Long-term, climate-driven phenological shift in a tropical large carnivore.

Understanding the degree to which animals are shifting their phenology to track optimal conditions as the climate changes is essential to predicting ecological responses to global change. Species at low latitudes or high trophic levels are theoretically expected to exhibit weaker phenological responses than other species, but limited research on tropical systems or on top predators impedes insight into the contexts in which these predictions are upheld. Moreover, a lack of phenological studies on top predators limits understanding of how climate change impacts propagate through entire ecosystems. Using a 30-y dataset on endangered African wild dogs (Lycaon pictus), we examined changes in reproductive phenology and temperatures during birthing and denning over time, as well as potential fitness consequences of these changes. We hypothesized that their phenology would shift to track a stable thermal range over time. Data from 60 packs and 141 unique pack-years revealed that wild dogs have delayed parturition by 7 days per decade on average in response to long-term warming. This shift has led to temperatures on birthing dates remaining relatively stable but, contrary to expectation, has led to increased temperatures during denning periods. Increased denning temperatures were associated with reduced reproductive success, suggesting that a continued phenological shift in the species may become maladaptive. Such results indicate that climate-driven shifts could be more widespread in upper trophic levels than previously appreciated, and they extend theoretical understanding of the species traits and environmental contexts in which large phenological shifts can be expected to occur as the climate changes.

A comparative test of inequity aversion in domestic dogs (Canis familiaris) and dingoes (Canis dingo).

Despite much recent empirical work on inequity aversion in nonhuman species, many questions remain about its distribution across taxa and the factors that shape its evolution and expression. Past work suggests that domestic dogs (Canis familiaris) and wolves (Canis lupus) are averse to inequitable resource distributions in contexts that call upon some degree of training such as 'give paw' and 'buzzer press' tasks. However, it is unclear whether inequity aversion appears in other canid species and in other experimental contexts. Using a novel inequity aversion task that does not require specific training, this study helps address these gaps by investigating inequity aversion in domestic dogs and a closely related but non-domesticated canid, the dingo (Canis dingo). Subjects were presented with equal and unequal reward distributions and given the opportunity to approach or refuse to approach allocations. Measures of interest were (1) subjects' refusal to approach when getting no food; (2) approach latency; and (3) social referencing. None of these measures differed systematically across the inequity condition and control conditions in either dogs or dingoes. These findings add to the growing literature on inequity aversion in canids, providing data from a new species and a new experimental context. Additionally, they raise questions about the experimental features that must be in place for inequity aversion to appear in canids.

First successful frozen semen of the maned wolf (Chrysocyon brachyurus).

This study aimed to describe successful cryopreservation of sperm from maned wolves (Chrysocyon brachyurus). Three ejaculates from 2 maned wolves were collected by digital manipulation of the penis and evaluated subjectively, centrifuged and frozen in BotuCrio® (Botupharma, Botucatu, Brazil) or Tris-yolk egg extender. Spermatozoa were thawed at 37ºC/30s or 70ºC/4s and evaluated for kinetics, morphology, plasma and acrosome membrane integrity, mitochondrial potential, hydrogen peroxide, superoxide anion and lipid peroxidation. From 5 thawed samples, two had sperm total motility >55% (56.0% and 64.0%) and progressive motility ~35% (35% and 40%), both frozen with Tris-yolk egg. Plasma and acrosome membrane integrity decreased and percentage of sperm defects increased post-thawing. We concluded that is possible to freeze spermatozoa from maned wolves using semen collection and processing methods applied for domestic dogs.

Understanding potential implications for non-trophic parasite transmission based on vertebrate behavior at mesocarnivore carcass sites.

High infection risk is often associated with aggregations of animals around attractive resources. Here, we explore the behavior of potential hosts of non-trophically transmitted parasites at mesocarnivore carcass sites. We used videos recorded by camera traps at 56 red fox (Vulpes vulpes) carcasses and 10 carcasses of other wild carnivore species in three areas of southeastern Spain. Scavenging species, especially wild canids, mustelids and viverrids, showed more frequent rubbing behavior at carcass sites than non-scavenging and domestic species, suggesting that they could be exposed to a higher potential infection risk. The red fox was the species that most frequently contacted carcasses and marked and rubbed carcass sites. Foxes contacted heterospecific carcasses more frequently and earlier than conspecific ones and, when close contact occurred, it was more likely to be observed at heterospecific carcasses. This suggests that foxes avoid contact with the type of carcass and time period that have the greatest risk as a source of parasites. Overall, non-trophic behaviors of higher infection risk were mainly associated with visitor-carcass contact and visitor contact with feces and urine, rather than direct contact between visitors. Moreover, contact events between scavengers and carnivore carcasses were far more frequent than consumption events, which suggests that scavenger behavior is more constrained by the risk of acquiring meat-borne parasites than non-trophically transmitted parasites. This study contributes to filling key gaps in understanding the role of carrion in the landscape of disgust, which may be especially relevant in the current global context of emerging and re-emerging pathogens.

The successful reintroduction of African wild dogs (Lycaon pictus) to Gorongosa National Park, Mozambique.

Large carnivores have experienced widespread extirpation and species are now threatened globally. The ecological impact of the loss of large carnivores has been prominent in Gorongosa National Park, Mozambique, after most were extirpated during the 1977-92 civil war. To remedy this, reintroductions are now being implemented in Gorongosa, initiating with endangered African wild dogs (Lycaon pictus), hereafter 'wild dogs'. We describe the first transboundary translocation and reintroduction of founding packs of wild dogs to Gorongosa over a 28-month study period and evaluate the success of the reintroduction based on five key indicator categories. We also assess how wild dog space use and diet influenced their success. We found that pre-release, artificial pack formation in holding enclosures aided group cohesion and alpha pair establishment. Post-release, we also observed natural pack formations as a result of multiple dispersal events. Founder and naturally formed packs produced pups in two of the three breeding seasons and packs successfully recruited pups. Survival rate for all wild dogs was 73% and all mortality events were from natural causes. Consequently, the population grew significantly over the study period. All indicators of success were fully achieved and this study documents the first successful reintroduction of wild dogs into a large, unfenced landscape in Mozambique and only the second on the continent. Potential mechanisms underlying these early successes were the avoidance of habitats intensively used by lions, dietary partitioning with lion, avoidance of human settlements, and Gorongosa's management strategy. We predict further population expansion in Gorongosa given that 68% of the park is still unused by wild dogs. This expansion could be stimulated by continued reintroductions over the short- to medium-term. Recovery of wild dogs in Gorongosa could aid in the re-establishment of a larger, connected population across the greater Gorongosa-Marromeu landscape.

Beauty or beast? Farmers' dualistic views and the influence of aesthetic appreciation on tolerance towards black-backed jackal and caracal.

Various species of wild, adaptable, medium-sized carnivores occur outside of protected areas, often coming into contact with people and their domestic animals. Negative human-carnivore interactions can lead to antagonistic attitudes and behavior directed at such species. In the South African Karoo, a semi-arid rangeland, the predation of small-livestock by mesopredators is common and farmers typically use a combination of non-lethal and lethal methods to try and prevent livestock losses. We used ethnographic field observations and semi-structured interviews as part of a mixed methods approach, including the quantitative and qualitative analysis of farmers' narratives to illustrate the nuanced ways in which sheep farmers relate to the two mesopredators that consume the most livestock on their farms; black-backed jackal and caracal. Overall, farmers attributed negative characteristics to jackal and caracal but farmers' narratives provided evidence of complex perceptions in that the animals were admired as well as disliked. Both species were seen as charismatic due to traits such as their physical appearance, their "cunning" nature and their remarkable adaptability to human activities, including lethal control. Aesthetic appreciation was an important predictor of tolerance towards both species whereas negative attitudes were associated with the perception that mesopredators should only occur within protected areas. Attitudes towards jackals also appeared to have been affected by cultural representations of them as "thieves". We showed that perceiving mesopredators as beautiful increased the average marginal probability of a farmer tolerating them, and that this strong relationship held when controlling for other covariates such as livestock predation. We advocate the importance of understanding the cultural and aesthetic aspects of predators and considering existing positive dimensions of human-wildlife relationships that may encourage increased farmers' tolerance, which might promote coexistence.

Rapid evolution of the primate larynx?

Tissue vibrations in the larynx produce most sounds that comprise vocal communication in mammals. Larynx morphology is thus predicted to be a key target for selection, particularly in species with highly developed vocal communication systems. Here, we present a novel database of digitally modeled scanned larynges from 55 different mammalian species, representing a wide range of body sizes in the primate and carnivoran orders. Using phylogenetic comparative methods, we demonstrate that the primate larynx has evolved more rapidly than the carnivoran larynx, resulting in a pattern of larger size and increased deviation from expected allometry with body size. These results imply fundamental differences between primates and carnivorans in the balance of selective forces that constrain larynx size and highlight an evolutionary flexibility in primates that may help explain why we have developed complex and diverse uses of the vocal organ for communication.

The brain of the African wild dog. II. The olfactory system.

Employing a range of neuroanatomical stains, we detail the organization of the main and accessory olfactory systems of the African wild dog. The organization of both these systems follows that typically observed in mammals, but variations of interest were noted. Within the main olfactory bulb, the size of the glomeruli, at approximately 350 µm in diameter, are on the larger end of the range observed across mammals. In addition, we estimate that approximately 3,500 glomeruli are present in each main olfactory bulb. This larger main olfactory bulb glomerular size and number of glomeruli indicates that enhanced peripheral processing of a broad range of odorants is occurring in the main olfactory bulb of the African wild dog. Within the accessory olfactory bulb, the glomeruli did not appear distinct, rather forming a homogenous syncytia-like arrangement as seen in the domestic dog. In addition, the laminar organization of the deeper layers of the accessory olfactory bulb was indistinct, perhaps as a consequence of the altered architecture of the glomeruli. This arrangement of glomeruli indicates that rather than parcellating the processing of semiochemicals peripherally, these odorants may be processed in a more nuanced and combinatorial manner in the periphery, allowing for more rapid and precise behavioral responses as required in the highly social group structure observed in the African wild dog. While having a similar organization to that of other mammals, the olfactory system of the African wild dog has certain features that appear to correlate to their environmental niche.

Selection and Constraints in the Ecomorphological Adaptive Evolution of the Skull of Living Canidae (Carnivora, Mammalia).

The association between phenotype and ecology is essential for understanding the environmental drivers of morphological evolution. This is a particularly challenging task when dealing with complex traits, such as the skull, where multiple selective pressures are at play and evolution might be constrained by ontogenetic and genetic factors. I integrate morphometric tools, comparative methods, and quantitative genetics to investigate how ontogenetic constraints and selection might have interacted during the evolution of the skull in extant Canidae. The results confirm that the evolution of cranial morphology was largely adaptive and molded by changes in diet composition. While the investigation of the adaptive landscape reveals two main selective lines of least resistance (one associated with size and one associated with functional shape features), rates of evolution along size were higher than those found for shape dimensions, suggesting the influence of constraints on morphological evolution. Structural modeling analyses revealed that size, which is the line of most genetic/phenotypic variation, might have acted as a constraint, negatively impacting dietary evolution. Constraints might have been overcome in the case of selection for the consumption of large prey by associating strong selection along both size and shape directions. The results obtained here show that microevolutionary constraints may have played a role in shaping macroevolutionary patterns of morphological evolution.

Functional morphology of the jaw adductor muscles in the Canidae.

Cranial form is closely allied to diet and feeding behavior in the Canidae, with the force and velocity of jaw-closing depending on the bony morphology of the skull and mandible, and the mass, architecture, and siting of the jaw adductor muscles. Previously, little has been reported on the details of the form and function of canid jaw adductor muscles, with earlier studies basing functional hypotheses on data derived from dry skull specimens. Here we use empirically derived muscle data from fresh-frozen specimens to explore the architecture of the muscles, and to inform finite element analyses models that predict bite force and strain energy in 12 species of wild canid. The inclusion of muscle architectural detail is shown to influence masticatory muscle force production capability calculations, indicating that muscles with longer fascicles were disadvantaged compared to muscles with shorter fascicles. No clear patterns of allometry were detected. Dietary groups were differentiated by temporalis fascicle angles, which, when allied with the differentiation of rostral length reported in previous studies, may further contribute to specializations of fast jaw-closing or forceful jaw-closing species. The most biomechanically demanding masticatory function is canine biting, and the highest strain energy values were reported in this loading condition, particularly in the zygomatic arches and caudal rostrum. Specific head shapes may be constrained by size, with scaled strain energy models predicting that some bony morphologies may only be viable in species with small body masses.

ECHOCARDIOGRAPHIC VARIABLES IN THE AFRICAN PAINTED DOG (LYCAON PICTUS).

The aim of this study was to screen for cardiac disease and report normal echocardiographic variables from a managed population of the African painted dog (Lycaon pictus; APD). The study included six male adult APDs under managed care with no evidence of significant structural cardiac disease. Echocardiography was performed with APD maintained under anesthesia with a combination of midazolam, butorphanol, and inhaled isoflurane. All animals underwent a full physical examination and two-dimensional (2D), M-mode, and Doppler echocardiography using standard views. Left ventricular volumes were calculated using the Simpson's method of discs. Medians with ranges were reported for the echocardiographic variables. No valvular structural abnormalities were noted. Trace mitral regurgitation was documented in five of six APD, trace tricuspid regurgitation in three of six, and mild pulmonic regurgitation in all APD. No aortic regurgitation was noted. Standard M-mode measurements fell within the normal ranges for allometrically scaled M-mode cardiac measurements for normal adult domestic dogs (Canis familiaris). Two-dimensional measurements were comparable with that of domestic dogs with similar body weight. The study demonstrates a similarity in cardiac size and function compared with normal adult domestic dogs and provides baseline echocardiographic variables for investigation of cardiac health in other populations of APDs in the future.

Carnivores, competition and genetic connectivity in the Anthropocene.

Current extinction rates are comparable to five prior mass extinctions in the earth's history, and are strongly affected by human activities that have modified more than half of the earth's terrestrial surface. Increasing human activity restricts animal movements and isolates formerly connected populations, a particular concern for the conservation of large carnivores, but no prior research has used high throughput sequencing in a standardized manner to examine genetic connectivity for multiple species of large carnivores and multiple ecosystems. Here, we used RAD SNP genotypes to test for differences in connectivity between multiple ecosystems for African wild dogs (Lycaon pictus) and lions (Panthera leo), and to test correlations between genetic distance, geographic distance and landscape resistance due to human activity. We found weaker connectivity, a stronger correlation between genetic distance and geographic distance, and a stronger correlation between genetic distance and landscape resistance for lions than for wild dogs, and propose a new hypothesis that adaptations to interspecific competition may help to explain differences in vulnerability to isolation by humans.

Spatial Subchondral Bone Density Reflecting Joint Loading of the Talus in Different Canidae.

OBJECTIVE: Subchondral bone density distribution can be used to study joint biomechanics non-invasively. Differences in joint loading between related species can aid in the understanding of joint loading and the development of certain types of orthopaedic pathology. This study was conducted to evaluate density distribution in the subchondral bone of the talus of different Canidae species, as a parameter reflecting the long-term joint loading in the tarsocrural joint. MATERIALS AND METHODS: The tarsal joints of cadaveric dogs of different breeds were included, that is, German Shepherd (n = 5), Bouvier des Flandres (n = 3) and Labrador Retriever (n = 6).Additionally, golden jackals (n = 5) (Canis aureus) and wolves (n = 5) (Canis lupus) were included. Consecutive computed tomography slices were made and the subchondral bone density distribution was evaluated using computer tomographic osteoabsorptiometry. Different breeds and species were visually compared. RESULTS: Differences were found in the subchondral bone density distribution of the talus between breeds and between species (Canis familiaris, Canis lupus and Canis aureus). DISCUSSION AND CONCLUSION: Based on the density distribution, there are differences in loading conditions of the tarsocrural joint in different species of Canidae. The joint loading distribution is very similar between dogs of the same breed and within the same species. Although between-breed differences can be explained by conformational differences, the between-species differences remain subject to further research.

Dog appeasing pheromone prevents the androgen surge and may reduce contact dominance and active submission after stressful interventions in African wild dogs (Lycaon pictus).

The endangered African wild dog (AWD; Lycaon pictus) is a highly social canid living in packs with a separate male and female hierarchy. Immobilisation, handling and translocations are acute stressors for AWDs, however such interventions are often needed for species management. In addition, new pack formation or temporary pack separation can lead to an increase in intra-pack aggression. The goal of this double-blinded placebo-controlled study conducted in captive zoo populations was to evaluate whether dog appeasing pheromone (DAP) reduces behavioural stress and faecal glucocorticoid metabolite levels (fGCM) normally associated with pack separation, immobilisation and reintroduction (SIR), and to assess whether this reduces aggressive behaviours and faecal androgen metabolite levels (fAM). Four packs (n = 11 males) were treated with DAP and 4 packs (n = 12 males) were treated with a placebo solution, applied at the end of anaesthesia. Behavioural interactions as well as fGCM and fAM were determined from 3 days before until 4-6 days after SIR. No effect of DAP on fGCM was observed, however, fAM increased after SIR in placebo but not DAP treated animals. Moreover, on the day of reintroduction, DAP treated packs tended to have lower rates of contact-dominance and active-submission behaviour, but higher rates of non-contact dominance behaviour. As these effects could decrease the risk of agonistic interactions, DAP may be a useful tool to help manage new pack formations and temporary pack separation.

Landscape correlates of space use in the critically endangered African wild dog Lycaon pictus.

Human-carnivore conflict can threaten human life and livelihoods, leading to retaliation that negatively affects carnivore conservation. The endangered African wild dog Lycaon pictus is prone to human-carnivore conflict. Therefore, it is imperative to understand which landscape features are associated with African wild dog occurrence since selection or avoidance of these features could predict the levels of conflict. We investigated resource selection in the African wild dog in relation to four anthropogenic landscape features (livestock density, agriculture, roads and human land use) within the landscape that may pose a mortality risk, as well as one natural feature (nature reserves). We compared spatio-temporal space use patterns of four African wild dog packs in north-eastern South Africa. Data were collected from one collared individual per pack. These packs constituted approximately 10% of the total remaining African wild dog population in South Africa. Two packs occurred outside of the Kruger National Park and had access to multiple areas with farmland and other anthropogenic features, whereas the remaining two packs mainly occurred within the boundaries of the Kruger National Park but made occasional forays outside of park boundaries. Utilising Resource Selection Functions and GIS analyses, we found that agricultural landscape features, roads and nature reserves were important predictors of African wild dog occurrence for all four packs. In addition to potential conflict with farmers, high odds of occurrence on roads with fast-moving traffic and road mortality was highlighted as a concern for three of the packs. While farms and areas that house livestock were readily available, pack presence occurred in areas with few farms and low livestock densities, pointing to avoidance of areas where human-carnivore conflict and resulting mortality could occur. Our study highlights potential threats to the persistence of the African wild dog, which can be used to inform future conservation efforts of the species.

Evidence of individual discrimination in the maned wolf long-distance extended-bark.

The acoustic channel is an efficient long-distance signalling system that may be especially effective for animals moving in the dark in a vast home range. The maned wolf's extended-bark is a long-range vocalization that functions as a mechanism to increase spatial distance among conspecifics as well as to enable pair-mate reunion. Individual variations in this vocalization have been reported, but the possibility that they can be perceived and used by the species has never been tested. In our study, we used ABAB playback experiments to test if captive maned wolves could perceive individual variations. We ran 19 playback sessions with ten different subjects from six different zoos. After discarding nine sessions that did not fulfil minimal experimental conditions, in all except two of the ten valid trials, the subjects responded with displaying Oriented Attention. To our knowledge, this is the first demonstration that the maned wolf is capable of discriminating among extended-barks of different individuals.

High accuracy at low frequency: detailed behavioural classification from accelerometer data.

Accelerometers are a valuable tool for studying animal behaviour and physiology where direct observation is unfeasible. However, giving biological meaning to multivariate acceleration data is challenging. Here, we describe a method that reliably classifies a large number of behaviours using tri-axial accelerometer data collected at the low sampling frequency of 1 Hz, using the dingo (Canis dingo) as an example. We used out-of-sample validation to compare the predictive performance of four commonly used classification models (random forest, k-nearest neighbour, support vector machine, and naïve Bayes). We tested the importance of predictor variable selection and moving window size for the classification of each behaviour and overall model performance. Random forests produced the highest out-of-sample classification accuracy, with our best-performing model predicting 14 behaviours with a mean accuracy of 87%. We also investigated the relationship between overall dynamic body acceleration (ODBA) and the activity level of each behaviour, given the increasing use of ODBA in ecophysiology as a proxy for energy expenditure. ODBA values for our four 'high activity' behaviours were significantly greater than all other behaviours, with an overall positive trend between ODBA and intensity of movement. We show that a random forest model of relatively low complexity can mitigate some major challenges associated with establishing meaningful ecological conclusions from acceleration data. Our approach has broad applicability to free-ranging terrestrial quadrupeds of comparable size. Our use of a low sampling frequency shows potential for deploying accelerometers over extended time periods, enabling the capture of invaluable behavioural and physiological data across different ontogenies.

Scaling of the corpus callosum in wild and domestic canids: Insights into the domesticated brain.

All domesticated mammals exhibit marked reductions in overall brain size, however, it is unknown whether the corpus callosum (CC), an integral white matter fiber pathway for interhemispheric cortical communication, is affected by domestication differentially or strictly in coordination with changes in brain size. To answer this question, we used quantitative magnetic resonance imaging to compare the midsagittal cross-sectional areas of the CC in 35 carnivore species, including eight wild canids and 13 domestic dogs. We segmented rostro-caudal regions of interest for the CC and evaluated correlations with brain mass. The results of this study indicate that under the influence of domestication in canids, the CC scales to brain size in an allometric relationship that is similar to that of wild canids and other carnivores, with relatively high correlation coefficients observed for all regions, except the rostrum. These results indicate that architectural and energetic considerations are likely to tightly constrain variation in caudal components of the CC relative to overall brain size, however fibers passing through the rostrum, putatively connecting prefrontal cortex, are less constrained and therefore may contribute more toward species-specific differences in connectivity. Given the species diversity of the Canidae and the resurgence of interest in the brain of the domestic dog, further studies aimed at characterizing the neural architecture in domesticated species is likely to provide new insights into the effects of domestication, or artificial selection, on the brain.

Linking trophic cascades to changes in desert dune geomorphology using high-resolution drone data.

Vegetation cover is fundamental in the formation and maintenance of geomorphological features in dune systems. In arid Australia, increased woody shrub cover has been linked to removal of the apex predator (Dingoes, Canis dingo) via subsequent trophic cascades. We ask whether this increase in shrubs can be linked to altered physical characteristics of the dunes. We used drone-based remote sensing to measure shrub density and construct three-dimensional models of dune morphology. Dunes had significantly different physical characteristics either side of the 'dingo-proof fence', inside which dingoes are systematically eradicated and shrub density is higher over vast spatial extents. Generalized additive models revealed that dunes with increased shrub density were higher, differently shaped and more variable in height profile. We propose that low shrub density induces aeolian and sedimentary processes that result in greater surface erosion and sediment transport, whereas high shrub density promotes dune stability. We speculate that increased vegetation cover acts to push dunes towards an alternate stable state, where climatic variation no longer has a significant effect on their morphodynamic state within the bi-stable state model. Our study provides evidence that anthropogenically induced trophic cascades can indirectly lead to large-scale changes in landscape geomorphology.

GONIOMETRIC MEASUREMENT OF JOINT ANGLES IN CRAB-EATING FOXES ( CERDOCYON THOUS).

This study aimed to assess joint angle values in the forelimbs and hind limbs of crab-eating foxes ( Cerdocyon thous) using a conventional manual goniometer. Nine captive crab-eating foxes, aged over 1 yr old, weighing from 5.9 to 7.5 kg were used. The maximum flexion and maximum extension of the shoulder, elbow, carpal, hip, stifle, and tarsal joints were measured. The range of motion was determined through the difference between the maximum flexion and the maximum extension. There was no significant difference between the right and left sides in the goniometric measurements. The mean values of the range of motion for shoulder, elbow, carpal, hip, stifle, and tarsal joints were respectively 108°, 99°, 131°, 79°, 103°, and 86°. The goniometric values of the crab-eating foxes can be used as means of comparison with other animals from the same species for diagnostic purposes, research documentation, and evaluation of therapeutic procedures.