The vomeronasal system in semiaquatic beavers.

In contrast to the main olfactory system that detects volatile chemicals in the nasal air, the vomeronasal system can detect nonvolatile chemicals as well as volatiles. In the vomeronasal system, chemicals are perceived by the vomeronasal organ (VNO) projecting axons to the accessory olfactory bulb (AOB). Beavers (Castor spp.) are semiaquatic mammals that have developed chemical communication. It is possible that the beaver's anal gland secretions, nonvolatile and insoluble substances, may work as a messenger in the water and that beavers may detect the nonvolatile chemicals floating on the water surface via the VNO. The present study aimed to clarify the specificities of the beaver vomeronasal system by histologically and immunohistochemically analyzing the VNO and AOB of 12 Eurasian beavers (C. fiber). The VNO directly opened to the nasal cavity and was independent of a narrow nasopalatine duct connecting the oral and nasal cavities. The VNO comprised soft tissues including sensory and nonsensory epithelium, glands, a venous sinus, an artery, as well as cartilage inner, and bone outer enclosures. The AOB had distinct six layers, and anti-G protein α-i2 and α-o subunits were, respectively, immunoreactive in rostral and caudal glomeruli layers indicating expressions of V1Rs and V2Rs. According to gene repertories analysis, the beavers had 23 and six intact V1R and V2R genes respectively. These findings suggested that beavers recognize volatile odorants and nonvolatile substances using the vomeronasal system. The beaver VNO was developed as well as in other rodents, and it had two specific morphological features, namely, disadvantaged contact with the oral cavity because of a tiny nasopalatine duct, and a double bone and cartilage envelope. Our results highlight the importance of the vomeronasal system in beaver chemical communication and support the possibility that beavers can detect chemicals floating on the water surface via the VNO.

Retention and loss of PIT tags and surgically implanted devices in the Eurasian beaver.

BACKGROUND: Passive integrated transponder devices (PIT tags) are a valuable tool for individual identification of animals. Similarly, the surgical implantation of transmitters and bio-loggers can provide useful data on animal location, physiology and behavior. However, to avoid unnecessary recapture and related stress of study animals, PIT tags and bio-loggers should function reliably for long periods of time. Here, we evaluated the retention of PIT tags, and of very high frequency (VHF) transmitters and bio-loggers that were either implanted subcutaneously or into the peritoneal cavity of Eurasian beavers (Castor fiber). RESULTS: Over a 21-year period, we implanted PIT tags in 456 individuals and failed to detect a PIT tag at recapture in 30 cases, consisting of 26 individuals (6% of individuals). In all instances, we were still able to identify the individual due to the presence of unique ear tag numbers and tail scars. Moreover, we implanted 6 VHFs, 36 body temperature loggers and 21 heart rate loggers in 28 individuals, and experienced frequent loss of temperature loggers (at least 6 of 23 recaptured beavers) and heart rate loggers (10 of 18 recaptured beavers). No VHFs were lost in 2 recaptured beavers. CONCLUSIONS: Possible causes for PIT tag loss (or non-detection) were incorrect implantation, migration of the tag within the body, a foreign body reaction leading to ejection, or malfunctioning of the tag. We speculate that logger loss was related to a foreign body reaction, and that loggers were either rejected through the incision wound or, in the case of temperature loggers, possibly adhered and encapsulated to intestines, and then engulfed by the gastro-intestinal tract and ejected. We discuss animal welfare implications and give recommendations for future studies implanting bio-loggers into wildlife.

Fears from the past? The innate ability of dogs to detect predator scents.

Throughout the animal kingdom, antipredator mechanisms are an evolutionary driving force to enable the survival of species classified as prey. Information regarding a predator's location can be determined through chemosensory cues from urine, faeces, visual and/or acoustic signals and anal gland secretions; and in several lab and field-based studies it has been seen that these cues mediate behavioural changes within prey species. These behaviours are often linked to fear and avoidance, which will in turn increase the prey's survival rate. In many studies dogs (Canis lupus familiaris) have been used as a predator species, however, no research has addressed a dog's innate ability to detect predator scents, hence the rationale behind this study. We assessed the innate ability of the untrained domestic dog to detect faecal scents of wild Eurasian brown bear (Ursus arctos arctos) and European lynx (Lynx lynx). The study monitored 82 domestic dogs across the UK and Norway. The dogs were exposed to the two predator faecal scents from Eurasian brown bear and European lynx, a herbivore faecal scent of Eurasian beaver (Castor fiber) and water control. Measurements were taken upon the time spent within a 40 cm radius of each scent and changes in the dog's heart rate when within this 40 cm radius. We found dogs spent a decreased length of time around the predator scents and had an increased heart rate in relation to their basal heart rate. We conclude that dogs can innately sense predator scents of brown bear and lynx and elicit fear towards these odours, as shown through behavioural and physiological changes.

Causes and consequences of inverse density-dependent territorial behaviour and aggression in a monogamous mammal.

Territoriality is an important process shaping population dynamics, and the defence of a territory is crucial for individuals to increase the duration of territory occupancy and, consequently, reproductive success. However, little is known about how the frequency of territory intrusions and subsequent territorial behaviours and aggression by territory owners are affected by external factors, such as population density. This is important because it can affect mate change (the replacement of one pair member) and dispersal, a key ecological process. The aim of this study was to investigate the behavioural and spatial response of territory owners to intruder pressure as a function of population density in a territorial, monogamous mammal, the Eurasian beaver (Castor fiber). Using a combination of GPS technology, scent experiments, camera trap data and tail scar observations from an individual-based long-term study, we investigated the factors influencing spatial movement patterns by territory owners in response to a simulated intruder and the factors affecting territory intrusions. We found consistent inverse density-dependent patterns in territorial behaviours and evidence of conspecific aggression. At lower densities, territory owners detected more simulated intrusions, showed more territorial reactions and experienced increased conspecific aggression as indicated by tail scars, suggesting increased intruder pressure. Inverse density-dependent territorial behaviour and aggression suggest a potential mechanistic link between inverse density-dependent natal dispersal and mate change. At low population densities, increased dispersal amplifies intruder pressure, leading to the observed increases in territorial behaviours, conspecific aggression and previously observed mate turnover, which in turn might increase natal dispersal. Our study demonstrates how population density can affect the behaviour and space use of individuals, which is important for territory occupancy and fitness.

Estimates for energy expenditure in free-living animals using acceleration proxies: A reappraisal.

It is fundamentally important for many animal ecologists to quantify the costs of animal activities, although it is not straightforward to do so. The recording of triaxial acceleration by animal-attached devices has been proposed as a way forward for this, with the specific suggestion that dynamic body acceleration (DBA) be used as a proxy for movement-based power. Dynamic body acceleration has now been validated frequently, both in the laboratory and in the field, although the literature still shows that some aspects of DBA theory and practice are misunderstood. Here, we examine the theory behind DBA and employ modelling approaches to assess factors that affect the link between DBA and energy expenditure, from the deployment of the tag, through to the calibration of DBA with energy use in laboratory and field settings. Using data from a range of species and movement modes, we illustrate that vectorial and additive DBA metrics are proportional to each other. Either can be used as a proxy for energy and summed to estimate total energy expended over a given period, or divided by time to give a proxy for movement-related metabolic power. Nonetheless, we highlight how the ability of DBA to predict metabolic rate declines as the contribution of non-movement-related factors, such as heat production, increases. Overall, DBA seems to be a substantive proxy for movement-based power but consideration of other movement-related metrics, such as the static body acceleration and the rate of change of body pitch and roll, may enable researchers to refine movement-based metabolic costs, particularly in animals where movement is not characterized by marked changes in body acceleration.

Knowing Me, Knowing You: Anal Gland Secretion of European Badgers (Meles meles) Codes for Individuality, Sex and Social Group Membership.

European badgers, Meles meles, are group-living in the UK, and demarcate their ranges with shared latrines. As carnivores, badgers possess paired anal glands, but olfactory information on the content of badger anal gland secretion (AGS) is largely uninvestigated. Here, we examined the volatile organic compounds (VOCs) of AGS samples from 57 free-living badgers using solid-phase microextraction (SPME) and gas chromatography-mass spectrometry. AGS was rich in alkanes (C7-C15, 14.3% of identified compounds), aldehydes (C5-C14, 9.7%), phenols (C6-C15, 9.5%), alcohols (C5-C10, 7.3%), aromatic hydrocarbons (C6-C13, 6.8%), ketones (C6-C13, 6.3%) and carboxylic acids (C3-C12, 5.6%) and contained a variety of esters, sulfurous and nitrogenous compounds, and ethers. The number of VOCs per profile ranged from 20 to 111 (mean = 65.4; ± 22.7 SD), but no compound was unique for any of the biological categories. After normalization of the raw data using Probabilistic Quotient Normalization, we produced a resemblance matrix by calculating the Euclidian distances between all sample pairs. PERMANOVA revealed that AGS composition differs between social groups, and concentration and complexity in terms of number of measurable VOCs varies between seasons and years. AGS VOC profiles encode individual identity, sex and vary with female reproductive state, indicating an important function in intraspecific communication. Because AGS is excreted together with fecal deposits, we conclude that chemical complexity of AGS enables particularly latrine-using species, such as badgers, to advertise more complex individual-specific information than in feces alone.

Sociodemographic factors modulate the spatial response of brown bears to vacancies created by hunting.

There is a growing recognition of the importance of indirect effects from hunting on wildlife populations, e.g. social and behavioural changes due to harvest, which occur after the initial offtake. Nonetheless, little is known about how the removal of members of a population influences the spatial configuration of the survivors. We studied how surviving brown bears (Ursus arctos) used former home ranges that had belonged to casualties of the annual bear hunting season in southcentral Sweden (2007-2015). We used resource selection functions to explore the effects of the casualty's and survivor's sex, age and their pairwise genetic relatedness, population density and hunting intensity on survivors' spatial responses to vacated home ranges. We tested the competitive release hypothesis, whereby survivors that increase their use of a killed bear's home range are presumed to have been released from intraspecific competition. We found strong support for this hypothesis, as survivors of the same sex as the casualty consistently increased their use of its vacant home range. Patterns were less pronounced or absent when the survivor and casualty were of opposite sex. Genetic relatedness between the survivor and the casualty emerged as the most important factor explaining increased use of vacated male home ranges by males, with a stronger response from survivors of lower relatedness. Relatedness was also important for females, but it did not influence use following removal; female survivors used home ranges of higher related female casualties more, both before and after death. Spatial responses by survivors were further influenced by bear age, population density and hunting intensity. We have shown that survivors exhibit a spatial response to vacated home ranges caused by hunting casualties, even in nonterritorial species such as the brown bear. This spatial reorganization can have unintended consequences for population dynamics and interfere with management goals. Altogether, our results underscore the need to better understand the short- and long-term indirect effects of hunting on animal social structure and their resulting distribution in space.

Socio-ecological features other than sex affect habitat selection in the socially obligate monogamous Eurasian beaver.

Habitat selection is a context-dependent mechanism, in which both the internal state as well as external factors affect the behavior and decisions of an individual. This is well known for polygamous mammals, which are typically sexually dimorphic, and often express great variability in behavior and habitat selection between individuals as well between the sexes. Among monogamous mammals, however, variability in habitat selection should be explained by group characteristics and the presence of offspring rather than by sex. We evaluated this hypothesis in a socially monogamous rodent, the Eurasian beaver (Castor fiber), in a saturated Norwegian population. For the first time in this species we applied GPS tracking devices (N = 22 adult beavers, in 15 territories, 2009-2013), and used resource selection functions (i) to document population-wide habitat selection and the importance of 'territory' therein, and (ii) to evaluate which socio-ecological factors explained potential individual differences in habitat selection. We found that variation in habitat selection was stronger between territories than between years or individuals nested by territory. We identified that family size and the presence of kits, but not sex, explained individual variation in habitat selection. Adults with kits and/or larger families tended to exhibit low risk-taking behavior (avoiding human-related variables such as roads, buildings, and agricultural land), and stayed close to their main lodge (parental care). Our results show that habitat selection is a context-dependent mechanism even in a species which expresses very little behavioral and morphological dimorphism.

Captive care and welfare considerations for beavers.

Beavers (Castor spp.) tend not to be a commonly held species and little published material exists relating to their captive care. We review published material and discuss husbandry issues taking into account the requirements of wild beavers. As social mammals with complex chemical communication systems and with such an ability to modify their environments, studies of wild counterparts suggest the captive requirements of beavers may actually be more sophisticated than generally perceived. Common field techniques may have practical application in the captive setting. Their widespread utilisation in conservation, including reintroductions, translocations and habitat management, also requires components of captive care. As welfare science advances there is increasing pressure on captive collections to improve standards and justify the keeping of animals. Conservation science is increasingly challenged to address individual welfare standards. Further research focusing on the captive care of beavers is required.

Coronary Artery Bypass Grafting in a Patient With Multivessel Disease and Dextrocardia With Situs Inversus Totalis.

Dextrocardia with situs inversus totalis is a rare hereditary condition characterized by reversed orientation of the major thoracic and abdominal organs. Though dextrocardia itself is not believed to increase the risk of coronary artery disease, the workup and surgical management of patients with this condition may be technically challenging to heart team clinicians. This report describes the case management of a high-risk 56-year-old man with dextrocardia who presented with multivessel coronary artery disease.

Individual recognition of Eurasian beavers (Castor fiber) by their tail patterns using a computer-assisted pattern-identification algorithm.

Individual recognition of animals is an important aspect of ecological sciences. Photograph-based individual recognition options are of particular importance since these represent a non-invasive method to distinguish and identify individual animals. Recent developments and improvements in computer-based approaches make possible a faster semi-automated evaluation of large image databases than was previously possible. We tested the Scale Invariant Feature Transform (SIFT) algorithm, which extracts distinctive invariant features of images robust to illumination, rotation or scaling of images. We applied this algorithm to a dataset of 800 tail pattern images from 100 individual Eurasian beavers (Castor fiber) collected as part of the Norwegian Beaver Project (NBP). Images were taken using a single-lens reflex camera and the pattern of scales on the tail, similar to a human fingerprint, was extracted using freely accessible image processing programs. The focus for individual recognition was not on the shape or the scarring of the tail, but purely on the individual scale pattern on the upper (dorsal) surface of the tail. The images were taken from two different heights above ground, and the largest possible area of the tail was extracted. The available data set was split in a ratio of 80% for training and 20% for testing. Overall, our study achieved an accuracy of 95.7%. We show that it is possible to distinguish individual beavers from their tail scale pattern images using the SIFT algorithm.

Reference-free SNP discovery for the Eurasian beaver from restriction site-associated DNA paired-end data.

In this study, we used restriction site-associated DNA (RAD) sequencing to discover SNP markers suitable for population genetic and parentage analysis with the aim of using them for monitoring the reintroduction of the Eurasian beaver (Castor fibre) to Scotland. In the absence of a reference genome for beaver, we built contigs and discovered SNPs within them using paired-end RAD data, so as to have sufficient flanking region around the SNPs to conduct marker design. To do this, we used a simple pipeline which catalogued the Read 1 data in stacks and then used the assembler cortex_var to conduct de novo assembly and genotyping of multiple samples using the Read 2 data. The analysis of around 1.1 billion short reads of sequence data was reduced to a set of 2579 high-quality candidate SNP markers that were polymorphic in Norwegian and Bavarian beaver. Both laboratory validation of a subset of eight of the SNPs (1.3% error) and internal validation by confirming patterns of Mendelian inheritance in a family group (0.9% error) confirmed the success of this approach.

The influence of mean climate trends and climate variance on beaver survival and recruitment dynamics.

Ecologists are increasingly aware of the importance of environmental variability in natural systems. Climate change is affecting both the mean and the variability in weather and, in particular, the effect of changes in variability is poorly understood. Organisms are subject to selection imposed by both the mean and the range of environmental variation experienced by their ancestors. Changes in the variability in a critical environmental factor may therefore have consequences for vital rates and population dynamics. Here, we examine ≥90-year trends in different components of climate (precipitation mean and coefficient of variation (CV); temperature mean, seasonal amplitude and residual variance) and consider the effects of these components on survival and recruitment in a population of Eurasian beavers (n = 242) over 13 recent years. Within climatic data, no trends in precipitation were detected, but trends in all components of temperature were observed, with mean and residual variance increasing and seasonal amplitude decreasing over time. A higher survival rate was linked (in order of influence based on Akaike weights) to lower precipitation CV (kits, juveniles and dominant adults), lower residual variance of temperature (dominant adults) and lower mean precipitation (kits and juveniles). No significant effects were found on the survival of nondominant adults, although the sample size for this category was low. Greater recruitment was linked (in order of influence) to higher seasonal amplitude of temperature, lower mean precipitation, lower residual variance in temperature and higher precipitation CV. Both climate means and variance, thus proved significant to population dynamics; although, overall, components describing variance were more influential than those describing mean values. That environmental variation proves significant to a generalist, wide-ranging species, at the slow end of the slow-fast continuum of life histories, has broad implications for population regulation and the evolution of life histories.

Intraoperative echocardiographic contrast opacifies the left atrial appendage and assists in surgical exclusion.

Surgical exclusion of the left atrial appendage (LAA) for stroke prevention in atrial fibrillation is frequently incomplete and remains to be optimized. We present a man who did not tolerate anticoagulant and failed percutaneous occlusion. Intraoperative echocardiographic contrast was used to assist surgical exclusion. Follow-up showed a persistent occlusion.

Pest detection dogs for wood boring longhorn beetles.

Invasive alien species are increasing due to globalization. Their spread has resulted in global economic losses. Asian [Anoplophora glabripennis (Motschulsky)] (ALB) and citrus [A. chinensis (Forster)] (CLB) longhorn beetles are two introduced wood borers which contribute to these economic losses e.g. the destruction of tree plantations. Early detection is key to reduce the ecological influence alongside the detrimental and expensive eradication. Dogs (Canis lupus familiaris) can detect these insects, potentially at an early stage. We trained two privately owned dogs to investigate their use as detection tools. We tested the dog's ability to discriminate ALB and CLB from native wood borers by carrying out double-blind and randomized experiments in three search conditions; (1) laboratory, (2) semi-field and (3) standardized field. For condition one, a mean sensitivity of 80%, specificity of 95% and accuracy of 92% were achieved. For condition two and three, a mean sensitivity of 88% and 95%, specificity of 94% and 92% and accuracy of 92% and 93% were achieved. We conclude that dogs can detect all types of traces and remains of ALB and CLB and discriminate them from native wood borers and uninfested wood, but further tests on live insects should be initiated.

The impact of bio-logging on body weight change of the Eurasian beaver.

Bio-logging is a common method to collect ecological data on wild animals, but might also induce stress, reduce body condition, and alter behavior. Eurasian beavers (Castor fiber) are a semi-aquatic and nocturnal species that are challenging to observe in the wild. Bio-loggers are hence useful tools to study their behaviour and movements, but this raises concerns of potential negative impacts of tagging. To investigate the potential negative impacts of glue-on tags, we compared body weight change for tagged and untagged Eurasian beavers. We hypothesized that tagged beavers would gain less body weight compared to untagged beavers, and that weight change might be affected by tagging length, tag weight, water temperature and the season of tagging. Daily percentage body weight change in relation to initial body weight during the first capture was compared during 57 tagging periods (18±7 days) and 32 controls periods (64±47 days). Body weight change varied between the two groups, with untagged beavers on average gaining daily weight whilst tagged beavers on average lost weight daily, indicating a negative effect of tagging. The average reduction in percentage body weight change per day for tagged beavers was small (0.1 ± 0.3%), and with large individual variation. Neither tag weight, number of tagging days, nor season were important in explaining body weight change of tagged animals. In other words, we found that tagging reduced daily body weight during the tagging period but were unable to determine the mechanism(s) responsible for this decline. Detrimental effects of tagging have important implications for animal welfare and can introduce bias in data that are collected. This calls for careful consideration in the use of tags. We conclude that studies investigating the effects of tagging should consider individual variation in the effects of tagging and, where possible, compare tagged animals with a control group.

Eurasian beaver (Castor fiber) health surveillance in Britain: Assessing a disjunctive reintroduced population.

BACKGROUND: Numerous translocations of Eurasian beavers have occurred with little implementation of standardised health screening. Pre-release health screening enables the selection of individuals with the best survival prospects and reduces potential health risks, but this is by-passed during unofficial releases. Beaver reintroduction to Britain has been haphazard and currently disjunctive populations of varying status exist. METHODS: This observational cross section study investigated the health status of three beaver populations, with 90 live beavers tested for a range of pathogens comprising 56 from Tayside (unofficially released Scotland), nine from Knapdale (officially released Scotland) and 25 from Devon (unofficially released England). In addition, a further 32 cadavers were screened (25 from Tayside and seven from Knapdale). RESULTS: All beavers were in good physical condition, did not harbour any non-native disease or parasites of concern and demonstrated remarkably low levels of any disease or parasite exposure. CONCLUSION: Beavers are establishing and adapting well to British landscapes and are not acting as reservoirs of significant zoonotic diseases. Official, licensed reintroduction programmes may appear overly convoluted; however, reputational damage of unofficial releases should be considered, along with the health and welfare of the animals involved and collateral damage to other wildlife, domestic animals and humans.

Conspecific recognition of pedal scent in domestic dogs.

Carnivores rely heavily on scent to communicate with conspecifics. Scent glands located on the underside of the feet provide an especially efficient way of leaving a scent trail. Although domestic dogs (Canis familiaris) are well-known for their olfactory abilities and scent marking behaviours, their use of pedal scent for communication remains unknown. We studied the reaction of intact dogs of both sexes to male and female pedal scent as well as a control sample of scent taken from the ground, using sniffing time and nostril usage as an indicator of interest level and emotional valence. In male subjects, only the sniffing duration for other males differed from the control samples, with no clear difference detected between male and female scent. Females showed no difference in the sniffing duration for any sample type. Conversely, male nostril use did not differ between the sample types, whereas females demonstrated a right nostril bias when sniffing the scent from other females and a left nostril bias when sniffing the control. We have shown that dogs recognize scent taken from the pedal glands from other dogs, although the extent to which they use this information to determine the sex of the scent depositor remains unclear.

Long-term capture and handling effects on body condition, reproduction and survival in a semi-aquatic mammal.

In long-term individual-based field studies, several parameters need to be assessed repeatedly to fully understand the potential fitness effects on individuals. Often studies only evaluate capture stress that appears in the immediate weeks or breeding season and even long-term studies fail to evaluate the long-term effects of their capture procedures. We investigated effects of long-term repeated capture and handling of individuals in a large semi-aquatic rodent using more than 20 years of monitoring data from a beaver population in Norway. To investigate the effects, we corrected for ecological factors and analysed the importance of total capture and handling events, years of monitoring and deployment of telemetry devices on measures related to body condition, reproduction and survival of individual beavers. Body mass of dominant individuals decreased considerably with number of capture events (107 g per capture), but we found no statistically clear short or long-term effects of capture and handling on survival or other body condition indices. Annual litter size decreased with increasing number of captures among older individuals. Number of captures furthermore negatively affected reproduction in the beginning of the monitoring, but the effect decreased over the years, indicating habituation to repeated capture and handling. By assessing potential impacts on several fitness-related parameters at multiple times, we can secure the welfare of wild animal populations when planning and executing future conservation studies as well as ensure ecologically reliable research data.

Landscape structure and population density affect intraspecific aggression in beavers.

Intraspecific competition plays an important role for territory acquisition and occupancy, in turn affecting individual fitness. Thus, understanding the drivers of intraspecific aggression can increase our understanding of population dynamics. Here, we investigated intraspecific aggression in Eurasian (Castor fiber) and North American (Castor canadensis) beavers that are both monogamous, territorial mammals. Combined, we examined tail scars from >1,000 beavers (>2,000 capture events) as part of two long-term studies in Norway and the USA. We investigated the influence of landscape structure, population density, sex, age, and (for Eurasian beavers only) social status and group size on the number of tail scars caused by conspecifics. The number of tail scars was affected by population density in well-connected landscape types (large lakes and rivers), but not in more isolated areas (ponds), where individuals generally had fewer tail scars. Further, the relationship of population density was not linear. In the North American beaver population occurring in large lakes, intraspecific aggression increased with population density. Conversely, in the saturated Eurasian beaver population, intraspecific aggression was in a negative relationship with population density (except at the highest densities), likely due to inverse density-dependent intruder pressure via dispersers. Our findings emphasize that population density can affect intraspecific aggression depending on landscape structure, which might have important consequences for local patterns of dispersal, mate change, and territory occupancy, all of which can affect population dynamics.