Development of technology for relocating kulans to preserve their population and restore the biodiversity of their habitatAbstract
The reintroduction of the kulan (Equus hemionus kulan) to the Ile-Balkhash State Nature Reserve aims to restore its population within its historical range in Kazakhstan. The study is motivated by the species' near extinction and the need for effective conservation strategies. The goal is to develop a biological basis for capturing kulans from the Altyn-Emel National Park and releasing them into the reserve. The researchers used population monitoring, capturing, and safe transport methodologies. The results show that the current population in Altyn-Emel has reached its ecological limit, justifying the relocation of up to 60 kulans annually. Preparations for reintroduction include corral improvements, transportation logistics, and rules for release. Despite risks from predators and pathogens, careful monitoring should ensure a sustainable new population in Southern Balkhash, contributing to regional biodiversity restoration. The conclusion emphasizes that reintroduction is critical to kulan conservation and maintaining biodiversity in Kazakhstan.
Keywords:
kulan reintroduction; Equus hemionus kulan; Ile-Balkhash State Nature Reserve; biodiversity restoration; conservation strategies; population monitoring
Resumo
A reintrodução do kulan (Equus hemionus kulan) na Reserva Natural Estadual de Ile-Balkhash visa restaurar sua população dentro de sua área de distribuição histórica no Cazaquistão. O estudo é motivado pela quase extinção da espécie e pela necessidade de estratégias de conservação eficazes. O objetivo é desenvolver uma base biológica para capturar kulans do Parque Nacional Altyn-Emel e soltá-los na reserva. Os pesquisadores utilizaram metodologias de monitoramento populacional, captura e transporte seguro. Os resultados mostram que a população atual em Altyn-Emel atingiu seu limite ecológico, justificando a realocação de até 60 kulans anualmente. Os preparativos para a reintrodução incluem melhorias nos currais, logística de transporte e regras para soltura. Apesar dos riscos de predadores e patógenos, o monitoramento cuidadoso deve garantir uma nova população sustentável no Balkhash Meridional, contribuindo para a restauração da biodiversidade regional. A conclusão enfatiza que a reintrodução é fundamental para a conservação dos kulans e a manutenção da biodiversidade no Cazaquistão.
Palavras-chave:
reintrodução de kulan; Equus hemionus kulan; Reserva Natural do Estado de Ile-Balkhash; restauração da biodiversidade; estratégias de conservação; monitoramento populacional
1. Introduction
Natural habitat destruction and extinction threaten many animal species (Davaa et al., 2017). One such endangered species is the kulan (Equus hemionus kulan), the number of subspecies of which is the subject of scientific debate in literature, where scientists (Kuznetsova and Zhegallo, 2009; Yasynetska, 2023; Lushchekina et al., 2024; Lukarevsky and Gorelov, 2007) distinguish between 3 and 8 subspecies. Classifying the subspecies is challenging since genetic research indicates that the distinctions between them are negligible (Berkinbai et al., 2023a). Early, it was found that three kulan subspecies lived in the territory of modern Kazakhstan: Mongolian (Equus hemionus Pallas, 1775), Kazakhstani (Equus hemionus finschi Matschie, 1911) and Turkmenian (Equus hemionus onager (or kulan) Boddaert, 1785). By the beginning of the 20th century, the Kazakhstani subspecies had completely died out, and the Mongolian and Turkmenian had disappeared from the territory of modern Kazakhstan. The first attempts at reintroduction were made in 1953. Later, four groups of kulans were formed in Kazakhstan: Barsa-Kelmes (Qyzylorda Region), Altyn-Emel (Jetisu Region), Andasai (Zhambyl Region) and Aktau-Buzachinsky (Mangistau Region), but the low level of animal protection in the Mangistau region led to the disappearance of kulans from both in the Aktau-Buzachinsky reserve and beyond (Bahloul et al., 2001; Berber, 2008; Berkinbai et al., 2023b).
Nowadays, kulans occupy less than 1% of their historical range and are still absent from the central steppe in Kazakhstan (Nurgaziyev et al., 2024). Various assumptions have been made about the causes of fluctuations in the kulan population. In particular, Solomatin (1973) associates the low population growth rate with various anthropogenic and especially climatic factors, which exists more at the level of a hypothesis and is not confirmed by other studies. Subsequent research, including the works of Myrzhieva et al. (2020) and Almanova et al. (2023), supports the prevailing view among conservation biologists that a combination of unfavorable climatic conditions and human-induced pressures has contributed to the limited expansion of kulan populations. Specifically, at times of poor climatic circumstances (such as severe droughts), kulans were purposefully eradicated nearly to the point of extinction.
Some contradicting findings arise from the examination of the data collected on the dynamics of observation, social and behavioral ecology, and the predominance and allegiance of kulans at various times. On the one hand, intense poaching was the primary cause of the low number of kulans. The persecution of kulans in spring leads to the mass death of newborn foals, negatively affects reproductive activity and often causes to abortions due to stress in females. Trying to escape, kulans leave their pastures, what also leads to high mortality of foals, especially at the age of 0.5 to 1.5 years. On the other hand, it is noted that an increase in the kulan population may conflict with agriculture in the reserve. Therefore, it is essential to implement a population control plan that includes the removal of certain animals as soon as the kulan population exceeds the maximum number of individuals for a given region in order to prevent the detrimental effects of population expansion.
In this century, active work on the reintroduction of kulans began in 2017 with the involvement of international partners in Kazakhstan. The goal of these efforts is to create a biological basis for capturing kulans from the population in the Altyn-Emel State National Nature Park and their subsequent reintroduction to the Ile-Balkhash State Nature Reserve. Within the framework of this program, a set of organizational, economic and biotechnical measures was developed. We aimed to demonstrate the rationale behind the selection of areas for kulan resettlement, the computation of the maximum number of animals that can be captured and subsequently removed from the population, the technology involved in animal transportation, and the potential risks of harming biological diversity. It enables us to follow every step of the kulan's reintroduction based on a natural experiment, this study is unique in Central Asia. seedlings.
2. Materials and Methods
2.1. Study design
This study was designed to assess the feasibility and ecological outcomes of reintroducing kulans (Equus hemionus kulan) from Altyn-Emel State National Park to the Ile-Balkhash State Nature Reserve. The research includes several integrated components: monitoring the existing population dynamics in the source region, identifying suitable relocation sites, developing and applying capture and transportation protocols, and evaluating adaptation risks at the reintroduction site. Data were collected through direct population counts, environmental assessments, logistical route planning, and analysis of predator and disease threats.
2.1.1. Ethical approval
All procedures involving animals in this study were conducted in accordance with national legislation on wildlife handling and approved by the relevant environmental and veterinary authorities. The capture, transport, and release of kulans were carried out under the guidelines set in the methodological recommendations for the reintroduction of kulans developed by the Institute of Zoology and the Ministry of Ecology and Natural Resources of the Republic of Kazakhstan.
2.2. Ecological and behavioral characteristics of kulans
The diet of kulans depends on the seasons of the year and the yield of the main food. When grass is insufficient, the kulan supplements its diet with shrubs, subshrubs, tree branches and switches to mixed feeding in certain seasons. The presence of watering places is a decisive factor in the distribution of kulans, especially in the warm season. In the summer months, this species is mainly found within the accessibility zone of permanent water bodies. They can safely drink water with a salt content of up to 20 g/L, but lactating females avoid drinking water with a salt content of more than 10 g/L. In general, the kulan should have regular access to water, drinking 12-15 liters per day and up to 24 liters on hot days.
The diet of kulans depends on the seasons of the year and the yield of the main food. When grass is insufficient, the kulan supplements its diet with shrubs (namely, Atraphaxis spinosa and Haloxylon ammodendron), subshrubs (Artemisia pauciflora, Salsola arbusculiformis, and Kochia prostrata), tree branches (Populus pruinosa, Elaeagnus angustifolia, and Tamarix ramosissima) and switches to mixed feeding in certain seasons. The presence of watering places is a decisive factor in the distribution of kulans, especially in the warm season. In the summer months, this species is mainly found within the accessibility zone of permanent water bodies. They can safely drink water with a salt content of up to 20 g/L, but lactating females avoid drinking water with a salt content of more than 10g/L. In general, the kulan should have regular access to water, drinking 12-15L/day and up to 24L on hot days.
Like other odd-toed ungulates adapting to arid conditions, the kulan apparently lives in groups that split and merge, with the only stable unit being females and their foals. Under favorable conditions, females give birth to one foal annually . In a study of the Barsa-Kelmes kulan grouping on Barsa-Kelmes Island, the average fertility of adult mares was estimated at 66.6%, but varied depending on pasture conditions and winter severity.
2.3. Study areas
2.3.1. Altyn-Emel State National Park
The Altyn-Emel State National Park is located in the Kerbulak and Panfilov districts of the Almaty region. The total area of the park is 459,620 hectares. The national park includes the northern part of the Kapchagay reservoir and the right bank of the Ili River, the deserted mountains of Malye and Bolshiye Kalkan, Aktau, Katutau, the outermost southwestern spurs of the Dzungarian Alatau (Sholak, Degeres, Matai), the southern slope of the Altyn-Emel ridge and part of the Koyandytau ridge, as well as the vast intermountain valley of Konyrolen.
2.3.2. Ile-Bakhlash State Natural Reserve
The Ile-Bakhlash State Nature Reserve is located in the Balkhash district of the Almaty region. It was created in 2017 and covers an area of 415,164.2 hectares (Figure 1), including two zones - “Pribalkhashsky” and “Ile River Delta”. The territory of the reserve is limited by Lake Balkhash in the north, the western area are limited by the Ili Delta, the eastern border is a bend in Balkhash, and the southern one is naturally not marked and passes through the sands of Sary-Yesik-Atyrau. The Ile-Balkhash includes a section of the Sary-Yesik-Atyrau Desert, previously identified as a promising territory, and corresponds to the general plan for the restoration of the species in Kazakhstan.
2.4. Methodology for counting kulans in reserves in Kazakhstan
The methodology for monitoring kulan populations in Kazakhstan is based on long-standing practices developed and implemented by state institutions, particularly the Institute of Zoology of the Ministry of Education and Science of the Republic of Kazakhstan. The study relies on official monitoring data and methods that are routinely used by qualified specialists in protected areas.
The method of counting kulans is complicated by various factors related to the animal's behavior. Therefore, various methods of counting the kulan population are used for four kulan habitats in Kazakhstan - Barsa-Kelmes, Altyn-Emel, Andasai and Aktau-Buzachinsky National Parks. Aerial visual counting by plane was used in the Andasaisky Reserve. Another effective method was aerial counting of kulans at watering places from 1962 to 1976 (Lukarevsky and Gorelov, 2007). Absolute counting at watering places in the summer season was used in the Barsa-Kelmes Reserve. Since 1994, the Institute of Zoology has carried out annual absolute counts of kulans in the Altyn-Emel State National Park using vehicle-based survey routes. These official monitoring activities are conducted in February, covering two consecutive days: the western part of the park is surveyed using 10 parallel routes (approximately 390 km), and the eastern part using 6 routes (about 300 km). The present study utilizes this verified dataset as a foundation for evaluating population dynamics and informing reintroduction decisions. Surveys are aimed at direct counting of the total number of individuals, on parallel survey routes for two days in February of each year; 10 parallel routes with a total length of about 390 km on the first day in the western part and 6 parallel routes with a total length of about 300 km on the second day in the eastern part of the State National Nature Park (Figure 2).
2.5. Reintroduction
The kulan is classified as a rare and endangered species, and there are no approved standards for catching. Therefore, the manual “Methodological recommendations. Reintroduction of kulans (capture, transportation, temporary holding, release)” were improved for the reintroduction.
Safe capturing techniques were improved to capture kulans in the Altyn-Emel. Kulans were transported at an average speed of 40 km / h, while the behavior and condition of the animals were monitored by a video camera. The release was carried out at night in a pre-selected location near a reservoir. As a result, not a single animal died or was even injured during the capture, temporary holding and transportation. Before the release, an oval enclosure with an area of 22 hectares was built at the reintroduction site in Ile-Bakhlash.
3. Results
3.1. Counting kulans
Currently, the total population of kulan in Kazakhstan is about 4100 individuals. The Altyn-Emel and Barsa-Kelmes groups of kulans maintain a tendency towards steady growth, while the Aktau-Buzachinsky group has apparently disappeared. According to the data, 3,607 kulans inhabit in Altyn-Emel National Park in 2021. The population has peaked in this region, based on the drastically reduced growth rate (Table 1).
However, a 2023 report by Kaczensky (2023) confirmed the ongoing presence of a small kulan group in the Andasai Wildlife Reserve, suggesting that this subpopulation may be more resilient than earlier assessments indicated.
Between 2012 and 2015, Altyn-Emel's population grew by 23.4%; between 2015 and 2018, it increased by 15.5%; and between 2018 and 2021, it expanded by 2.8%. Figure 3 shows how the growth of kulans has sharply slowed down to almost complete cessation. This suggests that the population density of kulans in the national park has reached its limit under current conditions and it is desirable to remove some animals to regulate their numbers. The optimal size of a healthy population seems to be the maximum level at which there was still a stable annual growth of about 7-9%, i.e. the level of 2012-2015 - about 2500-3000 animals. The current level exceeds it by more than 600 individuals or 20%.
3.2. Rationale for the selection of the territory for the reintroduction of kulans
The Sary-Esik-Atyrau Desert in the Almaty Region was previously identified as a promising area for the reintroduction of the kulan, since this is their historical habitat. The main criteria for selecting areas for the release of the kulan in Kazakhstan are:
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The presence of protection, the inaccessibility of the area for poaching and low prospects for economic development. Experience from previous reintroduction efforts has shown that reliable protection is critical to success. For example, in the absence of effective security measures, kulan populations have disappeared from both the Aktau-Buzachi and Andasai reserves. In contrast, well-protected areas such as Altyn-Emel have seen stable population growth. This demonstrates that without sufficient enforcement, other environmental advantages become ineffective;
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Physical-geographical conditions. They must comply the environmental requirements for the kulan (Table 2);
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Pasture-watering and protective conditions. The main thing is the availability of water sources, since kulans need it daily during the hot season;
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Low numbers or absence of livestock, which compete with the kulan for pastures and watering places.
The territory selected for reintroduction in the Ile-Balkhash State Natural Park meets these requirements.
3.3. Calculation of the maximum permissible catch quantity of kulans for reintroduction
There are standards for the removal of other large ungulates such as elk and maral, which make up to 10% of the total population during hunting. It is perfectly permissible to remove up to 5-7% of any group of Kulan due to their somewhat lower fertility assuming the objective is not to diminish the population but to sustain its expansion (Kozhanov et al., 2023). With a population of 3,607 individuals, 5% would be 180 animals. It is also necessary to take into account the stability of the habitat in Altyn-Emel, the presence of a developed security service, which allows for a sharp reduction in animal mortality. For the purposes of reintroduction to the Ile-Balkhash, it is planned to catch and transport up to 60 kulans annually in 2021-2026. This will make the annual removal from the group in Altyn-Emel 3 times less than what is easily acceptable, that is, about 1.6%.
Thus, such a scale of catches will not have any impact on the population of kulans in Altyn-Emel. It should be mentioned that because of the rivalry for food supplies and watering spots with goitered gazelle, as well as other issues that are already starting to arise, it is imperative that the Kulan population in Altyn-Emel be reduced (Nauanova et al., 2024). It is obvious that a sharp increase in the scale of kulan transportation is desirable to create new or support other existing populations of the species. The Sary-Esik-Atyrau Desert located in Ile-Balkhash State Natural Park is the closest area to the national park, where it is possible to carry out fairly large-scale transportation. At the same time, since the capture of 60 animals will not help reduce the number of kulan in Altyn-Emel and is practically the limit of the technical capabilities of the project, it is necessary to plan additional captures and transportation to other regions of Kazakhstan, in order to reach the general level of resettlement of at least 200 animals per year for at least 4-5 years. We emphasize that capture for reintroduction is not actually removal from nature - animals are resettled from one habitat to another, and the population of the species in Kazakhstan as a whole is not affected in any way (Akhylbekova et al., 2022).
3.4. Improved methodology for capture of kulans
The main issue of the reintroduction is the physical capture of individuals. A successful method of capturing the kulan using a corral was used to translocate the kulan from Barsa-Kelmes Island to Altyn-Emel State National Park in recent years, specifically, in 2019 when the population in Altyn-Emel was still growing and had not yet reached its ecological threshold. This transfer was part of a national conservation initiative to consolidate kulan populations in regions with stronger protection infrastructure. Transport cages were installed around the perimeter of the corral and food was placed in them. After the kulans entered them, the cages were slammed shut. Inside the corral, the animals were sorted and immobilized, then loaded into a transport cage.
The trap itself has two entrances and guide wings on both sides, which will facilitate the capture of animals from different parts of the park. Both guide wings are curved and directed approximately to the west. The distance between the posts is 4-5 meters, the chain-link mesh used is 2.5 mm thick, 1.8 meters high and 10 meters long in a roll. The wire rod is 6.5 mm thick and stretched along the entire length in 3 rows. The net is stretched with an indent of 3 meters from the main enclosure. Figure 4 presents design specifics based on the experience of building traps in Altyn-Emel in 2017 and Barsa-Kelmes in 2019.
Recommended design of an additional trap-pen for catching kulans in the Altyn-Emel State National Park.
3.5. Transport of kulans
The planned route from the capture location in Altyn-Emel State National Park to the release site in Ile-Balkhash State Natural Park was thoroughly covered as part of the kulan transportation preparations in order to assess the road conditions and outline the route for usage in November 2021. The total distance was 595 km. The following difficulties for trucks were identified: steep climbs and descents when crossing dry gullies. The average vehicle speed on a section of a field road with difficult sections will be on average 20-25 km/h; the average speed on desert or steppe roads is on average 30 km/h; on asphalt roads on average 60-65 km/h and on the highway on average 90 km/h (Figure 5).
Scheme of the transportation route from the Altyn-Emel State Research and Production Enterprise to the holding place in the Ile-Balkhash State Nature Reserve, indicating the type of roads. (1) Starting point (capture site, Altyn-Emel); (2) Checkpoint 1 (desert segment); (3) Checkpoint 2 (steppe road); (4) Release site (Ile-Balkhash Reserve).
3.6. Keeping and releasing kulans
In release areas without natural boundaries, temporary holding is necessary. It is necessary for the animals to become established and accustomed to the area, in order to avoid immediate long-distance reconnaissance movements of the animals. In this case, the release is carried out after the birth of the young, at which time the females with the cubs cannot go far and the group is established in the area. The release was carried out in a paddock with central coordinates N46˚14’45.0 E075˚34’46.9), has an oval shape without corners, an area of 22 hectares. The length of the paddock fence is 2079 m. The paddock has 2 sliding gates, 2 drinking troughs, 2 canopies for sheltering the animals in bad weather. The paddock includes areas of saxaul, a small dune, open areas and a small area of a clearing of dense grassy vegetation. The kulans will be kept in the adaptation pen for up to 5-7 months, depending on winter and spring conditions and the social dynamics in the pen.
The proposed area for the reintroduction of the kulan, the Ile-Balkhash State Nature Reserve, is located within the former range of the species and fully meets its needs. The original population in the Altyn-Emel State National Park has reached its maximum, its growth rate has slowed down sharply, and the transportation of part of the population from there is not only possible, but also highly desirable to improve its condition. The proposed capture and movement of up to 60 kulans annually in 2021-2026 will amount to only about 1.6% of the current annual population in Altyn-Emel and will not affect its condition in any way, although, obviously, they will contribute to its recovery to some extent.
4. Discussion
The kulan is not a species alien to the release area, since it lived there before. Kulans lived in the Balkhash region until the beginning of the last century. The last kulan in Kazakhstan was killed on the northwestern shore of Lake Balkhash in 1930-1935 . Kulans have always moved, depending on feeding, watering and protective conditions. Therefore, it is presumed that the newly introduced kulans may also show quite active movements.
Competition with ungulates living in the Ile-Balkhash State Natural Park - wild boar, goitered gazelle, roe deer, saiga, as well as with the reintroduced Bukhara deer, is not to be expected, since they occupy different food niches. Problems could arise only when long-distance movements of animals are restricted, as is the case in Altyn-Emel, but in the Southern Balkhash region there are no obstacles to the migration of ungulates (Berkinbai et al., 2023b). Thus, there is no obvious mutual influence between mammals and the kulan, except for potential interactions with wolves. They are the only predators in the desert and steppe ecosystems of Central Asia that hunt the kulan. However, the quantitative predation of wolves on the kulan has not been studied, and it is believed that wolves mainly hunt foals, old or sick animals. Nevertheless, it can be assumed that the kulan will provide an additional food base for wolves. At the same time, it is unlikely that the wolf will pose a serious threat to the reintroduction of the kulan. If this problem arises, the wolf population can be quickly regulated. If a tiger appears in the region, the potential threat from wolves will decrease sharply, since tigers consume wolves. Tigers also hunt kulans, but a serious level of preying is not expected, since they will occupy generally different habitats (kulans - open areas, tigers - fairly dense plantings), encountering only at watering places.
Poisonous animals also pose a threat to the growth of the kulan population. Among them, the pit viper is found in the Ile-Bakhlash State Natural Park, although its numbers are low. Of the arachnids, the tarantula and black widow spider are dangerous. However, these species also inhabit the Altyn Emel State National Natural Park and have not led to the extinction of kulans.
It is important to note that very few cases of infectious diseases have been recorded in kulans (Delling et al., 2023; Kachenski et al., 2017). The only disease listed by the International Epizootic Bureau (OIE) that can affect the kulan is anthrax, a disease caused by the spore-forming bacteria Bacillus anthracis. Also, increasing pressure from grazing livestock on wild odd-toed ungulates on the remaining ecological islands is a serious concern in many parts of the world. Recent and repeated outbreaks of equine influenza A in Mongolia and the Xinjiang Autonomous Region of China have directly threatened reintroduced Przewalski's horses in southwestern Mongolia (Great Gobi Nature Reserve) and in the Kalamali Nature Reserve in Xinjiang, China (Kachenski et al., 2017), since special control of captured kulans is necessary. It is crucial to assess the age and health of captured animals in order to prevent old, sick or weakened animals from being transported. At the place of release, according to general requirements, a quarantine of at least 30 calendar days must be looked after with constant veterinary control.
In addition to the risks of reintroduction, there are also benefits from returning the kulan to the territory of its former habitat. In the wild, a comparative study was conducted on the feeding ecology of the kulan with wild and domestic ungulates. A study of the kulan, Przewalski's horse and the domestic horse (Equus caballus) noted that all three are predominantly herbivorous, but the kulan consumes most of the shrubs (Xu et al., 2012; Burnik Šturm et al., 2016). Therefore, moderate grazing by the kulan may improve the quality of pastures for other ungulates (Nasiyev and Bekkaliyev, 2019; Mukhambetov et al., 2023). A mosaic of grazing intensities is expected to increase bird biodiversity, as it allows species adapted to different grazing intensities to coexist. It is supposed that kulan grazing will increase the habitat of grazing-dependent bird species, such as the saxaul jay, but will not interfere with livestock production. There will be no problems with the division and use of feed.
5. Conclusions
This study aimed to evaluate the feasibility and ecological justification for the reintroduction of kulans (Equus hemionus kulan) from Altyn-Emel State National Park to the Ile-Balkhash State Nature Reserve. Through the analysis of population trends, landscape characteristics, capture and transport logistics, and environmental risks, the study has demonstrated that such reintroduction is both necessary and viable.
The Ile-Balkhash State Nature Reserve is located within the former range of the species and fully meets its needs. The analysis of long-term monitoring data indicates that the kulan population in Altyn-Emel has reached ecological saturation, necessitating population regulation to avoid resource competition and slow growth. The proposed capture and relocation of up to 60 kulans annually in 2021-2026 will amount to only about 1.6% of the current annual population size in Altyn-Emel. Additional preparations were also carried out to ensure successful reintroduction of the kulans, including improving the design of the corral, describing the features of the transportation route, and rules for keeping and releasing kulans into a new territory. The reintroduction strategy is supported by improved corral capture techniques, detailed transport route planning, and a carefully prepared release site. While potential threats from predators, disease, and climate variability remain, the identified risks are manageable under conditions of continuous veterinary monitoring and adaptive management. This methodology should lead to the creation of a new sustainable population of this species in the Southern Balkhash region, which will be an important step towards restoring the biodiversity of the region.
This reintroduction initiative represents a significant step toward restoring kulan populations and enhancing regional biodiversity in Kazakhstan. The study provides a practical framework for future translocations and contributes to broader conservation planning for desert-adapted ungulates.
Data Availability Statement
Data will be available on request.
References
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AKHYLBEKOVA, B., SEREKPAYEV, N., NOGAYEV, A. and ZHUMABEK, B., 2022. Pasture productivity depending on the method of pasture use in the steppe zone of northern Kazakhstan. Online Journal of Biological Sciences, vol. 22, no. 4, pp. 476-483. http://doi.org/10.3844/ojbsci.2022.476.483
» http://doi.org/10.3844/ojbsci.2022.476.483 -
ALMANOVA, Z., KENZHEGULOVA, S., KASHKAROV, A., KEKILBAYEVA, G., USSALINOV, E., YERZHAN, D., ZHAKENOVA, A. and ZVYAGIN, G., 2023. Changes in soil fertility indicators after long-term agricultural use in northern Kazakhstan. International Journal of Design & Nature and Ecodynamics, vol. 18, no. 5, pp. 1045-1053. http://doi.org/10.18280/ijdne.180504
» http://doi.org/10.18280/ijdne.180504 -
BAHLOUL, K., PERELADOVA, O.B., SOLDATOVA, N., FISENKO, G., SIDORENKO, E. and SEMPÉRÉ, A.J., 2001. Social organization and dispersion of introduced kulans (Equus hemionus kulan) and Przewalski horses (Equus przewalski) in the Bukhara Reserve, Uzbekistan. Journal of Arid Environments, vol. 47, no. 3, pp. 309-323. http://doi.org/10.1006/jare.2000.0714
» http://doi.org/10.1006/jare.2000.0714 - BERBER, A.P., 2008. Kulan in the betpak-dala desert: population, distribution, and condition. Vestnik Ohotovedenia, vol. 5, no. 3, pp. 286-289.
-
BERKINBAI, O., OMAROV, B.B., SULEIMENOV, M.Z.H., TASHIBAYEV, E.S. and BAYMUKHANBETOV, E.B., 2023a. Fauna of kulan (Equus hemionus Pallas, 1775) parasites in the State National Natural Park “Altyn-Emel”. Kazakhstan Zoological Bulletin, vol. 4, no. 1, pp. 42. http://doi.org/10.54944/kzbpm723hq44
» http://doi.org/10.54944/kzbpm723hq44 - BERKINBAI, O.B., OMAROV, B.B., SULEIMENOV, M.Z.H., DZHUSUPBEKOVA, N.M., BERDIKULOV, M.A., TUGANBAY, A. and BARBOL, B.I., 2023b. Formation of the parasite fauna of re-acclimatized Turkmenian kulan in Kazakhstan. Theory and Practice of Combating Parasitic Diseases, vol. 24, pp. 85-89.
-
BURNIK ŠTURM, M., GANBAATAR, O., VOIGT, C.C. and KACZENSKY, P., 2016. Sequential stable isotope analysis reveals differences in multi-year dietary history of three sympatric equid species in SW Mongolia. Journal of Applied Ecology, vol. 54, no. 4, pp. 1110-1119. http://doi.org/10.1111/1365-2664.12825 PMid:28717255.
» http://doi.org/10.1111/1365-2664.12825 - DAVAA, L., BATSAIKHAN, N., FAGAN, W., GHANDAKLY, E., KACZENSKY, P., MUELLER, T., SAMIYA, R., SCHAFBERG, R., STUBBE, A., STUBBE, M. and ANSORGE, H., 2017. First assessment of the population structure of the Asiatic wild ass in Mongolia. European Journal of Wildlife Research, vol. 64, pp. 3. http://doi.org/10.1007/s10344-017-1162-x.
-
DELLING, C., HELM, C.H., HEINZE, P., FRIEDMAN, M. and BÖTTCHER, D., 2023. First report of infection with metacestode stages of Echinococcus multilocularis in a kulan (Equus hemionus kulan) from Slovakia. International Journal for Parasitology. Parasites and Wildlife, vol. 22, pp. 80-83. http://doi.org/10.1016/j.ijppaw.2023.09.003 PMid:37736617.
» http://doi.org/10.1016/j.ijppaw.2023.09.003 - KACHENSKI, P., DOLDIN, R., ENCKE, D., LINNELL, J.D.K., LUKANOWSKI, O., SIDOROVA, T., SKLYARENKO, S., KISEBAYEV, T., WALZER, K., WARD, S., ZUTER, S., 2017. Feasibility study for kulan (Equus hemionus kulan) reintroduction into the central steppe of Kazakhstan. Trondheim: Norwegian Institute for Nature Research.
-
KACZENSKY, P., 2023 [viewed 28 October 2024]. Status and conservation of the asiatic wild ass across its range. Report prepared for the CMS Secretariat [online]. Bonn: Convention on the Conservation of Migratory Species of Wild Animals. Available from: https://www.cms.int/sites/default/files/document/cms-bfn_AWA%20TW_overview_report_en_final_corrected%202908.pdf
» https://www.cms.int/sites/default/files/document/cms-bfn_AWA%20TW_overview_report_en_final_corrected%202908.pdf -
KOZHANOV, Z., SERIKBAYEVA, A., KOZHANOVA, N., SYDYKOV, D., SADVAKASSOV, K. and MUKHAMETKALIEV, M., 2023. Impact of functional foods on improving the health of the Kazakh population. Advancements in Life Sciences, vol. 10, no. 4, pp. 555. http://doi.org/10.62940/als.v10i4.1844
» http://doi.org/10.62940/als.v10i4.1844 -
KUZNETSOVA, T. and ZHEGALLO, V., 2009. Equus (Hemionus) nalaikhaensis from the Pleistocene of Mongolia, the earliest kulan of Central Asia. Paleontological Journal, vol. 43, no. 5, pp. 574-583. http://doi.org/10.1134/S0031030109050141
» http://doi.org/10.1134/S0031030109050141 - LUKAREVSKY, V. and GORELOV, Y., 2007. Kulan (Equus hemionus Pallas 1775) in Turkmenistan. Exploration into the Biological Resources of Mongolia, no. 10, pp. 231-240.
-
LUSHCHEKINA, A., KARIMOVA, T. and NERONOV, V., 2024. The current state of kulan populations (Equus hemionus Pallas, 1775) in Central Asia countries. Arid Ecosyst, vol. 14, no. 1, pp. 79-88. http://doi.org/10.1134/S2079096124010104
» http://doi.org/10.1134/S2079096124010104 - MUKHAMBETOV, B., NASIYEV, B., ABDINOV, R., KADASHEVA, Z. and MAMYROVA, L., 2023. Influence of soil and climatic conditions on the chemical composition and nutritional value of Kochia prostrata feed in the arid zone of Western Kazakhstan. Caspian Journal of Environmental Sciences, vol. 21, no. 4, pp. 853-863.
-
MYRZHIEVA, A.B., SHABDARBAEVA, G.S., TURGANBAEVA, G.E., BALGIMBAEVA, A.I. and IBAZHANOVA, A.S., 2020. Ixodid ticks: epizootic status and methods for tick population size reduction. Online Journal of Biological Sciences, vol. 20, no. 4, pp. 166-175. http://doi.org/10.3844/ojbsci.2020.166.175
» http://doi.org/10.3844/ojbsci.2020.166.175 - NASIYEV, B. and BEKKALIYEV, A., 2019. The impact of pasturing technology on the current state of pastures. Annals of Agri Bio Research, vol. 24, no. 2, pp. 246-254.
-
NAUANOVA, A., SHAIKHIN, S., OSPANOVA, S., MAKENOVA, M., SHUMENOVA, N. and BOSTUBAYEVA, M., 2024. Enhancing spring barley grain yield with local biofertilizers in the semi-arid steppe zone of Northern Kazakhstan. International Journal of Design & Nature and Ecodynamics, vol. 19, no. 2, pp. 371-378. http://doi.org/10.18280/ijdne.190202
» http://doi.org/10.18280/ijdne.190202 -
NURGAZIYEV, R., IRMULATOV, B., NASIYEV, B., SIMIC, A., ZHANATALAPOV, N., BEKKALIYEV, A., KHIYASOV, M. and AIDARBEKOVA, T., 2024. Influence of organic fertilizers on the restoration of the biological resource potential of natural degraded pastures in the steppe zone of Northern Kazakhstan. Online Journal of Biological Sciences, vol. 24, no. 4, pp. 848-857. http://doi.org/10.3844/ojbsci.2024.848.857
» http://doi.org/10.3844/ojbsci.2024.848.857 - SOLOMATIN, A.O., 1973. Kulan Moscow: Nauka.
-
XU, W., XIA, C., YANG, W., BLANK, D., QIAO, J. and LIU, W., 2012. Seasonal diet of Khulan (Equidae) in Northern Xinjiang, China. The Italian Journal of Zoology, vol. 79, no. 1, pp. 92-99. http://doi.org/10.1080/11250003.2011.620635
» http://doi.org/10.1080/11250003.2011.620635 -
YASYNETSKA, N., 2023. Craniological characteristics of turkmenian Kulan Equus hemionus kulan Groves and Mazak, 1967 of the local population of the biosphere reserve “Askania Nova”. Biosphere Reserve "Askania Nova" Reports, vol. 25, pp. 164-176. http://doi.org/10.53904/1682-2374/2023-25/19
» http://doi.org/10.53904/1682-2374/2023-25/19
Edited by
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Editor:
Takako Matsumura Tundisi
