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Climate oscillations and prehistoric human activity during the Middle-Late Pleistocene profoundly affected the population fluctuations and extinctions of megafauna, especially the extinct woolly rhinoceros. Fordham et al. (2024) recently proposed new solutions based on fossil records, paleoclimates, and prehistoric human activities data to reconstruct an explicit process-driven model, resulting in high-resolution population dynamics of the woolly rhinoceros. This study revealed the mechanisms of the woolly rhinoceros extinction: climate-driven habitat fragmentation combined with low but persistent levels of human hunting weakened metapopulation processes, leading to their extinction., (© 2024 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

In mammals, the patella is the biggest sesamoid bone of the skeleton and is of crucial importance in posture and locomotion, ensuring the role of a pulley for leg extensors while protecting and stabilizing the knee joint. Despite its central biomechanical role, the relation between the shape of the patella and functional factors, such as body mass or locomotor habit, in the light of evolutionary legacy are poorly known. Here, we propose a morphofunctional investigation of the shape variation of the patella among modern rhinoceroses and more generally among perissodactyls, this order of ungulates displaying a broad range of body plan, body mass and locomotor habits, to understand how the shape of this sesamoid bone varies between species and relatively to these functional factors. Our investigation, relying on three dimensional geometric morphometrics and comparative analyses, reveals that, within Rhinocerotidae and between the three perissodactyl families, the shape of the patella strongly follows the phylogenetic affinities rather than variations in body mass. The patellar shape is more conservative than initially expected both within and between rhinoceroses, equids and tapirs. The development of a medial angle, engendering a strong mediolateral asymmetry of the patella, appears convergent in rhinoceroses and equids, while tapirs retain a symmetric bone close to the plesiomorphic condition of the order. This asymmetric patella is likely associated with the presence of a "knee locking" mechanism in both equids and rhinos. The emergence of this condition may be related to a shared locomotor habit (transverse gallop) in both groups. Our investigation underlines unexcepted evolutionary constraints on the shape of a sesamoid bone usually considered as mostly driven by functional factors., Competing Interests: The authors declare that they have no competing interests., (© 2024 Mallet and Houssaye.)

Artificial water provisioning is a common practice in southern African nature reserves, where different game species exhibit preferences for specific waterhole types. The movement patterns and behaviour of elephants and rhinos are closely linked to water availability, with these mega-herbivores noticeably influencing the environment and other species they interact with at waterholes. Since there is limited research on this topic, understanding preferences for different types of artificial waterholes is crucial, particularly during periods of water scarcity. This knowledge enables reserve managers to effectively manage the numbers and types of waterholes. In this study, we investigate artificial waterhole selection and preferences by elephants and rhinos in the Olifants West Nature Reserve, South Africa. The study area featured various waterhole types, including earth dams, concrete pans, reservoirs, and troughs. By employing camera traps, we analysed visitation frequency, timing of visits, and factors influencing visit frequency. Our findings revealed distinct preferences for specific waterhole types among different social groupings of the study species. Breeding herds of elephants predominantly utilized reservoirs and occasionally visited troughs, while bachelor herds favoured earth dams. Black rhinos showed a preference for earth dams, whereas white rhinos selected troughs and earth dams, with bachelor groups favouring troughs and female rhinos favouring earth dams. The outcomes of this study have significant implications for the development of comprehensive conservation plans in areas where these species reside, and for potential release sites., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Smith et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Heavy animals incur large forces on their limb bones, due to the transmission of body weight and ground reaction forces, and the contractions of the various muscles of the limbs. This is particularly true for rhinoceroses, the heaviest extant animals capable of galloping. Several studies have examined their musculoskeletal system and the forces their bones incur, but no detailed quantification has ever been attempted. Such quantification could help understand better the link between form and function in giant land animals. Here we constructed three-dimensional musculoskeletal models of the forelimb and hindlimb of Ceratotherium simum, the heaviest extant rhino species, and used static optimisation (inverse) simulations to estimate the forces applied on the bones when standing at rest, including magnitudes and directions. Overall, unsurprisingly, the most active muscles were antigravity muscles, which generate moments opposing body weight (thereby incurring the ground reaction force), and thus keep the joints extended, avoiding joint collapse via flexion. Some muscles have an antigravity action around several joints, and thus were found to be highly active, likely specialised in body weight support (ulnaris lateralis; digital flexors). The humerus was subjected to the greatest amount of forces in terms of total magnitude; forces on the humerus furthermore came from a great variety of directions. The radius was mainly subject to high-magnitude compressive joint reaction forces, but to little muscular tension, whereas the opposite pattern was observed for the ulna. The femur had a pattern similar to that of the humerus, and the tibia's pattern was intermediate, being subject to great compression in its caudal side but to great tension in its cranial side (i.e. bending). The fibula was subject to by far the lowest force magnitude. Overall, the forces estimated were consistent with the documented morphofunctional adaptations of C. simum's long bones, which have larger insertion areas for several muscles and a greater robusticity overall than those of lighter rhinos, likely reflecting the intense forces we estimated here. Our estimates of muscle and bone (joint) loading regimes for this giant tetrapod improve the understanding of the links between form and function in supportive tissues and could be extended to other aspects of bone morphology, such as microanatomy., (© 2024 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

We examined the entanglement of biodiversity conservation, human-animal interactions, zootherapy, and local beliefs among Sumatran Healers and their local community by completing an ethnography of 43 Indigenous Healers across 8 tribes in Bengkulu Province, Sumatra, Indonesia. Data collection tools were interviews, observations, videos, photographs, and a researcher journal. Of the 43 Healers, 30 used animals and mentioned 62 species. Of the animals identified, the International Union for Conservation of Nature Red List lists 34% (n = 21) as endangered, decreasing, or vulnerable, including Sumatran tiger (Panthera tigris sumatrae), Sumatran elephant (Elephas maximus sumatranus), and Sumatran rhinoceros (Dicerorhinus sumatrensis). Of the 30 Healers using animals, 50% (n = 15) practiced healing with at least one endangered, decreasing, or vulnerable animal. We defined 3 personas: Healer self-persona, Healer-imposed persona, and community-imposed persona. A persona represented a group's opinions and sentiments related to Healers killing animals for medicinal purposes. Using an iterative data analysis process, we grouped the data across the 3 personas into 5 themes: ease of killing and preparing animals, emotions related to killing animals, animal value, relationship to religion, and Healers are tricksters. The complexity of merging the identities of Healers and the community within an actor-network embodies the relationality of actions, interactions, and feelings among Healers, between Healers and animals, and between Healers and the community. Conservationists should be cognizant of Healers' medicinal use of animals, views of human-animal interactions, and zootherapy from all social and emotional perspectives. The data led to defining Indigenous Healer ecological knowledge components of zootherapy, human-animal interactions, and biodiversity conservation., (© 2024 Society for Conservation Biology.)

Although black rhinoceros Diceros bicornis are mostly solitary in the wild, the Hiroshima City Asa Zoological Park (Asa Zoo) has kept a family group together during the daytime, with good reproductive performance over five decades. Management procedures at the zoo include temporary single housing of the mother before and after giving birth, which facilitates maintenance of a compatible family group. We recorded intra-group spatial relationships for 4 years and 4 months, during which time an adult female reared two consecutive calves. During daytime she remained in an enclosure with her new calf, one or two older offspring, and an adult male, the sire of all her offspring. Proximity (within two adult body-lengths) scores between the mother and her two calves were especially high during the first year after birth, and only slightly lower for her older offspring. The adult male had the lowest proximity scores. The spatial relationships were visualized by applying multidimensional scaling (MDS) to the proximity scores. Mother and calves were plotted close to each other, with older offspring slightly farther apart on the two-dimensional MDS representation; the adult male was more distant from the other group members. These findings indicate clear follower-type characteristics in the mother-calf pair and also older immature offspring, albeit to a lesser degree. Although black rhinoceros are generally solitary in the wild, our results duplicate observations of some wild black rhinoceros groups containing an adult female, her calf, and an older immature, with adult males being largely solitary., (© 2024 Wiley Periodicals LLC.)

Bite force is often associated with specific morphological features, such as sagittal crests. The presence of a pronounced sagittal crest in some tapirs (Perissodactyla: Tapiridae) was recently shown to be negatively correlated with hard-object feeding, in contrast with similar cranial structures in carnivorans. The aim of this study was to investigate bite forces and sagittal crest heights across a wide range of modern and extinct tapirs and apply a comparative investigation to establish whether these features are correlated across a broad phylogenetic scope. We examined a sample of 71 specimens representing 15 tapir species (5 extant, 10 extinct) using the dry-skull method, linear measurements of cranial features, phylogenetic reconstruction, and comparative analyses. Tapirs were found to exhibit variation in bite force and sagittal crest height across their phylogeny and between different biogeographical realms, with high-crested morphologies occurring mostly in Neotropical species. The highest bite forces within tapirs appear to be driven by estimates for the masseter-pterygoid muscle complex, rather than predicted forces for the temporalis muscle. Our results demonstrate that relative sagittal crest height is poorly correlated with relative cranial bite force, suggesting high force application is not a driver for pronounced sagittal crests in this sample. The divergent biomechanical capabilities of different contemporaneous tapirids may have allowed multiple species to occupy overlapping territories and partition resources to avoid excess competition. Bite forces in tapirs peak in Pleistocene species, independent of body size, suggesting possible dietary shifts as a potential result of climatic changes during this epoch., (© 2022 International Society of Zoological Sciences, Institute of Zoology/Chinese Academy of Sciences and John Wiley & Sons Australia, Ltd.)

Objective: To determine the pharmacokinetics of a single bolus of intravenous (IV) propofol after intramuscular administration of etorphine, butorphanol, medetomidine, and azaperone in 5 southern white rhinoceros to facilitate reproductive evaluations. A specific consideration was whether propofol would facilitate timely orotracheal intubation., Animals: 5 adult, female, zoo-maintained southern white rhinoceros., Procedures: Rhinoceros were administered etorphine (0.002 mg/kg), butorphanol (0.02 to 0.026 mg/kg), medetomidine (0.023 to 0.025 mg/kg), and azaperone (0.014 to 0.017 mg/kg) intramuscularly (IM) prior to an IV dose of propofol (0.5 mg/kg). Physiologic parameters (heart rate, blood pressure, respiratory rate, and capnography), timed parameters (eg, time to initial effects and intubation), and quality of induction and intubation were recorded following drug administration. Venous blood was collected for analysis of plasma propofol concentrations using liquid chromatography-tandem mass spectrometry at various time points after propofol administration., Results: All animals were approachable following IM drug administration, and orotracheal intubation was achieved at 9.8 ± 2.0 minutes (mean ±SD) following propofol administration. The mean clearance for propofol was 14.2 ± 7.7 ml/min/kg, the mean terminal half-life was 82.4 ± 74.4 minutes, and the maximum concentration occurred at 2.8 ± 2.9 minutes. Two of 5 rhinoceros experienced apnea after propofol administration. Initial hypertension, which improved without intervention, was observed., Clinical Relevance: This study provides pharmacokinetic data and insight into the effects of propofol in rhinoceros anesthetized using etorphine, butorphanol, medetomidine, and azaperone. While apnea was observed in 2 rhinoceros, propofol administration allowed for rapid control of the airway and facilitated oxygen administration and ventilatory support.

Translocation is a valuable conservation tool, but poses significant risks for the transported rhinoceroses. Interventions reducing these risks are required to ensure positive welfare during transportation. The aim of this study was to evaluate the effect of journey duration and feeding during the transport of white rhinoceroses ( Ceratotherium simum simum ). A total of 32 animals were transported by road during two events, five days apart. Fifteen rhinoceroses in the first transport event (37.0 ± 2.4 hr duration) were not fed, while 17 rhinoceroses in the second event (32.2 ± 1.5 hr duration) were offered lucerne. Blood samples were collected at capture and after transport for the evaluation of changes in serum clinical chemistry analytes. The Wilcoxon rank-sum test was used to compare differences between the groups. In all rhinoceroses, transport resulted in changes in serum electrolyte, metabolite and enzyme concentrations, indicating a loss in total body water, nutritional shifts, stress and fatigue. Fed rhinoceroses, transported over a shorter time, displayed greater changes in osmolality ( p < 0.006), serum sodium and chloride concentrations ( p = 0.005 and = 0.001, respectively) indicating a greater degree of total body water loss than non-fed rhinoceroses. Feeding and a shorter transport duration reduced, but did not prevent, nutritional challenges. A greater increase in the muscle enzymes CK and AST ( p = 0.027 and = 0.001, respectively), indicated greater fatigue in non-fed rhinoceroses transported over a longer time. Further work to distinguish the effects of feeding and journey duration is required to better understand the role feeding may play in mitigating welfare challenges during rhinoceros translocation.

For over a century, researchers have assumed that the plane of the lateral semicircular canal of the inner ear lies parallel to the horizon when the head is at rest, and used this assumption to reconstruct head posture in extinct species. Although this hypothesis has been repeatedly questioned, it has never been tested on a large sample size and at a broad taxonomic scale in mammals. This study presents a comprehensive test of this hypothesis in over one hundred "ungulate" species. Using CT scanning and manual segmentation, the orientation of the skull was reconstructed as if the lateral semicircular canal of the bony labyrinth was aligned horizontally. This reconstructed cranial orientation was statistically compared to the actual head posture of the corresponding species using a dataset of 10,000 photographs and phylogenetic regression analysis. A statistically significant correlation between the reconstructed cranial orientation and head posture is found, although the plane of the lateral semicircular canal departs significantly from horizontal. We thus caution against the use of the lateral semicircular canal as a proxy to infer precisely the horizontal plane on dry skulls and in extinct species. Diet (browsing or grazing) and head-butting behaviour are significantly correlated to the orientation of the lateral semicircular canal, but not to the actual head posture. Head posture and the orientation of the lateral semicircular canal are both strongly correlated with phylogenetic history.

The appendicular skeleton of tetrapods is a particularly integrated structure due to the shared developmental origin or similar functional constraints exerted on its elements. Among these constraints, body mass is considered strongly to influence its integration but its effect on shape covariation has rarely been addressed in mammals, especially in heavy taxa. Here, we propose to explore the covariation patterns of the long bones in heavy animals and their link to body mass. We investigate the five modern rhinoceros species, which display an important range of bodyweight. We used a 3D geometric morphometric approach to describe the shape covariation of the six bones composing the stylopodium and zeugopodium both among and within species. Our results indicate that the appendicular skeleton of modern rhinos is a strongly integrated structure. At the interspecific level, the shape covariation is roughly similar between all pairs of bones and mainly concerns the muscular insertions related to powerful flexion and extension movements. The forelimb integration appears higher and more related to body mass than that of the hind limb, suggesting a specialization for weight support. The integration of the stylopodium elements does not seem to relate to body mass in our sample, which suggests a greater effect of shared developmental factors. Conversely, the covariation of the zeugopodium bones seems more associated with body mass, particularly for the radius-ulna pair. The fibula appears poorly integrated with other bones, especially within non-Rhinoceros species, which may represent a case of parcellation due to a functional dissociation between the hind limb bones. The exploration of the integration patterns at the intraspecific level also highlights a more prominent effect of age over individual body mass on shape covariation within C. simum. This study lends support to previous hypotheses indicating a link between high body mass and high integration level., (© 2020 Anatomical Society.)

Vocal communication networks can be linked to social behaviour, allowing a deeper understanding of social relationships among individuals. For this purpose, the description of vocal dyads is fundamental. In group-living species, this identification is based on behavioural indicators which require a high level of reactivity during social interactions. In the present study, we alternatively established a proximity-based approach to investigate whether sex-specific differences in vocal communication reflect social behaviour in a species with rather loose social associations and low levels of reactivity: the Southern white rhinoceros (Ceratotherium simum simum). We performed audio- and video recordings of 30 captive animals from seven groups. Vocal networks for the four most common call types were constructed by considering conspecifics at close distance (≤ 1 body length) to the sender as potential receivers. The analysis of the resulting unidirectional structures showed that not only the sex of the sender but also the sex of the potential receiver, the quality of social interactions (affiliative or agonistic) as well as association strength predict the intensity of vocal interactions between group members. Thus, a proximity-based approach can be used to construct vocal networks providing information about the social relationships of conspecifics-even in species with loose social associations where behavioural indicators are limited.

Feeding practice in herbivorous mammals can impact their dental wear, due to excessive or irregular abrasion. Previous studies indicated that browsing species display more wear when kept in zoos compared to natural habitats. Comparable analyses in tapirs do not exist, as their dental anatomy and chewing kinematics are assumed to prevent the use of macroscopic wear proxies such as mesowear. We aimed at describing tapir chewing, dental anatomy and wear, to develop a system allowing comparison of free-ranging and captive specimens even in the absence of known age. Video analyses suggest that in contrast to other perissodactyls, tapirs have an orthal (and no lateral) chewing movement. Analysing cheek teeth from 74 museum specimens, we quantified dental anatomy, determined the sequence of dental wear along the tooth row, and established several morphometric measures of wear. In doing so, we showcase that tapir maxillary teeth distinctively change their morphology during wear, developing a height differential between less worn buccal and more worn lingual cusps, and that quantitative wear corresponds to the eruption sequence. We demonstrate that mesowear scoring shows a stable signal during initial wear stages but results in a rather high mesowear score compared to other browsing herbivores. Zoo specimens had lesser or equal mesowear scores as specimens from the wild; additionally, for the same level of third molar wear, premolars and other molars of zoo specimens showed similar or less wear compared specimens from the wild. While this might be due to the traditional use of non-roughage diet items in zoo tapirs, these results indicate that in contrast to the situation in other browsers, excessive tooth wear appears to be no relevant concern in ex situ tapir management., Competing Interests: The authors have declared that no competing interests exist.

Paleontologists and paleoanthropologists have long debated relationships between cranial morphology and diet in a broad diversity of organisms. While the presence of larger temporalis muscle attachment area (via the presence of sagittal crests) in carnivorans is correlated with durophagy (i.e. hard-object feeding), many primates with similar morphologies consume an array of tough and hard foods-complicating dietary inferences of early hominins. We posit that tapirs, large herbivorous mammals showing variable sagittal crest development across species, are ideal models for examining correlations between textural properties of food and sagittal crest morphology. Here, we integrate dietary data, dental microwear texture analysis, and finite element analysis to clarify the functional significance of the sagittal crest in tapirs. Most notably, pronounced sagittal crests are negatively correlated with hard-object feeding in extant, and several extinct, tapirs and can actually increase stress and strain energy. Collectively, these data suggest that musculature associated with pronounced sagittal crests-and accompanied increases in muscle volume-assists with the processing of tough food items in tapirs and may yield similar benefits in other mammals including early hominins.

Evolutionary theory expects social, communicative species to eavesdrop most on other species' alarm calls [e.g., 1, 2] but also that solitary-living species benefit most from eavesdropping [3, 4]. Examples of solitary species responding to the alarm calls of other species, however, are limited and unconvincing [3-5]. The Swahili name for the red-billed oxpecker (Buphagus erythrorynchus) is Askari wa kifaru, the rhinos' guard [6]. Black rhino (Diceros bicornis) are a solitary-living, non-vocal species and are critically endangered through hunting. We searched Hluhluwe-iMfolozi Park, South Africa, for rhinoceros for 27 months with and without the aid of radio telemetry and conducted 86 experimental, unconcealed approaches to 11 rhino, without or with varying numbers of resident oxpecker. Oxpeckers enabled rhinos to evade detection by us in 40% to 50% of encounters. Alarm-calling by oxpeckers significantly improved the rate and distance that rhinos detected our approach from 23% to 100% and 27 ± 6 m to 61 ± 4 m, respectively. Every additional oxpecker improved detection distance by 9 m. Rhinos alerted by oxpeckers' alarm calls never re-oriented in our direction but moved to face downwind. Thus, oxpeckers' calls communicate only threat proximity, not direction, and rhinos assume the hunter is stalking from downwind. We confirm that oxpeckers guard rhinos and the importance of depredation, not sociality, in the evolution of eavesdropping [4, 7]. Conservationists should consider reintroducing oxpeckers to rhino populations, reinstating their anti-human sentinel [8]. VIDEO ABSTRACT., Competing Interests: Declaration of Interests The authors declare no competing interests., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Malayan tapirs are listed as endangered and are bred in captivity under governmental management. The success of captive breeding programs varies and the underlying causes are unclear. Here, we investigate how tapir reproduction is affected by previous breeding experience, enclosure type/size and visitor numbers so that appropriate steps can be taken to achieve self-sustaining captive populations. Data on social and reproductive behaviors were collected from six tapirs (three males, three females), from different breeding centers in Peninsular Malaysia for 18 weeks. Results revealed that social and reproductive behavior of both sexes was significantly influenced by social and environmental conditions. Larger enclosure size tended to increase social and reproductive behaviors, whereas high number of visitors reduced initial interaction between males and females. No specific breeding month was confirmed; however, reproductive behaviors were highest in April. Overall, this study contributes to a better understanding of the relationships between social and reproductive behaviors, and captive environments on Malayan tapirs. In future, frequency of sexual interactions should be monitored regularly to identify animals exhibiting below-average frequency and who might, therefore, be prone to reproductive difficulties.

The Pliocene site of Kanapoi is key to our understanding of the environmental context of the earliest species of Australopithecus. Various approaches have been used to reconstruct the environments of this site, and here we contribute new data and analyses using mesowear and hypsodonty. The dental traits of 98 bovids, suids and rhinocerotids from Kanapoi were analyzed using these proxies. Results indicate that most of the animals analyzed had a relatively abrasive diet. Bovids in the assemblage incorporated more grass into their diet than do modern species of the same tribe or genus. Although Pliocene Kanapoi likely had complex environments, our analysis indicates that grassy habitats were a dominant component of the ecosystem, a conclusion that supports the results of previous investigations of the paleoecology of the site., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

All extant species in the Rhinocerotidae family are experiencing escalating threats in the wild, making self-sustaining captive populations essential genetic reservoirs for species survival. Assisted reproductive technologies (ARTs) will become increasingly important for achieving and maintaining ex situ population sustainability and genetic diversity. Previous reports have shown that a large proportion of captive southern white rhinoceros (SWR) females are irregularly cyclic or acyclic, and that cycling females display two different estrous cycle lengths of approximately 30 or 70 days. It has been suggested that the longer estrous cycle length is infertile or subfertile, as no term pregnancies have been observed following long cycles. Here we report the achievement of two pregnancies following long luteal phases, using ovulation induction and artificial insemination with either fresh or frozen-thawed semen. One female SWR conceived on the first insemination attempt and gave birth to a live offspring. A second female conceived twice in consecutive long cycles although the first embryo was resorbed by 33 days post-insemination. A pregnancy from this female's second insemination is ongoing with expected parturition in November 2019. Whether prolonged estrous cycles in SWR are subfertile or infertile in natural breeding situations remains unclear. However, our findings demonstrate that the application of ARTs following prolonged cycles can result the successful establishment of pregnancies in SWR. Therefore, with ARTs, female SWR otherwise considered nonreproductive due to long estrous cycles may still have the potential for representation and contribution to the ex situ population., (© 2019 The Authors. Zoo Biology Published by Wiley Periodicals, Inc.)

The volume fraction (VF) of a given brain region, or the proper mass, ought to reflect the importance of that region in the life of a given species. This study sought to examine the VF of various brain regions across 61 different species of mammals to discern if there were regularities or differences among mammalian orders. We examined the brains of carnivores (n = 17), ungulates (n = 8), rodents (n = 7), primates (n = 11), and other mammals (n = 18) from the online collections at the National Museum of Health and Medicine. We measured and obtained the VF of several brain regions: the striatum, thalamus, neocortex, cerebellum, hippocampus, and piriform area. We refined our analyses by using phylogenetic size correction, yielding the corrected (c)VF. Our groups showed marked differences in gross brain architecture. Primates and carnivores were divergent in some measures, particularly the cVF of the striatum, even though their overall brain size range was roughly the same. Rodents predictably had relatively large cVFs of subcortical structures due to the fact that their neocortical cVF was smaller, particularly when compared to primates. Not so predictably, rodents had the largest cerebellar cVF, and there were marked discrepancies in cerebellar data across groups. Ungulates had a larger piriform area than primates, perhaps due to their olfactory processing abilities. We provide interpretations of our results in the light of the comparative behavioral and neuroanatomical literature., (© 2020 S. Karger AG, Basel.)

In vivo and ex vivo sensors have the potential to aid tracking and anti-poaching endeavours and provide new insights into rhinoceros physiology and environment. However, the propagation of electromagnetic signals in rhinoceros tissue is currently not known. We present simulation and agar models of the rhinoceros that allow the investigation of electromagnetic propagation by in vivo and ex vivo devices without the need for surgery. Since the dielectric properties of rhinoceros tissue have not been documented, the conductivity and permittivity of the skin, fat, muscle, blood and other organs are first approximated by means of a meta-analysis that includes animals with similar physical properties. Subsequently, we develop anatomical models that include dermal layers, internal organs and a skeleton. We also develop a flank model that serves as an approximation of the anatomical model in certain situations. These models are used to determine the viability of communication between an in vivo device and an ex vivo device attached to the hind leg of the animal. Two types of antenna (microstrip-fed planar elliptical monopole antenna and printed inverted-F antenna) and three feasible implant locations (back, neck and chest) are considered. In addition to the computer models, phantom recipes using salt, sugar and agar are developed to match the dielectric properties of each tissue type at the industrial, scientific and medical (ISM) frequencies of 403MHz, 910MHz and 2.4GHz. The average error between the measured and theoretically predicted dielectric values was 6.22% over all recipes and 4.49% for the 2.4 GHz recipe specifically. When considering the predicted efficiency of the transmitting and receiving antennas, an agreement of 67.38% was demonstrated between the computer simulations and laboratory measurements using the agar rhinoceros flank models. Computer simulations using the anatomical model of the rhinoceros indicate that the chest is the optimal implant location and that best signal propagation is achieved using the planar inverted-F antenna (PIFA). Using this configuration, the simulations indicate that communication between the implant and an ex vivo device attached to the hind leg is challenging but possible. Furthermore, we find that the inclusion of factors such as the density and temperature of the phantom materials were found to be critical to the achievement of good agreement between practice and simulation., Competing Interests: The authors have declared that no competing interests exist.

A three-species population system under a facultative mutualistic relationship of one of the species is studied. The considered interactions are as follows: facultative between the first species and the second species, obligatory mutualism between the second species and the first one, and the third species is a predator of the first species. For this purpose, we extend the model proposed by Morozov et al., originally used to describe obligatory mutualism, to consider obligatory and facultative mutualism and prove that under adequately selected parameters this system produces a spatial patchy spread of populations or continuous wave fronts. Since the analytical treatment of a three-species model is often prohibitive, we first analyze the interaction between two mutualist species without diffusion and without the presence of the predator. Some parameters are fixed in the bistable regime of the mutualistic species to further consider the influence of the third species. The remaining parameters are then selected to produce patchy patterns under different mortality rates. Finally, the equations of the final three-species system are numerically solved to test the influence of different initial conditions in the formation of patchy populations. It is confirmed that the velocity and the profile of a traveling front are independent on the initial conditions. Our approach opens the way to study more general biological situations., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Background: Host microbiomes play a role in hormone production and subsequent fertility in humans, but this is less well understood in non-model organisms. This is of particular relevance to species in zoo-based conservation breeding programmes, as relationships between host microbiome composition and reproductive output may allow for the development of microbial augmentation strategies to improve success. Here, we characterise faecal bacterial communities of breeding and non-breeding eastern black rhino (Diceros bicornis michaeli) using 16S rRNA gene amplicon sequencing and quantify progestagen and glucocorticoid metabolite concentrations through enzyme immunoassays to identify such relationships., Results: We identified significant differences in black rhino gut microbiome composition according to ID, institution, breeding success and ovarian cycle phase. In particular, the gut microbiome during pregnancy and post-parturition was significantly altered. Around a third of bacterial genera showed more than ± 10% correlation with either progestagen and/or glucocorticoid concentration, and in general, microbial genera correlated with both hormones in the same direction. Through a combination of analyses, we identified four genera (Aerococcaceae, Atopostipes, Carnobacteriaceae and Solobacterium) that were significantly associated with breeding success, pregnancy and/or post-parturition, and higher faecal progestagen metabolite concentrations. These genera had a lower-than-average relative abundance in the gut microbiome., Conclusion: Our results indicate that many members of the gut microbiome of black rhino are associated with hormone production and breeding success, and some members of the rare microbiota appear to be particularly important. Although the directionality of the relationship is unclear, the variation in gut microbiome communities represents a potential biomarker of reproductive health. We identified four genera that were associated with multiple indicators of reproductive output; these could be candidate probiotics to improve the breeding success of black rhino in zoo-based conservation breeding programmes. Further work is required to understand the efficacy and feasibility of this, either directly through microbial augmentation (e.g. probiotics) or indirectly via dietary manipulation or prebiotics.

Unrelenting poaching to feed the illegal trafficking of rhinoceros (rhino) horn remains the principle threat to the persistence of south-central black and southern white rhino that live in the Kruger National Park (Kruger), South Africa. Other global environmental change drivers, such as unpredictable climatic conditions, impose additional uncertainties on the management and persistence of these species. The drought experienced in Kruger over the 2015/2016 rainy season may have affected rhino population growth and thus added an additional population pressure to the poaching pressure already occurring. Under drought conditions, reduced grass biomass predicts increased natural deaths and a subsequent decrease in birth rate for the grazing white rhino. Such variance in natural death and birth rates for the browsing black rhino are not expected under these conditions. We evaluated these predictions using rhino population survey data from 2013 to 2017. Comparisons of natural deaths and birth rates between pre- (2013/2014 and 2014/15), during- (2015/2016) and post-drought (2016/2017) periods in Kruger showed increased natural mortality and decreased births for white rhino, but no significant changes for black rhino, supporting our predictions. As a result, despite reduced poaching rates, the total mortality rate of white rhino remains significantly higher than the birth rate. Decreased poaching, decreased natural deaths and no apparent drought effects in black rhino resulted in a lower total mortality rate than the estimated birth rate in 2017. Active biological management and traditional anti-poaching initiatives together therefore represent the most likely way to buffer the impacts of decreased population growth through climate change and wildlife crime on the persistence of rhinos., Competing Interests: The authors have declared that no competing interests exist.

This report describes weekly repeated anesthesia in a 7-yr-old, 1,030 kg, female Eastern black rhinoceros ( Diceros bicornis michaeli), that was immobilized six times using a combination of 2 mg etorphine (0.002 mg/kg), 5 mg medetomidine (0.005 mg/kg), 25 mg midazolam (0.024 mg/kg), and 300 mg ketamine (0.29 mg/kg) delivered intramuscularly (IM) via remote dart to facilitate long-term medical care of a bilateral, obstructive Actinomyces sp. rhinitis. The drug combination described in this study resulted in reliable, rapid recumbency of the animal within 2-8 min after initial administration via dart and produced deep anesthesia for 34-78 min without supplemental anesthetic administration. Antagonist drugs (100 mg naltrexone [0.1 mg/kg] and 25 mg atipamezole [0.024 mg/kg] IM) produced reliable and uneventful recoveries in all the procedures. During each anesthetic procedure, the animal was intubated and provided intermittent positive pressure ventilation with a megavertebrate demand ventilator. Tachycardia and hypoxia noted after induction resolved after positive pressure ventilation with oxygen. This report provides useful information on a novel anesthetic protocol used repeatedly for intensive medical management in a black rhinoceros.

The effects of etorphine on the pulmonary vascular system of white rhinoceros ( Ceratotherium simum) have not been described and could play a role in the severe hypoxemia that develops after immobilization with etorphine-based drug combinations. Characterization of these effects requires measurement of pulmonary vascular pressures and cardiac output (CO). To refine a technique for pulmonary arterial catheterization, five boma-habituated white rhinoceros (three females and two males weighing 1,012-1,572 kg) were immobilized by remote injection with etorphine plus azaperone followed by butorphanol. This afforded the opportunity to perform a pilot study and acquire preliminary measurements of pulmonary arterial pressure (PAP) and CO before and after supplemental etorphine given intravenously. Ultrasonographic guidance was used to insert a sheath introducer into a linguofacial branch of a jugular vein. A 160-cm-long pulmonary artery catheter with a balloon and thermistor was then passed through the introducer and positioned with its tip in the pulmonary artery. It was not long enough to permit wedging for measurement of pulmonary artery occlusion pressure. Mean PAP was 35 mm Hg (minimum, maximum 32, 47 mm Hg) and increased ( P = 0.031) by 83% (28, 106%) after supplemental etorphine. Thermodilution CO was 120 L/min (92, 145 L/min) and increased 27% (3, 43%) ( P = 0.031). Heart rate was 100 (88, 112) beats/min and increased 20% (4, 45%) ( P = 0.031), whereas arterial partial pressure of oxygen was 35 mm Hg (30, 94 mm Hg) and decreased 47% (20, 72%) ( P = 0.031). The cardiovascular observations could result from etorphine-induced generalized sympathetic outflow, as has been reported in horses. Further studies of etorphine in isolation are needed to test this suggestion and to discern how the changes in pulmonary vascular pressures and blood flow might relate to hypoxemia in etorphine-immobilized white rhinoceros.

In the present study, a quantitative proteomic approach was used to analyze and compare the proteome in horns from endangered species (rhinoceros, Saiga antelope, and Tibetan antelope) and common species (yak, water buffalo, and goat) based on the isobaric tag for relative and absolute quantification techniques. In total, 591 proteins were identified, and 321 were quantified and categorized based on molecular function, cellular component, and biological process. Principal component analysis and hierarchical clustering analysis results based on differences in the amount of protein identified three major clusters, and proteins including transglutaminase, desmocollin, and elongation factors were selected as trait components from proteomic patterns of horn samples from different species. Quantitative proteomic analysis based strategies can therefore provide further evidence for sustainable alternatives to replace animal horn from threatened species., (© 2018 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim.)

The white rhinoceros ( Ceratotherium simum ) has a discontinuous African distribution, which is limited by the extent of sub-Saharan grasslands. The southern population (SWR) declined to its lowest number around the turn of the nineteenth century, but recovered to become the world's most numerous rhinoceros. In contrast, the northern population (NWR) was common during much of the twentieth century, declining rapidly since the 1970s, and now only two post-reproductive individuals remain. Despite this species's conservation status, it lacks a genetic assessment of its demographic history. We therefore sampled 232 individuals from extant and museum sources and analysed ten microsatellite loci and the mtDNA control region. Both marker types reliably partitioned the species into SWR and NWR, with moderate nuclear genetic diversity and only three mtDNA haplotypes for the species, including historical samples. We detected ancient interglacial demographic declines in both populations. Both populations may also have been affected by recent declines associated with the colonial expansion for the SWR, and with the much earlier Bantu migrations for the NWR. Finally, we detected post-divergence secondary contact between NWR and SWR, possibly occurring as recently as the last glacial maximum. These results suggest the species was subjected to regular periods of fragmentation and low genetic diversity, which may have been replenished upon secondary contact during glacial periods. The species's current situation thus reflects prehistoric declines that were exacerbated by anthropogenic pressure associated with the rise of late Holocene technological advancement in Africa. Importantly, secondary contact suggests a potentially positive outcome for a hybrid rescue conservation strategy, although further genome-wide data are desirable to corroborate these results., (© 2018 The Author(s).)

Many mammals enhance their olfactory signals visually by depositing them in conspicuous locations such as well-travelled paths. It is also possible to enhance the odour itself through behaviours aimed at modifying odour emission rates. White rhinos defecate in communal middens. While defecating, territorial males kick sharply with their back feet which disperses their dung. Despite being a ubiquitous trait of territorial male white rhinos, the reason behind this behaviour is unclear. We hypothesised that the purpose of dung kicking was for olfactory signal amplification (OSA) in terms of an increased emission of volatile compounds (i.e. increased signal strength). Using dung collected from non-territorial adult males (because it is not possible to collect whole dung from territorial males), we show that the dispersal of male white rhino dung causes OSA by increasing the emission of hydrocarbon acids. The dung odour of territorial and non-territorial males differs only quantitatively, hence it is likely that the same emission patterns occur for territorial male dung odours following dung dispersal. The volatile compound indicating age of intact dung was toluene, but for dispersed dung it was acetophenone (similar to territorial male dung). Despite the benefits of OSA, dung dispersal carried a cost of decreased odour longevity. Thus, signal detectability is temporally reduced. However, territorial males likely counteract this by defecating in middens both before and during peak visitation times by other individuals (15:00-23:00). As a result, we suggest that dung kicking by territorial males amplifies signal strength, such that their dung odours are the most prominent and easily detectable by individuals visiting the middens. This would then better signal territorial ownership to both potential rivals and potential mates.

The objective of this study was to determine the major nutrient composition of Indian rhinoceros milk ( Rhinoceros unicornis) over the first 13 mo of an 18-mo lactation period and to compare the results to those of previous studies on rhinoceros, African elephant ( Loxodonta africana), and horse milk ( Equus ferus caballus). The following parameters were measured: dry matter (DM), crude ash (ASH), crude protein (CP), ether extract (EE), nitrogen-free extract (NFE; calculated), lactose, calcium (Ca), phosphorus (P), magnesium (Mg), fatty acids (FAs), and gross energy (GE). DM, ASH, CP, and EE were determined with a proximate analysis, lactose with infrared spectroscopy and an enzymatic method, minerals with an autoanalyzer, FA with gas chromatography, and GE with bomb calorimetry. Milk samples were collected from two Indian rhinoceros cows from Zoo Basel. Rhino A gave birth to her third calf on 10 September 2012; three samples were collected and analyzed (colostrum, milk 1 wk and 2 wk postpartum). Rhino B gave birth to her eighth calf on 05 October 2013; samples were collected and 15 were chosen for the analyses (from colostrum to 13 mo postpartum). The composition of rhino B's colostrum was 13.8% DM (wet-weight basis), 4.8% ASH, 61.8% CP, 0.7% EE, 32.6% NFE, 26.7% lactose, 0.59% Ca, 0.54% P, 0.2% Mg (DM basis), and 20.3 MJ GE/kg DM. Rhino B's sample collected 13 mo postpartum averaged 8.0% DM (wet-weight basis), 3.6% ASH, 16.3% CP, 1.8% EE, 78.3% NFE, 84.7% lactose, 0.54% Ca, 0.48% P, 0.09% Mg (on DM basis), and 17.43 MJ GE/kg DM. The main FAs in rhino B's and rhino A's samples were C10 : 0, C12 : 0, C16 : 0, C18 : 1n9c, and C18 : 2n6c. Milk of the Indian rhinoceros is low in fat and protein but high in lactose, which is comparable to the milk composition of other rhinoceros species and horses, but not African elephants.

The Oldowan site HWK EE (Olduvai Gorge, Tanzania) has yielded a large fossil and stone tool assemblage at the transition from Lower to Middle Bed II, ∼1.7 Ma. Integrated tooth wear and stable isotope analyses were performed on the three most abundant ungulate taxa from HWK EE, namely Alcelaphini, cf. Antidorcas recki (Antilopini) and Equus oldowayensis (Equini), to infer dietary traits in each taxon. Some paleodietary changes were observed for cf. A. recki and E. oldowayensis based on tooth wear at the transition from the Lemuta to the Lower Augitic Sandstone (LAS) interval within the HWK EE sequence. Stable carbon and oxygen isotope data show no significant changes in bulk diet or hydroclimate between the Lemuta and LAS intervals. The combined tooth wear and stable isotope data suggest similar paleoecological conditions across the two HWK EE intervals, but that differences in vegetation consumed among ungulates may have resulted in changes in dietary niches. Integrating tooth wear and stable isotope analyses permits the characterization of ungulate diets and habitats at HWK EE where C 4 dominated and minor mixed C 3 and C 4 habitats were present. Our results provide a better understanding of the paleoenvironmental conditions of the Lemuta and LAS intervals. The LAS assemblage was mostly accumulated during relatively dry periods at Olduvai Gorge when grasses were not as readily available and grazing animals may have been more nutritionally-stressed than during the formation of the Lemuta assemblage. This helps to contextualize variations in hominin and carnivore feeding behavior observed from the faunal assemblages produced during the two main occupations of the site., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Despite their size and potentially dangerous demeanor, the rhinoceros has been a preferred subject of wildlife reproductive scientists. Several factors contribute to this taxon's popularity including the ability to utilize insightful tools like non-invasive hormone metabolite monitoring and transrectal ultrasonography, the necessity for mate introductions to coincide with the female's estrus when breeding certain species or individuals, and the desire to develop assisted reproductive technologies to facilitate the genetic management and ultimate sustainability of small, managed populations in human care. The resulting profusion of rhinoceros reproductive studies has revealed significant species-specific characteristics and exposed the prevalence of aberrant reproductive activity within this taxon. Of equal importance, it has guided necessary intervention and enhanced our success in overcoming challenges associated with breeding rhinoceroses., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Describing vocal repertoires represents an essential step towards gaining an overview about the complexity of acoustic communication in a given species. The analysis of infant vocalisations is essential for understanding the development and usage of species-specific vocalisations, but is often underrepresented, especially in species with long inter-birth intervals such as the white rhinoceros. Thus, this study aimed for the first time to characterise the infant and juvenile vocal repertoire of the Southern white rhinoceros and to relate these findings to the adult vocal repertoire. The behaviour of seven mother-reared white rhinoceros calves (two males, five females) and one hand-reared calf (male), ranging from one month to four years, was simultaneously audio and video-taped at three zoos. Normally reared infants and juveniles uttered four discriminable call types (Whine, Snort, Threat, and Pant) that were produced in different behavioural contexts. All call types were also uttered by the hand-reared calf. Call rates of Whines, but not of the other call types, decreased with age. These findings provide the first evidence that infant and juvenile rhinoceros utter specific call types in distinct contexts, even if they grow up with limited social interaction with conspecifics. By comparing our findings with the current literature on vocalisations of adult white rhinoceros and other solitary rhinoceros species, we discuss to which extent differences in the social lifestyle across species affect acoustic communication in mammals.

Videos recorded with infrared camera traps placed in petroleum contaminated areas of the Peruvian Amazon have shown that four wildlife species, the most important for indigenous peoples' diet (lowland tapir, paca, red-brocket deer and collared peccary), consume oil-contaminated soils and water. Further research is needed to clarify whether Amazonian wildlife's geophagy can be a route of exposure to petrogenic contamination for populations living in the vicinity of oil extraction areas and relying on subsistence hunting., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Extant rhinoceroses share the characteristic nasal horn, although the number and size of horns varies among the five species. Although all species are herbivores, their dietary preferences, occipital shapes, and common head postures vary. Traditionally, to predict the "usual" head posture (the most used head posture of animals during normal unstressed activities, i.e., standing) of rhinos, the occipital shape was used. While a backward inclined occiput implies a downward hanging head (often found in grazers), a forward inclined occiput is related to the horizontal head posture in browsing rhinos. In this study, the lateral semicircular canal (LSC) of the bony labyrinth was virtually reconstructed from µCT-images in order to investigate a possible link between LSC orientation and head posture in extant rhinoceroses. The usual head posture was formerly reconstructed for several non-rhinoceros taxa with the assumption that the LSC of the inner ear is held horizontal (parallel to the ground) during normal activity of the living animal. The current analysis of the LSC orientation resulted in a downward inclined usual head posture for the grazing white rhinoceros and a nearly horizontal head posture in the browsing Javan rhinoceros. The other three browsing or mixed feeding species show subhorizontal (closer to horizontal than a downgrade inclination) head postures. The results show that anatomical and behavioral aspects, like occipital shape, presence and size of horns/tusk-like lower incisors, as well as feeding and feeding height preferences influence the usual head posture. Because quantitative behavioral data are lacking for the usual head postures of the extant rhinos, the here described relationship between the LSC orientation and the resulting head posture linked to feeding preferences gives new insights. The results show, that the inner ear provides additional information to interpret usual head postures linked to feeding preferences that can easily be adapted to fossil rhinoceroses., (© 2017 Wiley Periodicals, Inc.)

The population of free-roaming white rhinoceroses (Ceratotherium simum) is under serious threat. Captive breeding of this species is therefore becoming more important, but this is challenging and often not successful. Obtaining reproductive reference values is a crucial aspect of improving these breeding results. In this study performed between 2008 and 2016, reproductive performance was analysed in 1,354 animals kept in a 8000 hectares game-ranched environment. Descriptive statistics of this captive population showed an average annual herd growth (%) of 7 .0±0.1 (min -9 -max 15). Average calving rates were calculated as an annual calving rate of 20% and biennial calving rate of 37% adult females calving per year. Females had a median age of 83.2 months at first calving (IQR 72.9-110.7) and inter-calving intervals of 29.2 (IQR 24.6-34.8) months. Furthermore, translocations of animals did not interfere with reproductive success in terms of inter-calving periods or age at first calving. Multivariate models showed a clear seasonal calving pattern with a significant increase of the number of calvings during December-April when compared to April-December. Our results did not show any significant skewed progeny sex ratios. Weather observations showed no significant influence of rain or season on sex ratios of the calves.

In order for an olfactory signal to be effective, it must persist in the environment for an extended period. White rhino dung odours transmit information about sex, age, territorial and oestrous states. As these odours relay important information, temporal changes in the odour emission rate and/or composition may be critical in order for other individuals to obtain this information. Here, we examine how the dung odours of adult white rhinos (male: territorial and non-territorial; female: oestrous and non-oestrous) change over the short (hours) and long (seasons) term using headspace extraction. Additionally, we measure seasonal midden visitation and defecation frequency to link behaviours to seasonal changes in odour longevity. We found that during the dry season, territorial male dung odours persisted for 48 hours, while the odours of all other adults persisted for >48 hours. The high temperature and humidity of the wet season did not affect odour longevity of oestrous females, but decreased dung odour longevity (i.e. increased emission) of territorial males, non-territorial males, and non-oestrous females. Despite this reduction, neither males nor females adjusted their seasonal visitation or defecation frequency. With regard to a temporal indicator, 3,7-dimethly-1,6-octadiene signalled dung age during the wet season, while acetophenone signalled dung age during the dry season. Ultimately, our results provide the first detailed account of temporal changes of dung VOCs, and highlight the limitations of dung as a signalling medium.

This study sought to determine whether the retinal organization of the white rhinoceros (Ceratotherium simum), a large African herbivore with lips specialized for grazing in open savannahs, relates to its foraging ecology and habitat. Using stereology and retinal wholemounts, we estimated a total of 353,000 retinal ganglion cells. Their density distribution reveals an unusual topographic organization of a temporal (2,000 cells/mm 2 ) and a nasal (1,800 cells/mm 2 ) area embedded within a well-defined horizontal visual streak (800 cells/mm 2 ), which is remarkably similar to the retinal organization in the black rhinoceros. Alpha ganglion cells comprise 3.5% (12,300) of the total population of ganglion cells and show a similar distribution pattern with maximum densities also occurring in the temporal (44 cells/mm 2 ) and nasal (40 cells/mm 2 ) areas. We found higher proportions of alpha cells in the dorsal and ventral retinas. Given their role in the detection of brisk transient stimuli, these higher proportions may facilitate the detection of approaching objects from the front and behind while grazing with the head at 45 °. Using ganglion cell peak density and eye size (29 mm, axial length), we estimated upper limits of spatial resolving power of 7 cycles/deg (temporal area), 6.6 cycles/deg (nasal area), and 4.4 cycles/deg (horizontal streak). The resolution of the temporal area potentially assists with grazing, while the resolution of the streak may be used for panoramic surveillance of the horizon. The nasal area may assist with detection of approaching objects from behind, potentially representing an adaptation compensating for limited neck and head mobility. J. Comp. Neurol., 525:2484-2498, 2017. © 2017 Wiley Periodicals, Inc., (© 2017 Wiley Periodicals, Inc.)

Kaldar Cave is a key archaeological site that provides evidence of the Middle to Upper Palaeolithic transition in Iran. Excavations at the site in 2014-2015 led to the discovery of cultural remains generally associated with anatomically modern humans (AMHs) and evidence of a probable Neanderthal-made industry in the basal layers. Attempts have been made to establish a chronology for the site. These include four thermoluminescence (TL) dates for Layer 4, ranging from 23,100 ± 3300 to 29,400 ± 2300 BP, and three AMS radiocarbon dates from charcoal samples belonging to the lower part of the same layer, yielding ages of 38,650-36,750 cal BP, 44,200-42,350 cal BP, and 54,400-46,050 cal BP (all at the 95.4% confidence level). Kaldar Cave is the first well-stratified Late Palaeolithic locality to be excavated in the Zagros which is one of the earliest sites with cultural materials attributed to early AMHs in western Asia. It also offers an opportunity to study the technological differences between the Mousterian and the first Upper Palaeolithic lithic technologies as well as the human behaviour in the region. In this study, we present a detailed description of the newly excavated stratigraphy, quantified results from the lithic assemblages, preliminary faunal remains analyses, geochronologic data, taphonomic aspects, and an interpretation of the regional paleoenvironment.

Little is known about the mechanisms causing tremors during immobilisation of rhinoceros and whether cardiorespiratory supportive interventions alter their intensity. Therefore, we set out to determine the possible mechanisms that lead to muscle tremors and ascertain whether cardiorespiratory supportive interventions affect tremor intensity. We studied tremors and physiological responses during etorphine-azaperone immobilisation in eight boma-held and 14 free-living white rhinoceroses. Repeated measures analysis of variance and a Friedman test were used to determine differences in variables over time and between interventions. Spearman and Pearson correlations were used to test for associations between variables. Tremor intensity measured objectively by activity loggers correlated well (p < 0.0001; r2 = 0.9) with visual observations. Tremor intensity was greatest when animals were severely hypoxaemic and acidaemic. Tremor intensity correlated strongly and negatively with partial pressure of oxygen (PaO2 ) (p = 0.0003; r2 = 0.9995) and potential of hydrogen (pH) (p = 0.02, r2 = 0.97). It correlated strongly and positively with adrenaline concentrations (p = 0.003; r2 = 0.96), and adrenaline correlated strongly and negatively with PaO2 (p = 0.03; r2 = 0.95) and pH (p = 0.03; r2 = 0.94). Therefore, hypoxaemia and acidaemia were likely associated with the intensity of tremors through their activation of the release of tremorgenic levels of adrenaline. Tremors can be reduced if circulating adrenaline is reduced, and this can be achieved by the administration of butorphanol plus oxygen insufflation. Furthermore, to assist with reducing the risks associated with rhinoceros immobilisation, tremor intensity could be used as a clinical indicator of respiratory and metabolic compromise.

Mammals commonly communicate olfactorily via urine. However, the extent to which they communicate via dung, another waste product, is unknown. Behavioural studies suggest that mammals can obtain information from dung odours but are unclear about the information transmitted. Moreover, an understanding of the volatile organic compounds (VOCs) released from dung is limited. To address this, we analysed the odours emitted from the dung of free-ranging white rhinos, and found that 2,3-dimethylundecane signalled an individual's sex, heptanal discriminated age class, nonane defined male territorial status and 2,6-dimethylundecane indicated female oestrous state. To validate these findings, we artificially reproduced key elements of the territorial and oestrous odour profiles (i.e. profiles likely to elicit behavioural responses from receivers). We then exposed free-ranging territorial males to these odours. In response, males elicited behaviours associated with the specific odours (e.g. territorial male (potential threat): reduced latency in assuming vigilance; oestrous female (potential mate): increased investigation). These results indicate that the VOCs identified from the dung of free-ranging individuals do transmit key information. Moreover, as white rhinos of all ages and sexes defecate communally, middens probably act as information centres. Furthermore, as many other mammals defecate communally, olfactory communication via dung odours is likely a widespread phenomenon., (© 2017 The Author(s).)

The spatial scale of animal space use, e.g. measured as individual home range size, is a key trait with important implications for ecological and evolutionary processes as well as management and conservation of populations and ecosystems. Explaining variation in home range size has therefore received great attention in ecological research. However, few studies have examined multiple hypotheses simultaneously, which is important provided the complex interactions between life history, social system and behaviour. Here, we review previous studies on home range size in ungulates, supplementing with a meta-analysis, to assess how differences in habitat use and species characteristics affect the relationship between body mass and home range size. Habitat type was the main factor explaining interspecific differences in home range size after accounting for species body mass and group size. Species using open habitats had larger home ranges for a given body mass than species using closed habitats, whereas species in open habitats showed a much weaker allometric relationship compared with species living in closed habitats. We found no support for relationships between home range size and species diet or mating system, or any sexual differences. These patterns suggest that the spatial scale of animal movement mainly is a combined effect of body mass, group size and the landscape structure. Accordingly, landscape management must acknowledge the influence of spatial distribution of habitat types on animal behaviour to ensure natural processes affecting demography and viability of ungulate populations., (© 2016 The Author(s).)

Forstercooperiines are a group of primitive rhinocerotoids with a relatively large body size in the Eocene, and normally considered to be closely related to Giant Rhinos. Here we report a new forstercooperiine, Pappaceras meiomenus sp. nov., from the late Early Eocene Arshanto Formation, Erlian Basin, Nei Mongol, China. Pappaceras is the earliest known unequivocal rhinocerotoid, and the holotype of the new species, represented by the most complete cranium of forstercooperiines known to date, shows the earliest evidence of reduction of the first upper premolar in rhinocerotoids, and resembles paraceratheriine Juxia in basicranial features, supporting the interpretation that the forstercooperiine clade is ancestral to paraceratheriines. The new species also displays some similarities with amynodontids in craniodental structures. Phylogenetic analysis identifies P. meiomenus as a basal taxon of the monophyletic forstercooperiines. It also reveals novel phylogenetic relationships of early rhinocerotoids that indicates Uintaceras is the sister group of paraceratheriids, to which amynodontids are more closely related than to any other group of rhinocerotoids. Furthermore, the eggysodontid clade is excluded from hyracodontids and placed as the sister group of rhinocerotids. Hyracodontidae, excluding paraceratheriids and eggysodontids, is placed as the most basal group of the rhinocerotoids.

When immobilising wildlife, adverse side effects can include hypoxaemia, acidosis and hypertension. Pulmonary gas exchange and acid-base status were evaluated during immobilisation of 25 free-ranging and one boma-held black rhinoceros (Diceros bicornis) in Zimbabwe. The effect of different body positions on arterial oxygenation was evaluated. A combination of the following drugs was used: an opioid (etorphine or thiafentanil), azaperone and an a2 -adrenoceptor agonist (detomidine or xylazine). Respiratory and heart rates, rectal temperature and pulse oximetry-derived haemoglobin oxygen saturation were recorded. Serial arterial blood samples were analysed immediately in the field. Marked hypoxaemia and hypercapnia were recorded in immobilised free-ranging black rhinoceroses. Arterial oxygenation was higher during sternal compared to lateral recumbency. Most rhinoceroses developed acidaemia of respiratory and metabolic origin. Initially high lactate concentrations in free-ranging rhinoceroses decreased during immobilisation. Pulse oximetry was unreliable in the detection of hypoxaemia. Positioning in sternal recumbency and routine use of oxygen supplementation are recommended in the management of immobilised rhinoceroses as measures to improve arterial oxygenation.

Forelimb morphology is an indicator for terrestrial locomotor ecology. The limb morphology of the enigmatic tapir (Perissodactyla: Tapirus) has often been compared to that of basal perissodactyls, despite the lack of quantitative studies comparing forelimb variation in modern tapirs. Here, we present a quantitative assessment of tapir upper forelimb osteology using three-dimensional geometric morphometrics to test whether the four modern tapir species are monomorphic in their forelimb skeleton. The shape of the upper forelimb bones across four species (T. indicus; T. bairdii; T. terrestris; T. pinchaque) was investigated. Bones were laser scanned to capture surface morphology and 3D landmark analysis was used to quantify shape. Discriminant function analyses were performed to reveal features which could be used for interspecific discrimination. Overall our results show that the appendicular skeleton contains notable interspecific differences. We demonstrate that upper forelimb bones can be used to discriminate between species (>91% accuracy), with the scapula proving the most diagnostic bone (100% accuracy). Features that most successfully discriminate between the four species include the placement of the cranial angle of the scapula, depth of the humeral condyle, and the caudal deflection of the olecranon. Previous studies comparing the limbs of T. indicus and T. terrestris are corroborated by our quantitative findings. Moreover, the mountain tapir T. pinchaque consistently exhibited the greatest divergence in morphology from the other three species. Despite previous studies describing tapirs as functionally mediportal in their locomotor style, we find osteological evidence suggesting a spectrum of locomotor adaptations in the tapirs. We conclude that modern tapir forelimbs are neither monomorphic nor are tapirs as conserved in their locomotor habits as previously described. J. Morphol. 277:1469-1485, 2016. © 2016 Wiley Periodicals, Inc., (© 2016 Wiley Periodicals, Inc.)

Preventing obesity in zoo animals is increasingly recognized as an important husbandry objective. To achieve this goal, body condition scoring (BCS) systems are available for an ever-increasing number of species. Here, we present a BCS for the greater one-horned rhinoceros (Rhinoceros unicornis) based on an evaluation (on a scale from 1 to 5) of seven different body regions, and report resulting scores for 62 animals from 27 facilities, based on digital photographs. In animals above 4 years of age, this BCS correlated with the body mass:shoulder height ratio. Although differences between the sexes for individual regions were noted (with consistently higher scores in males for the neck and shoulder and in parous females for the abdomen), the average BCS of all regions did not differ significantly between males (4.3 ± 0.4) and females (4.1 ± 0.5). Linking the BCS to results of a questionnaire survey and studbook information, there were no differences in BCS between animals with and without foot problems or between parous and non-parous females. In a very limited sample of 11 females, those eight that had been diagnosed with leiomyoma in a previous study had a higher BCS (range 3.9-4.9) than the three that had been diagnosed as leiomyoma-free (range 3.5-3.7). The BCS was correlated to the amount of food offered as estimated from the questionnaire. Adjusting the amounts and the nutritional quality of the diet components is an evident measure to maintain animals at a target BCS (suggested as 3-3.5). Zoo Biol. 35:432-443, 2016. © Wiley Periodicals, Inc., (© 2016 Wiley Periodicals, Inc.)

Objectives: To adapt the algorithm for the calculation of oxygen saturation to the blood characteristics of the white rhinoceros by two different methods and to determine the accuracy of conventional pulse oximetry measurements., Study Design: Adaptation of two mathematical models of the oxygen dissociation curve (ODC)., Animals: Twenty-five captive white rhinoceros (Ceratotherium simum), including 12 males and 13 females, aged 6-32 years., Methods: During 33 anaesthetic events, 94 arterial blood gas samples with 72 simultaneous pulse oximetry measurements were analysed. The calculation of oxygen saturation was adapted to the characteristics of rhinoceros blood using two different methods. Firstly, a mathematical model developed in 1984 and, secondly, an oxygen status algorithm (OSA) produced by the same developer in 2005 were tested for their applicability for clinical use., Results: When arterial partial pressure of oxygen is >7.98 kPa (60 mmHg), oxygen saturation exceeds 95%. At partial pressures of 6.12-6.52 kPa (46-49 mmHg) Method 1 determined oxygen saturations of 92.5-95.3% and Method 2 oxygen saturations of 90.2-91.6%. Both methods resulted in similar ODCs and accounted for the low p50 value of rhinoceros blood. Method 1 provided better adaptation in respect to the physiological parameters of the rhinoceros, especially with regard to the Bohr effect, than Method 2. Pulse oximetry was an unreliable method of monitoring arterial oxygen saturation during general anaesthesia in this species., Conclusion: Adapting the oxygen saturation algorithm to consider the left shift of the ODC provides a useful tool for monitoring oxygen status, especially as pulse oximetry is insufficiently accurate. Experimental determination of the complete Hill curve is required to further validate and optimize the algorithm for use in the white rhinoceros., Clinical Relevance: The method will facilitate the accurate interpretation of oxygen saturation calculated by blood gas analysis in white rhinoceros., (© 2016 Association of Veterinary Anaesthetists and the American College of Veterinary Anesthesia and Analgesia.)

The objective of this study was to design an artificial insemination (AI) protocol using cryopreserved spermatozoa to obtain pregnancies in captive Indian rhinoceroses (Rhinoceros unicornis). Four methods developed varied by timing and approach, as follows; Method 1: females (n=2) were inseminated pre- and post-ovulation under general anesthesia, Method 2: females (n=2) were inseminated pre-ovulation without anesthetic via endoscopy, Method 3: females (n=1) were inseminated pre-ovulation without anesthetic via manual insertion of an insemination catheter, Method 4: females (n=2) were inseminated same as Method 3 with the addition of standing sedation. Semen deposition site varied as a result of changes in AI technology and experience. All females conceived following intrauterine AI using three methods. Four pregnancies (n=3 females) produced via Method 3 and 4 resulted in term births (n=2 male calves, n=2 female calves) at 481.8±12.8days post-AI. Unfortunately, two early pregnancy losses were documented in a fourth female conceiving via Method 2. Pregnancy rates were 0%, 22%, 17%, and 50% for Method 1-4, respectively. Method 3 and 4 rates improved to 29% and 67%, respectively when accounting for AI's conducted only on ovulatory estrous cycles. Spermatozoa (n=5 males) were cryopreserved 0.3-9.3 y prior to successful AI procedures. The lowest dose of frozen-thawed sperm resulting in conception was 500×10(6) motile sperm. Mean time from AI to ovulation in conceptive and non-conceptive cycles was 26±11.8h and 66±80.7h, respectively., (Copyright © 2016 Elsevier B.V. All rights reserved.)

With only three living individuals left on this planet, the northern white rhinoceros (Ceratotherium simum cottoni) could be considered doomed for extinction. It might still be possible, however, to rescue the (sub)species by combining novel stem cell and assisted reproductive technologies. To discuss the various practical options available to us, we convened a multidisciplinary meeting under the name "Conservation by Cellular Technologies." The outcome of this meeting and the proposed road map that, if successfully implemented, would ultimately lead to a self-sustaining population of an extremely endangered species are outlined here. The ideas discussed here, while centered on the northern white rhinoceros, are equally applicable, after proper adjustments, to other mammals on the brink of extinction. Through implementation of these ideas we hope to establish the foundation for reversal of some of the effects of what has been termed the sixth mass extinction event in the history of Earth, and the first anthropogenic one. Zoo Biol. 35:280-292, 2016. © 2016 The Authors. Zoo Biology published by Wiley Periodicals, Inc., (© 2016 The Authors. Zoo Biology published by Wiley Periodicals, Inc.)