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.

Aggressive interactions among female, semi-captive pampas deer (Ozotoceros bezoarticus) increase within the hierarchy and after short-term removal of the male.

Two trials were carried out to determine: (a) if there is a positive relation between the frequency of aggressive interactions among female pampas deer and their position within the hierarchy (HI), (b) if short-term removal of the male triggers an increase in the frequency of aggression, and (c) if the magnitude of this increase is related to the individual rank position of the female. Each of 19 breeding groups comprised one adult male and from four to six adult females. The HI was determined for each female and all aggressive interactions were recorded. These were recorded while the male was present (i.e., all of Trial 1 and the "with male" period in Trial 2) and after removal of the male (i.e., the "without male" period in Trial 2). The individual percentage change in the frequency of aggressive interactions after male removal was calculated. In Trial 1 the HI, the frequency of different types of aggressive interaction and the total of aggressive interactions were positively related (all: p < .001; General Linear Mixed Model [GLMM]). In Trial 2, the frequency of total aggressive interactions increased after male removal (F(1, 27) = 3.5; p < .001; GLMM). The individual percentage changes in aggressive interaction between periods were positively related to HI (F(6, 24) = 2.56; p = .05; GLMM). For female pampas deer maintained in breeding groups, we conclude that the frequency of aggressive interactions increases within the hierarchy. Aggression also increases after the short-term removal of the male, mainly among females of higher social status.

A wolfdog point of view on the impossible task paradigm.

To elucidate the role of domestication, we used the impossible task paradigm to test Czechoslovakian Wolfdogs with a known proportion of 'wolfblood' in their DNA and, as a control group for our subjects, we used German shepherd dogs. We hypothesized that the difference between wolves and domestic dogs is based on genetics and modified by obedience; if so, the looking back performance of the subject should be linked to its proportion of wolf-genes. To prove that, we observed 73 Czechoslovakian Wolfdogs, and 27 German shepherd dogs, and analysed their human-directed gazing behaviour during our test. Our apparatus consisted of a glass container placed upside down over a small amount of food. The test proceeded with three solvable trials, in which the subject could obtain the food by manipulating the container, followed by an unsolvable one in which the container was fixed onto the board. Our results suggest that there is no significant correlation between the probability of looking back in Czechoslovakian Wolfdogs and their proportion of 'wolf blood'. However, the probability of looking back was higher in German Shepherd dogs than in Czechoslovakian Wolfdogs (odds ratio = 9.1). German Shepherd dogs showed not only a higher frequency of looking back, but also the duration of their looks was longer.

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.

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.

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.

Effects of Genotype and Housing System on Rabbit Does' Aggressive Behaviors and Injuries in Smallholding Conditions.

The objective of the study was to investigate the effects of housing (deep litter + concrete floor vs. deep litter + ground soil with the possibility to dig burrows), and genotype (Mecklenburg or Hyplus) on aggressive behaviour, social contacts, does' and kits' injuries, and progeny mortality. Twelve groups of six rabbit does (n = 72) were assigned to four treatments (two housing systems and two genotypes). Aggressive behaviour of does, number of injuries on does and kits, and postnatal kit mortality were recorded. The effects of housing and genotype were tested using multivariate GLMM Models. We found that the housing treatment in interaction with the genotype had a significant effect on aggressive behaviours in group housed does (F3,12 = 14.34, p = 0.0003), where the lowest incidence of aggression was in Mecklenburg does housed on ground soil. Reduced aggression was reflected in a lower number of injuries in does (F3,68 = 10.51, p < 0.0001), number of injuries in kits, and kit mortality (F3,1 = 4.59, p < 0.0001, F3,54 = 43.94, p < 0.0001). The results indicate that the proper combination of genotype and housing should be carefully considered for breeding to reduce aggression and injury in group housed does.

Evolutional background of dominance/submissivity in sex and bondage: the two strategies?

OBJECTIVES: We theorize that sexual arousal by dominance and submission may be connected to a reproduction strategy respecting a reached social dominance rank (a common reproduction strategy in socially living mammals), while the preference for "bondage" may be derived from an opportunistic strategy when being unable to compete for hierarchic rank (an alternative reproductive strategy that co-occurs frequently with the above-named main strategy). The answers to questions dealing with hierarchy in character should correlate exclusively with sexual arousal connected to any kind of expression of a hierarchy, but not with bondage. DESIGN AND SETTINGS: The data were obtained from young adults (157 males and 183 females aged 18-20, with mean 18.4 years) via questionnaires. RESULTS: Seven out of eight questions dealing with hierarchy correlated with sexual arousal by dominance and submission in men (Spearman's r=0.169-0.313; p<0.05 - p<0.001), two questions correlated with sexual arousal by dominance and submission in women (Spearman's r=0.32-0.166, p<0.001, p<0.05). THE MAIN FINDINGS: The questions dealing with hierarchy correlated with sexual arousal by dominance and submission while no answers correlated with bondage, neither in men nor in women. CONCLUSION: The preference for sexual arousal by dominance and submission may be connected to strategy respecting rank, while the preference for "bondage" may be derived from an opportunistic strategy that may be essential for possible partner problems solution. From the evolutionary biology point of view, these patterns of sadomasochistic sex appear as adaptive rather than as pathology.

The transition from the female-like great calls to male calls during ontogeny in southern yellow-cheeked gibbon males (Nomascus gabriellae).

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.

Long-term trends in the body condition of parents and offspring of Tengmalm's owls under fluctuating food conditions and climate change.

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.

Turning preference in dogs: North attracts while south repels.

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.

Developmental instability of ano-genital distance index: Implications for assessment of prenatal masculinization.

Prenatal androgens are generally assumed to permanently affect the morphology of external genitalia. In laboratory rodents, the ano-genital distance index (AGDI) has become widely used as a marker of natural prenatal masculinization. A greater value should indicate a greater masculinization. The aim of this study was to assess the developmental stability of the AGDI in female mice during pre- (Days 1-21) and postweaning (Days 21-61) periods. Presuming that the AGDI reflects a permanent effect of prenatal masculinization, we hypothesized that the AGDI will be a developmentally stable morphometric measurement (interindividual differences in the AGDI measurement should persist over time). In contrast to our prediction, the AGDI showed poor repeatability both during pre- (.15) and postweaning (.22) periods, indicating developmental instability. (A value of 1 indicates perfect repeatability and a value of 0 indicates no repeatability.) The AGDI thus does not seem to be a reliable marker of prenatal female masculinization.

Interactive influences of fluctuations of main food resources and climate change on long-term population decline of Tengmalm's owls in the boreal forest.

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.

Magnetic alignment enhances homing efficiency of hunting dogs.

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.

Early ontogeny social deprivation modifies future agonistic behaviour in crayfish.

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.

Specialized use of two fingers in free-ranging aye-ayes (Daubentonia madagascariensis).

The aye-aye (Daubentonia madagascariensis) possesses a highly specialized hand with two fingers, the third and the fourth, being used in a way unparalleled by any other primate. We observed the use of the third and the fourth fingers in various activities in four free-ranging aye-ayes. We found that the thin third finger was used exclusively or preferably for tapping, inserting into the mouth (probably for cleaning the teeth) and probing for nectar, kernels and insects in bamboo, twigs and live wood. In contrast, the robust fourth finger was used preferably when eating jackfruit (Artocarpus heterophyllus). When probing for invertebrates in soft plant tissues and in dead wood, both fingers were used in high proportions. To extract the contents from coconuts, the two fingers were apparently used for different tasks. From this small (686 observations), but unique, study of free-ranging aye-ayes, we conclude that the third finger appears to be specialized for use in tasks requiring high mobility, sensitivity and precision, whereas the fourth finger appears to be specialized for tasks requiring strength, scooping action and deep access.

Increased resistance to sudden noise by audio stimulation during early ontogeny in German shepherd puppies.

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<0.01). Interestingly, gender was seen to affect response, with the males scoring more than the females (P<0.1). In conclusion, the results suggested that audio stimulation early in life improved the response of the dogs to intense sudden noise, as caused by the shovel. Therefore, acoustic stimulation during the very early period of life has the potential to raise the necessary skills of dogs for military and police purposes, or civilian life.

Estimation of the probability of fighting in fallow deer (Dama dama) during the rut.

Fighting between males is a frequent component of the rutting behavior of Cervidae. Frequent conflicts are exhausting; fighting may be risky and can lead to serious injuries or even death. We focused on the process of assessment of the opponent's fighting ability and escalation of the combat, estimating the probability of fighting based on the encounter components such as groaning and parallel walk. In this study, we observed the agonistic behavior of fallow deer bucks (Dama dama) during the rut over four seasons. During this time, we recorded 205 encounters between bucks. Non-contact display, which allows contestants to assess their opponents fighting ability, occurred in 83% of the encounters. The highest predicted probability of a fight was found when both of the males vocalized and turned into the parallel walk. The chance of a clear outcome decreased when the males were fighting in comparison to when they did not fight. The initiator of the competitive encounter won 41% of the cases, while the attacked buck won 23% of the encounters. If the contestants avoided fighting, however, the initiator won 78% of encounters. Therefore, the initiator was more successful when no fight occurred compared to when the encounters escalated into fighting. In most cases where ritualized behavior occurred, one of the opponents left after vocalization or parallel walk occurred. Thus, vocalization and parallel walk increased the probability for a clear outcome. The probability of a fight was lowest in situations where the males displayed asymmetric behavior. Increased symmetry of the contestants' behavior was strongly correlated with a higher probability of a fight. Thus, these results indicate that fallow deer bucks use efficient tactic during the rut, which, in turn, minimizes the chance of injury while fighting during the breeding season.

Pre-breeding food restriction promotes the optimization of parental investment in house mice, Mus musculus.

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.