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Five periods of antler growth showing which parts were developing at the same time. Adapted from Bartoš and Losos (1997). 32  

Five periods of antler growth showing which parts were developing at the same time. Adapted from Bartoš and Losos (1997). 32  

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In this review we offer a synthesis of a 30-year-long investigation focussed on the relationship between dominance rank-related behaviour and the timing and growth of antlers in deer. Our studies related primarily to red and fallow deer. We present evidence to suggest that dominance-related behaviour in male deer is strong enough to influence both...

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... this basis, the entire phase of antler growth was divided into five segments of time (periods). Periods were defined to cover the time during which a single antler characteristic developed (Fig. 4). 'Period 5' represents the time of antler elongation, which lasts ~70% of the entire velvet period; see for example Muir et al. 27 The periods were determined for each animal separately, according to the individual speed of antler growth. Upon knowing the final shape and size of the antler, the periods were extrapolated Bartoš and ...

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... In turn, high concentrations of individual cholesterol fractions in red deer plasma may cause serious health consequences [79]. Finally, the maintenance of the dominant male position in a herd of farmed hinds also generates a strong stress reaction [80], which may be reflected in the parameters studied. ...
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As a reactive species, the red deer is sensitive to both negative exogenous and endogenous stimuli. An intensive hunting period may have a particularly negative impact on game animals. The aim of this study was to determine the plasma cortisol level and biochemical parameters in 25 wild red deer (Cervus elaphus) harvested during stalking hunts in correlation with the sex and age of the animals. The mean cortisol concentrations in the stags and hinds analyzed in this study were similar (20.2 and 21.5 ng/mL, respectively). Higher HDL cholesterol values were found in the blood of the hinds than in stags (p < 0.05). Similarly, the mean levels of LDL cholesterol, lactate dehydrogenase, and alanine aminotransferase were higher by 21%, 16%, and 42%, respectively, in the blood of the hinds. In contrast, the levels of alkaline phosphatase, bilirubin, and aspartate aminotransferase were higher in the stags (by 30%, 49%, and 36%, respectively). There was a negative correlation of the cortisol concentration with urea and bilirubin and a positive correlation between cortisol and aspartate aminotransferase in the stags (p < 0.05). In turn, a negative correlation was found between the cortisol and urea levels in the hinds (p < 0.05). In summary, the stress caused by stalking hunts and the characteristic behavior of red deer during the mating season had an impact on chosen biochemical parameters. The increased concentration of cortisol resulted in a decrease in the carcass mass, which may lead to the deterioration of the physical condition of animals on hunting grounds.
... The positive effect of the increasing total of attacked conspecifics on antler growth (Fig. 3c) also suggested involvement of an effect of dominance throughout increased aggression of the males. Such a role of the "invisible", but nonetheless present dominance is in agreement with our previous studies 3, 55 . Details obtained in the current study showed that association with tolerant individuals or attacking Non-Associates was a more potent factor than dominance, as we previously thought 10,55 . ...
... Such a role of the "invisible", but nonetheless present dominance is in agreement with our previous studies 3, 55 . Details obtained in the current study showed that association with tolerant individuals or attacking Non-Associates was a more potent factor than dominance, as we previously thought 10,55 . ...
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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.
... In cervids, increased CORT levels are typical in the winter [15,24,[26][27][28][29]. However, recent reports indicate an increased concentration of this hormone in deer during the growth and mineralization of antlers, i.e., in early spring and late summer [30][31][32]. It is likely related to species differences and sensitivity to climatic factors [26]. ...
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Background The European roe deer ( Capreolus capreolus ) is a species particularly reactive to all kinds of negative stimuli. Hunting activity is one of the most potent stressors that disturbs the welfare of wild animals. During stress, various endocrine responses are elicited to improve the physical performance of the affected individual. A commonly assessed hormone for overcoming stressful situations is cortisol (CORT). In this study, plasma CORT levels in roe deer were assessed during the season of the most intense stalking hunts in Poland (summer vs. late autumn), the sex of the harvested animals (males vs. females), and age of harvest animal. In addition, the health status of the roe doe was evaluated on the basis of selected indices of blood chemistry, which could be associated with circulating cortisol levels. Results The mean cortisol levels were 58.066 ng/ml in the male group (summer) and 27.694 ng/ml in the female group (late autumn). Higher CORT levels were associated with a significantly lower of total cholesterol, lactate dehydrogenase, and uric acid ( p < 0.05). Moreover, the mean concentration of uric acid was negatively correlated with the level of CORT in the male and female groups ( p < 0.05). Together with the increase in mean CORT level, the HDL cholesterol of all the tested animals increased significantly ( p < 0.05). Conclusions Higher CORT in males during the summer than in females during the late autumn were most likely due to the arousal with the mating season. The level of CORT increased with the animals’ age. Uric acid and age are both predictors of roe deer’s serum CORT level.
... Outcomes of this study can be applied to many fields of roe deer biology since this species is an excellent model for evolutionary studies, biodiversity and social organization (Bartos and Bubenik 2011). In combination with interspecies cytogenetic studies, SNPs markers can also be used for the mapping of genes involved in the variation of unique organ development (i.e. ...
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Blood samples from forty-six roe deer ( Capreolus capreolus) acquired during officially approved hunting in six hunting divisions throughout Poland were used to isolate the genomic DNA. All individuals were genotyped by MD_Bovine BeadChip (Illumina) for 46.750 Single Nucleotide Polymorphism (SNP) markers. SNPs of inappropriate clusters, with a marker call rate lower than 90% and with a minor allele frequency (MAF) lower than 0.01, located on sex chromosomes and mitochondrial DNA, were removed. Altogether, 21.033 SNP markers were included for further analysis. Observed and expected heterozygosity amounted to 0.098 and 0.119, respectively. Among 21.033 markers, a panel of 148 SNPs were selected for relationship analysis. They were unlinked and had a MAF higher than 0.2. This set of SNPs showed a probability of parentage exclusion of 1.29x10 -6 and 2.37x10 -19 for one, and two known parents, respectively. The probability of identity was estimated at 1.8x10 -40. The probabilities obtained in this study are sufficient for the monitoring and effective management of the genetic diversity of roe deer in Poland and is a cost-effective complementary tool for forensic applications.
... This usually happens among animals that know each other (Bartoš, 1982) and where social dynamics do not suffer from unexpected events (e.g., a dominant deer suffers severe injuries or arrival of an unknown animal) (Clutton-Brock et al., 1979). Even though some animals were consistently more dominant than other animals, the hierarchy of our bachelor group was not linear, which is rare in this species (Appleby, 1983;Bartoš and Bubenik, 2011;but see Clutton--Brock et al., 1982;Lincoln, 1972;and Mitchell et al., 1977 for linear hierarchies). Interestingly, not all dominant individuals perceived agonistic interactions in the same way (Esattore et al., 2020), and these inter-individual differences in interaction-proneness might be related to personality. ...
Article
Personality is not a uniquely human characteristic and it has been documented in a wide range of organisms, from mammals to birds, reptiles, fish and invertebrates. However, personality is still poorly understood in Cervids. Therefore, our study aimed to fill this gap by i) investigating personality and ii) exploring its links to dominance hierarchy, assessed by behavioral observations in 11 captive and tame male red deer (Cervus elaphus). Using questionnaires to assess personality, three trained volunteers rated these animals in 15 behaviorally composed adjectives with detailed descriptions, based on their overall impression at the end of the observation period. Behavioral data from animals was collected across three different situations, namely “feeding” (i.e., high competition for a scarce resource), “normal” (i.e., no external stimuli) in a group setting, and “handling” (i.e., stressful situation due to human manipulation) in an individual setting. We estimated dominance hierarchies between the individuals based on situations of average and high competition (i.e., “normal” and “feeding”) via the Clutton-Brock Index (CBI). Using Fleiss’ Kappa for inter-rater reliability, only five of our 15 behavioral adjectives showed acceptable reliability. Using principal component analysis, four of these adjectives formed one personality component labelled “Confidence/Aggressiveness”. We found that although “Confidence/Aggressiveness” did not correlate with CBI, ratings of two adjectives loading onto this component, namely “Confident” and “Submissive”, significantly correlated with the CBI, indicating that questionnaire ratings reflect real behavioral variation in red deer males. Our study provides the first assessment of personality in male red deer and adds to the growing literature on Cervid personality, offering the basis for future personality research in ungulates.
... Prior studies of sika deer in Japan (namely C. n. nippon and C. n. yesoensis) concluded that food habits were different among sex and age groups. For example, large adult males tended to feed more frequently but consume lower quality forage, whereas the smaller adult females and fawns consumed less, higher quality forage (Padmalal and Takatsuki, 1994;Asada and Ochiai, 1996). Another study found that the composition of the herd and size of the group is flexible, and varied seasonally and among different habitats (Jayasekara and Takatsuki, 2000). ...
... Antler size and the number of male antler points are usually influenced by multiple factors, including age, nutrition and physiological status; they generally correspond to social rank in cervids (Hayden et al., 1994;Barto s and Bubenik, 2011). In sika deer, the first antler for males is a simple straight antler (spike), which develops between 1.5 and 2.5 years old. ...
... Forest habitats at our study site have been shown to yield foods with much better quality for deer than grasslands (Pei and Chen, 2004). The phenomenon of sexual segregation therefore could in part be to reduce the competition for food resources among individuals (Padmalal and Takatsuki, 1994;Asada and Ochiai, 1996;Pei and Lee, 1999;Jiang et al., 2006). Alternatively, the presence of free-ranging dogs (Canis lupus familiaris) in the area are considered a potentially serious threat to free-ranging sika deer (Jeng, 2003). ...
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The Formosan sika deer was declared extinct in the wild in Taiwan around 1969. In 1994, a series of reintroductions was initiated in Taiwan’s Kenting National Park. Today a wild population is established in the core area of the Park, and even continues to expand. Between October 2016 to March 2018, we used Unmanned Aerial Vehicles (UAV) to survey the herd composition of the sika deer in the Shuiwaku Grassland, a forest-grassland mosaic habitat inside the park. We completed a total of 56 early morning surveys (range: 2–7 per month) during the study period. The photographs and videos collected by UAV were used to identify the age and/or sex of each individuals recorded, and to generate data on deer group size, sex ratio, age-group ratio for each herd, and solitary animals. Overall, we cumulatively detected 394 non-independent deer groups comprised of 4732 individuals, and an additional 130 solitary animals. We found that outside of the rutting season (February–September), adult females remained in the adjacent forest the majority of the time. When females moved into the grassland, they formed small-stable groups with young deer (average size: 6.7 ± 4.5 during the post-rutting season; 6.3 ± 3.2 during the calving season). Males either occupied mixed sex groups (average size: 18.0 ± 9.1 during the post-rutting season; 13.8 ± 7.3 during the calving season), or formed bachelor groups (average size: 5.6 ± 4.3 during the post-rutting season; 5.7 ± 3.8 during the calving season) in the open grassland during this same period. During the territorial rut, the proportion of solitary adult males increased significantly to 24.2 ± 15.4%, whereas females also formed gregarious but unstable mixed sex groups with males (average size: 17.0 ± 12.0). Although we observed the first newborns in the open grassland in August (2017), the ratio of fawns to adult females was highest (56.8%) in October, suggesting that this may be the optimal time to conduct annual birth rate surveys. Overall, we found sexual segregation was present in the Formosan sika deer largely driven by sex-based differences in habitat preference, and the non-random assortment of sexes in response to their reproductive status.
... Deer are excellent models for studying evolution, biodiversity, interspecies hybridization (Abernethy 2008;Derr et al. 1991), social organization (i.e. hierarchical status) (Bartos and Bubenik 2011), unique organ development (i.e. fully regenerable antlers) (Li et al. 2009) and habitat selection (Blix 2016;Qiao et al. 2006). ...
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Deer species has both scientific research and economic value, and half of these species, however, are listed as endangered animals. For the conservation purpose, we designed a novel universal deer-specific PCR primer pair based on an evolutionarily conservative coding sequence (i.e., CEP295NL gene) across some deer species. This primer pair was successfully amplified and sequenced, showing around ~ 540 bp in cervids. Validation results showed that it can be utilized to develop a reliable and simple diagnostic tool for distinguishing other closely related species, as well as possibly interspecific identification amongst cervids.
... For instance, some birds have partial migration (Wingfield, 2005). In male elk, variation in ornaments signals male status and is related to androgen level (Bartos & Bubenik, 2011;Bartos et al., 2012). In termites, caste specialization is induced by interaction with other individuals via modification of juvenile hormone titer, and this contributes to the maintenance of the colony (Watanabe et al., 2014). ...
Article
Competitive interaction among individuals of a single population may result in the differentiation of two or more distinct life-history tactics. For example, although they exhibit unimodal size distribution, male juveniles of salmonids differentiate into those going down to the ocean to grow and returning to the natal stream after several years to reproduce (migratory tactic) and those staying in the stream and reproducing for multiple years (resident tactic). In this study, we developed a simple mathematical model for the positive feedback between hormonal and behavioral dynamics, with the expectation of establishing multiple discrete clusters of hormone levels leading to differentiation of life-history tactics. The assumptions were that probability of winning in fighting depends both on the body size and hormone level of the two contestants. An individual with a higher hormone level would be more likely to win the competition, which further enhanced hormone production, forming a positive feedback loop between hormone level and fighting ability. If the positive feedback was strong but not excessive, discrete clusters of hormone levels emerged from a continuous distribution. In contrast, no clear clustering structure appeared in the distribution of hormone levels if the probability of winning in fighting was controlled by the body size.
... Previous studies have shown that agonistic activity associated with rank position acquisition and https://doi.org/10.1016/j.ygcen.2020.113552 Received 11 February 2020; Received in revised form 13 July 2020; Accepted 14 July 2020 maintenance affected antler growth in red and fallow deer Dama dama (Bartoš and Bubenik, 2011) because it modified testosterone (T) and cortisol (C) concentrations (Bartoš et al. 2009; Bartoš et al. 2010; Bartoš et al. 2000). In the current study, there is also a focus on the period of antler growth, although studying antler growth itself is not covered here. ...
Article
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.
... The growth rate and size of the first antlers developed by young males in the second year of life depend on the quality of food provided by their mothers and by the habitat and ecological conditions existing during the winter period (García et al. 1999;Gómez et al. 2008;Janiszewski et al. 2008). Bartoš (1980) as well as Bartoš and Bubenik (2011) have shown that behavioural aspects, e.g. dominance in group and social structures or other stress factors, can also be a cause of formation of larger antlers in deer. ...
... Moreover, the final antler size was negatively related to the leukocyte count and the body condition had a more important effect on the final antler size, which was bigger than that exerted by the body weight, as also confirmed by Gaspar-López et al. (2011). However, one cannot forget that the size of antlers can also be influenced by animal behaviour and their social structure (Bartoš 1980;Bartoš and Bubenik 2011). ...
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The aim of the study was to assess the effect of supplementation of feed rations with increased calcium and phosphorus doses on the haematological parameters and plasma zinc (Zn), phosphorus (P), magnesium (Mg), copper (Cu), calcium (Ca), and iron (Fe) content as well as the body weight and the growth and development of the first antler in farmed fallow deer (Dama dama Linnaeus, 1758). The mean level of erythrocytes (RBC), haemoglobin (HGB), and haematocrit (HCT) was increased in the Ca- and P-supplemented group after the treatment period. The change was statistically significant (p < 0.05) in the case of RBC and HCT. The other haematological parameters (mean corpuscular volume (MCV), mean corpuscular haemoglobin (MCH), mean corpuscular haemoglobin concentration (MCHC), and platelet count (PLT)) were reduced. An increase in the Zn content was observed in the plasma of slaughtered animals. The concentration of other minerals (P, Mg, and Cu only in group II receiving a higher level of Ca and P in the feeding dose; Ca and Fe only in group I supplemented with a lower content of Ca and P in a nutritional dose) in blood plasma decreased slightly after the supplementation period and declined further after the slaughter. Noteworthy, there was a significant increase in the plasma Cu and Fe levels in group I in group II, respectively, in the post-supplementation period. No significant differences were observed in the body weight between the groups, but there was a beneficial effect of the higher Ca and P dose in the feed ration for the farmed fallow deer on the length of the first antler (p < 0.05). The antlers of animals in group II were on average 2.3 cm longer than in group I.