right hemisphere (i.e. left eye). Motor lateralization can be identified when a foal will suck predominantly from one side, either left or right. We found no population trend in the preferred suckling side, but we detected significant differences amongst individual foals. One-third (35.4 %) of 79 foals showed a strong, either right or left side preference which increased with age. The mothers did not influence the foals' suckling side preferences either by side-biased rejection or termination of suckling. According to our findings, a general pattern of sucking with the left eye open for better danger detection and recognition is unlikely in foals up to 7 months old. Foals of this age are probably young or fully focused on suckling and rely on their mothers' vigilance. Individual side preferences amongst foals are suggested to be based on motor lateralization.

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.