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Global climate change, habitat fragmentation, and human interference have resulted in a significant, ongoing decline in the population of goitered gazelles. Effective conservation strategies require an understanding of resource requirements of threatened species, such as dietary needs. Therefore, we aimed to elucidate the food composition and seasonal dietary changes of goitered gazelles through microhistological analyses of fresh feces. Fabaceae (11.5%), Gramineae (9.4%), Chenopodiaceae (20.2%), Asteraceae (10.1%), and Rosaceae (19.5%) formed the primary dietary components of goitered gazelle. Additionally, Krascheninnikovia arborescens (13.4%) and Prunus sibirica (16.3%) were identified as the key forage plants. Forbs (50.4%) were the predominant plants for grazing throughout the year, particularly in the spring (72.9%). The proportion of trees in the diet was highest in the autumn (36.7%) and comparatively lower in other seasons. Furthermore, the proportions of shrubs (22.0%) and graminoids (14.8%) both reached their peaks in the winter. Our findings indicate that goitered gazelles strategically forage seasonally to cope with resource bottlenecks, enhancing their adaptability to arid and semi-arid habitats. Our study provides essential ecological information for the conservation of goitered gazelles and emphasizes the importance of dietary studies of species of ecological significance in environmentally sensitive areas.

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Interspecific hybridization increasingly occurs in the course of anthropogenic actions, such as species translocations and introductions, and habitat modifications or occurs in sympatric species due to the shortage of conspecific mates. Compared with anthropogenically caused hybridization, natural hybridization is more difficult to prove, but both play an important role in conservation. In this study, we detected hybridization of two gazelle sister species, Gazella bennettii (adapted to dry areas) and Gazella subgutturosa (adapted to open plains), in five habitat areas, where G. bennettii naturally occur in central Iran. The hybrids have a nuclear genomic identity (based on two introns), habitat preference, and phenotype of G. bennettii, but the mitochondrial identity (based on cyt b) of G. subgutturosa. We suggest that natural hybridization of female G. subgutturosa and male G. bennettii happened twice in central Iran in prehistoric times, based on the haplotype pattern that we found. However, we found indications of recent hybridization between both species under special circumstances, for example, in breeding centers, due to translocations, or in areas of sympatry due to the shortage of conspecific mates. Therefore, these two species must be kept separately in the breeding centers, and introduction of one of them into the habitat of the other must be strictly avoided.

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The phylogeny and species boundaries of Gazella subgutturosa and G. marica have been long debated. The achievements of past conservation efforts have been compromised by a lack of knowledge about the phylogeny and taxonomic status of different populations. We integrated the recent genetic findings by previous studies with morphometric analyses and ecological niche modeling (ENM) to assess discreteness among populations of these gazelle species in Asia. Taxonomic diversity of gazelles was investigated by using principal components analysis (PCA) based on 14 cranial measures of male skulls. Ecological niche divergence was examined based on a PCA on climatic factors and a species distribution modeling (SDM) with environmental variables. Morphometric results indicated substantial differentiation in size between skulls of the western Zagros Mountains including west and south-western Iran and Arabian Peninsula from all other samples east of the Zagros Mountains from Iran to China. ENM also revealed that gazelles in the east and west of Zagros Mountains occupy distinct niches and that there are apparent areas of disconnection across the goitered gazelle suitable range. A complete divergent niche occupation was also observed between goitered gazelles of northern Mongolia and other populations of the species, except those in China. Taking the inferences from ENM and morphology together with previous genetics results, we conclude that gazelles in the west and south-west of Iran may represent G. marica. Also, our combined analyses revealed divergence among gazelles of Iran, Central Asia, and Mongolia/China. These results may pave the way for future studies and have conservation implications particularly for reintroduction/supplementation programs.

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Predation is a very powerful force that shapes many ungulate traits. It is widely known that increasing vigilance intensity is costly and leads to a decline of forage intake. Consequently, ungulates gather into larger groups to reduce an individual vigilance burden by using the "many eyes" effect and to enhance their survival through the "safety in numbers" effect. Vigilance rate and related aggregation changes are two of the most discussed antipredator responses of ungulates to predation risk, but less considered factors also have a significant impact. To enhance our knowledge on these impact factors, I investigated the antipredator responses (vigilance, staring, and escape running) of goitered gazelles. I found that: a) adult females demonstrated vigilance, staring and escape running significantly more often than adult males, sub-adults and fawns; b) adult gazelles (females and males), having more experience in responding to threats, were more vigilant than young individuals (sub-adults and fawns), having likely more involvement in finding food; c) during the birthing and hiding periods, mothers displayed higher rates of vigilance than in other seasons, while staring and escape running were observed most often at the end of the hiding period, when fawns altering their hiding strategy to following one, started to walk a lot in the absence of mothers, as well; and d) goitered gazelles changed their vigilance rate in response to predation risks before changing group size. The events of the yearly biological cycle (birthing/rearing young for females; rutting for males) also had significant impacts on the antipredator responses (vigilance rate and aggregation) in adult gazelles.

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Predation is usually the primary cause of infant death among ungulate species, with the annual variation in the survival of neonates over their first summer a major factor in the population dynamics of many ungulates. Consequently, the maternal rearing strategy of a species is crucial for its reproductive success. Since the roles mothers and fawns play in the implementation of antipredator strategies in hider species have been poorly understood until now, this paper considers this behavior in the goitered gazelle, which is a typical hider species. I found that within the first month after birth goitered gazelle mothers largely controlled the behavior of their fawns both during the active period (determining the timing of separation and reunion with fawns, movement direction and speed, and bed site location) and the hiding period (keeping the hiding fawns under continuous watch from a distance, especially right after birth). With age the fawns' mobility increased and cases of independent behavior of fawns apart from their mothers were found more often, though females continued to control their fawns' behaviors. The main elements of the goitered gazelles' maternal care strategy - generally related to protecting fawns from potential predator attack - are very similar to other hider species in both bovids and cervids, which demonstrates a standard set of maternal care behaviors, irrespective of predator type or its behavioral peculiarities. Such maternal behaviors, however, likely only decrease the predation losses to a certain extent.

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In ungulates, predation is often a major cause of infant mortality and likely plays an important role in shaping maternal care strategies that favor progeny survival. The anti-predator strategies of ungulates can be broadly categorized into two groups, hiding infants and following infants. We studied the maternal behavioral strategies of goitered gazelle, which is a typical representative of a hiding species. We found that shortly after birth, goitered gazelle mothers (1) stayed at the greatest distances from their hiding fawns; (2) spent the shortest amount of time together with their fawns, and then only for suckling (during the active phase) which was also the longest inter-suckling intervals (during the hiding phase); (3) kept twins separated in different hiding places and suckled them individually one after the other; (4) changed fawns' hiding places after every active period, travelling with them during the whole suckling bout; (5) oriented their body and muzzle most often toward their hiding young during the fawn's hiding phase; and (6) demonstrated the highest level of vigilance during their approaches to their hiding fawns. The anti-predator strategy of goitered gazelle females was similar to that observed in some North American (Antilocapra americana, Odocoileus hemionus and Odocoileus virginianus) and Eurasian (Dama dama and Capreolus capreolus) ungulate species that also demonstrate hiding behavior. Females of these species live on different continents with disparate environments and different predatory threats, but share anti-predator strategies, which are likely the result of convergent evolution in Bovids and Cervids.

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The mother-offspring social unit is a universal feature in the social life of all mammals and nursing is the most direct and vital component of maternal investment in young. Living in diverse environments, various ungulate species have different strategies for rearing offspring, from bearing a single, relatively large newborn and supplying only limited amounts of milk, to bearing several relatively small newborns with intensive post-partum lactation. In this paper, we consider the rearing strategy of goitered gazelle with a focus on suckling behavior, which, until now, has never been a subject of special investigation. Adult females of this species in their reproductive prime typically bear twins when environmental conditions are favorable, but the proportion of singletons increases when conditions are unfavorable. We expected that in goitered gazelles suckling intensity would be maximal during the first weeks after birth, and then decrease with the growth of the young; we also expected that twins would demand more energy, but receive significantly less maternal investment per young than singletons. We found that, indeed, suckling behavior had similar dynamics as typical of all bovid species, but our expectation for less maternal investment in twins vs. singletons was wrong. In reality, female goitered gazelles suckled twins significantly more intensively and terminated suckling less often compared to singletons. We concluded that in favorable situations females of high quality have the ability to show significantly more maternal investment in each twin, while singletons are typically born to weaker females. This ability of females to produce mostly twins allows goitered gazelles to take advantage of any favorable opportunity to quickly increase their population in an environment with unpredictable and abrupt yearly changes typical of the arid regions of Central Asia.

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Aggression serves a great variety of social functions, one of which is protection of individual territories from intruders. Territorial males of many antelope species show aggressive noncontact displays, and only rarely fight. It has been suggested that ungulate males tend to have more frequent and longer aggressive interactions with rivals of similar age or social status than with males of dissimilar status. In the present paper, we test whether territorial and non-territorial males behave in a similar manner and avoid fights, and whether or not they preferentially direct aggressive and longer agonistic interactions towards males of similar age or social status, rather than towards other classes of males. We found that territorial males usually avoided straight fights with peers, and instead mainly used noncontact displays in aggressive interactions. In contrast, non-territorial males used fights considerably more often, especially during the onset of territoriality in April to May, when these males had their most frequent aggressive interactions. Territorial bucks aggressively interacted most frequently with non-territorial males and significantly less often with other territorial males, but agonistic noncontact displays between territorial males lasted the longest. In contrast, non-territorial males addressed their aggressive noncontact displays and fights most often to peers and less frequently to sub-adults. Asymmetry in the social status of territorial vs. non-territorial males was likely responsible for the distinctively different agonistic behaviors shown by the two types of males, which in turn are likely due to the different costs and benefits each male can accrue from these aggressive interactions.

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Aggression in ungulates is a very common behavior which serves a great variety of social functions, the most important of which is territorial protection from intruders. Typically during the rut, territorial males in Antilopinae species have access to mating females, and territoriality leads to a drastic change in the males' lifestyle as they spend most of their aggressive efforts on protecting their territories from other males. In contrast, non-territorial males generally behave in a constant manner all year round, with most of their aggression spent on hierarchical interactions within groups. In this paper, we discuss the differences in aggressive behaviors between territorial and non-territorial males of goitered gazelles (Gazella subgutturosa). We found that territorial males of this species demonstrated most of their aggressive displays, such as threat postures and space-claim patterns, to other territorial males-neighbors, but rarely chased after them at the end of a conflict. Inversely, territorial males frequently chased non-territorial and immature males while only occasionally demonstrating aggressive patterns. Non-territorial males mostly chased other males in their agonistic interactions, displaying threat postures less often than territorial males, but also showing aggressive patterns not found in territorial males. So we concluded that territorial and non-territorial males of goitered gazelles had different repertoires of aggressive behaviors, and when adult males switched their social status from territorial during the rut to non-territorial outside the rut, they obviously changed their preference in aggressive behaviors accordingly.

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Most of the vocalizations of Antilopinae males are soft and usually only heard from a very close distance. The goitered gazelle is a rare exception to this rule, and during the rutting period territorial males of this species are among the noisiest antelopes. Rutting vocalization is such an essential part of the rutting behavior in goitered gazelle that adult males have a hypertrophic larynx, the muscle tissues of which increase considerably in size during the rut. We were interested in the frequency and variance with which male goitered gazelles emit their calls depending on an animal's age, reproductive status and time of the year in order to understand the main function of the rutting vocalizations. We found that roaring was mostly related to courting displays, while vocalizations during aggressive displays were less frequent in male-male interactions. Acoustic signals likely enhance courtship displays and also may aid in accelerating female ovulation, promoting synchronization of breeding cycles during the mating and birthing periods, which last only several days for most of the females in our population. We discuss the potential benefits of such behaviors and compare it to other species living in similarly extreme environments.

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We studied object-horning behaviour in goitered gazelles in the natural, arid environment of Kazakhstan over a 6-year period. We found that object-horning was used by adult males mostly as a threat display during territorial conflicts. Therefore object-horning was observed most frequently in territorial single males during the rut in November-December. Object-horning, though, also had a marking effect, with the males' use of this behaviour leaving visible traces that advertized the location of preorbital and urination-defecation scent marks. Therefore, this pattern also was observed linked with preorbital marking and urination-defecation marking behaviours, especially during the rut. Goitered gazelle males chose the most abundant and eatable shrubs for object horning. In contrast to other gazelle species, object-horning in goitered gazelle was observed much more frequently and at the same rate as preorbital and urination-defecation scent markings. This, then, proved a more vigorous and aggressive level of rutting behaviour of the goitered gazelle compared to tropical gazelles, and most likely connected to the short rutting period in the studied species. We concluded, therefore, that object-horning was a manifold phenomenon that played a very important role in goitered gazelle agonistic displays, but without loosing the marking intention of this behaviour.

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