Abstract

It is well known that the earth's climate changes that occur as a result of global climate
change are affecting the biosphere of our planet as a whole. As a result, many animal
species change their geographical ranges, seasonal activity, migration patterns, population
and interaction with other species, including microorganisms, the facts of which are
observed in the deserts of Kazakhstan. The main vector of the plague-causing agent in the
desert natural focus of the Betpak-Dal plague in Kazakhstan is Great gerbil, where the first
plague epizootic was discovered in 1959, whose initial area of this focus was 30000 km2
until 1990. Due to rapid industrial development in the Betpak-Dala desert, the impact of
global climate change, and other factors over the past 30 years, the area of this natural
plague outbreak has increased by 50.12% (60.14 thousand km2) in the northern direction.
This expansion resulted from the biological transformation of the main carrier's habitat,
leading to the increased prevalence of plague among rodents. At the same time, the plague
microbe was also involved in the transformation process, increasing its natural area of the
pistil (the area of distribution of the microbe), thereby elevating the risk of human infection
with the plague.

The paper presents the results of our epizootiological monitoring from 2009 to 2022,
covering an area of 50,000 km2 and laboratory studies of 522,376 samples of field material
for plague. The paper includes the structure of the focus and the reasons for the expansion
of the settlement area of the rodent population and also identification of sectors with
varying levels of epidemic danger and epizootic activity, and an analysis of the main
phenotypic and genotypic properties of Yersinia pestis strains circulating in this natural
plague focus.

Keywords: plague, plague foci, area of focus, Rhombomys opimus, biotransformation,

epizootological monitoring.

Introduction
Plague is an acute natural focal infectious disease of a group of quarantine infections,
occurring with an exceptionally severe general condition, fever, damage to lymph nodes,
lungs, and other internal organs, often with the development of sepsis. In the past, the
disease was characterized by a high, almost 100% lethality and very high contagiousness.
The causative agent of the infection is the plague bacillus (Yersinia pestis). In natural foci of
the plague, the sources and reservoirs of the causative agent of infection are mainly rodents
– marmots, ground squirrels, and gerbils, mouse-like rodents, as well as hares. Carriers of
the causative agent of infection are fleas of 55 different species (Source: The Plague, 2023).

Yersinia pestis circulates in animal reservoirs, particularly in rodents, in the natural foci of
infection found on all continents except Australia. The natural foci of plague are situated in a
broad belt in the tropical and sub-tropical latitudes and the warmer parts of the temperate
latitudes around the globe, between the parallels 55° N and 40° S (Plague Manual:
Epidemiology, Distribution, Surveillance and Control, 2023).
Various types of natural plague foci (desert, steppe, high-altitude, and mixed) on the
territory of Kazakhstan occupy 1083.9 thousand square kilometers, which is about 40% of
the territory, including cross-border high-altitude plague foci (Kyrgyzstan, Mongolia, China,
Russia), where outbreaks of plague among the population have repeatedly occurred in the
past (Abdeliyev et al., 2022; Atshabar et al., 2015). The studied desert plague focus of
Betpak-Dal (Betpak-Dal autonomous plague center) is part of the Central Asian desert
natural plague focus of Kazakhstan, where the main carrier is a great gerbil (Rhombomys
opimus).

Originally, the Betpak Dala desert was used as a seasonal pasture by livestock breeders
keeping cattle to the south, in the Moyinkum desert and on the foothill plains. Cattle tracks
passed through Betpak Dala, along which cattle were driven north in the spring to the vast
steppes of Saryark, south of the Kazakh low-rise area and back to the sands of Minkum in
the autumn. Thanks to the millions of herds of saiga, in the past, a hunting industry
developed here. Since the 70s of the last century, mining of uranium, gold, and other
minerals began in the Betpak Dala desert. However, against the background of global
warming, anthropogenic impact, or partial aridization in some parts of the territory, the
great gerbil population is progressing due to the coverage of the territories of Ulytau and
Karaganda regions of Kazakhstan beyond the 47th parallel of northern latitude. The active
study of rodents in the Betpak Dala desert began in the 30-60s of the XX century and
continues to the present time, where a large amount of material was collected (Kashkarov,
1935; Beloslyudov, 1948; Ismagilov, 1961; Ismagilov, 1974; Burdelov, 1974; Burdelov, 1977;
Dubrovsky et al., 1980; Shubin, 1981; Rapoport and Rakhimov, 2006; Kulemin et al., 2017;
Sazhnev et al., 2008; Rapoport, 2013; Sazhnev et al., 2013; Sazhnev et al., 2016; Sazhnev et
al., 2018; Abdel et al., 2021)."
Unfortunately, to date, there is insufficient data on the settlements of the great gerbil on a
cartographic basis in Betpakdal, and, due to a various reasons, the distribution of the great
gerbil on the territory is devoid of detalisation (Ismagilov, 1961, Dubrovsky et al., 1980,
Sazhnev et al., 2008). Therefore, our task was to concretize all known gerbil settlements by
2022 with a brief description of them within the western part of the Betpak Dala desert of
Kazakhstan.

Thus, the significant persistence and transformation of natural plague foci, the variety of
mechanisms and ways of transmission of the causative agent of infection, the spread of the
pathogen by carriers in nature, the involvement in its circulation of a large number of warm-
blooded animals and arthropods, an increase in the risk of human infection with plague
determine the importance of improving monitoring of natural foci. Delineation of the
modern range of the great gerbil (Rhombomys opimus) is the basis of epizootiological
examination and is necessary to clarify the boundaries of the natural focus of the plague in
order to develop effective monitoring and preventive measures to ensure biological safety
and epidemic well-being of the country's population as a whole.

Materials and Methods

Epizootiological monitoring and collection of field material

The research was carried out according to generally accepted methods of epizootiological
monitoring (Stepanov et al., 1990; Smolensky et al., 2019) using modern tools of the GPS
system and GIS technology. The experimental base of the study was the Shymkent Anti-
Plague Station branch and the Central Reference Laboratory of the Masgut Aikimbayev’s
National Scientific Center for Especially Dangerous Infections of the Ministry of Health of the
Republic of Kazakhstan (NSCEDI, MoH of the RK).

Field and laboratory studies were carried out on the basis of the Resolution of the Chief
State Sanitary Doctor of the Ministry of Health of the Republic of Kazakhstan “On carrying
out sanitary and anti-epidemic and sanitary-preventive measures on the enzootic plague
territory of the Republic of Kazakhstan for 2021-2025” No 8 from 26.02.2021, in accordance
with Article 36 of the Code of the Republic of Kazakhstan “On public health and healthcare
system,” taking into account the requirements of the International Health Regulations
(2005) in order to ensure epidemiological well-being for plague on the territory of the
Republic of Kazakhstan and approved by the Bioethics Committee of the NSCEDI of the MoH
of the RK.

The materials are based on accumulated long-term observations (2009-2022) in the process
of epizootiological examination within the Turkestan region (Sozak district) and Ulytau
region (Ulytau district) of Kazakhstan. The survey area in the South passed along the Shu
River, in the North to the Zhideli sands. The length was about 300 km. In the West, the
border was the Sarysu River, in the East, the survey area reached the border with the
Zhambyl region of Kazakhstan, which was about 200 km from West to East. The survey area
was more than 50.0 thousand km2. In total, 36,203 mammals and 486,173 ectoparasites
were studied during the analyzed period (total – 522,376 samples).

Study of the properties of isolated strains

To study phenotypic and molecular genetic studies (Polymerase Chain Reaction, PCR) of this,
49 strains of Yersinia pestis were used, isolated from the Betpak Dala autonomous plague
focus of Kazakhstan obtained from the depository and museum of living cultures of the
NSCEDI. All manipulations with Y. pestis strains were carried out in accordance with
biological safety standards (Nekrasova et al., 2012) and techniques for working with
pathogens (Nekrasova et al., 2001). DNA extraction of Y. pestis strains was carried out using
the QIAamp DNA Mini Kit (Qiagen, USA) (Halos et al., 2004; Billeter et al., 2011). Genotyping
of Y. pestis was carried out using whole genome sequencing. DNA sample preparation was
performed using the Nextera XT DNA Library Preparation Kit (catalog number: FC-131-1024),
according to the manufacturer's instructions. Sequencing was performed on a high-
performance MiSeq sequencer, Illumina platform, and a set of chemical reagents MiSeq
Reagent Kit v3, 600 Cycles (catalog number: MS-102-3003) according to the manufacturers'
instructions.

Research Quality Control

Quality control was ensured by testing control strains deposited in the museum of living
cultures of the NSCEDI: reference strains of Y. pestis from various autonomous foci of
Kazakhstan, Y. pestis EV strain, Y. pseudotuberculosis strain, and fragments of loci of four
well-studied strains representing the main biovars of the plague microbe were used as
reference samples: Pestoides F (biovar Microtus/Antiqua), Nepal516 (biovar Antiqua),
KIM10+ (biovar Mediaevalis), and CO92 (biovar Orientalis).
Results
General Characteristics of the Desert Zone
Betpak Dala (Kaz. betpak – unscrupulous, shameless, fierce, turk. batnak – swampy,
turkic/kaz. dala – plain) is a desert in the Ulytau, Karaganda, Turkestan, and Zhambyl regions
of Kazakhstan. It is located between the lower reaches of the Sarysu River, the Chu River,
and Lake Balkhash. To the north, it borders the Kazakh small hills. Betpakdala belongs to the
gravelly-clayey and clayey-salt deserts. The area is approximately 75,000 km², with a length
from west to east of about 500 km and from north to south of up to 170 km (Betpak-Dala,
2023).
Geographical Characteristics and Climatic Conditions of the Landscape
The Betpak Dala Desert is a flat and gently undulating plain with average heights of 300-350
m and a general slope to the southwest. In the east, the elevations are greater, and in the
southeast, there is the Zheltau elevation up to 974 m (Zhambyl city). The western part of
Betpakdala is composed of folded Mesozoic and horizontally lying Paleogene loose rocks
(sands, sandstones, clays, pebbles), while the eastern part has a folded structure and is
composed of lower Paleozoic sedimentary-metamorphic strata and granites. Desert gray-
brown saline and solonetzic soils predominate. The climate is sharply continental, with
precipitation averaging 100-150 mm per year, of which only 15% occurs in summer. The
region experiences sharp contrasts between winter and summer, with dry and hot summers
and moderately cold winters with little snow. The average temperature in January is -13-
14°C, while in July, it is +23.0-24.5°C. The sum of temperatures above 10°C is 3100-3400°C,
the hydrothermal coefficient of moisture availability is 0.2-0.3, and the duration of the
period with air temperatures above 10°C is 170-177 days. In winter, absolute minimum
temperatures reach –42 to –46°C.
A stable snow cover is formed in the first ten days of December and lasts 2.5-3 months, but
its height does not exceed 10-15 cm. Despite the large thermal resources, a significant lack
of moisture limits the cultivation of agricultural crops (Betpak-Dala, 2023).
Flora and fauna of the territory: A significant part of the territory is covered with thickets of
boyalych (Salsola arbuscula), in depressions replaced by wormwood-ephemeral and
solyanka formations (black wormwood, gray wormwood, etc.). Solyanka formations are
represented by boyalych (Xylosalsola arbuscula), biyurgin (Anabasis salsa), tasbiurgun
(Nanophyton erinaceum), kokpek (Atriplex cana), itsigek sarsazan (Halocnemum
strobilaceum (Pall.)), etc. There are many holophytes – soleros (Salicornia), solyanka (Salsola
tragus). Quite a lot of shrubs – tobylga (Filipendula vulgaris), karagany (Caragana), gingila
(Tamarix gallica), meadowsweet (Spiraeanthus), buzzard (Calligonum). Occasionally, in the
depressions, there are thickets of saxaul (Haloxylon), turangi (Populus euphratica), etc.
Ephemeral plants develop poorly in spring because, at a time when the soil is best soaked
due to the melting of seasonal snow and early spring precipitation, it does not have time to
warm up sufficiently yet. In spring, flowers such as tulips and onions develop, but ephemeral
vegetation does not form on a dense cover (Kashkarov, 1935; Beloslyudov, 1948; Ismagilov,
1961; Popova and Kutyrev, 2022).

The herpetofauna is represented by the following species: Central Asian tortoise

(Agrionemys horsfieldi), steppe agama (Agama sanguinolenta), swift lizard (Eremias velox),
arrow-snake (Psammophis lineolatum), and cottonmouth (Agkistrodon halys).

Among the four-legged predators found in the saxaul forests are the wolf (Canis lupus), fox
(Vulpes vulpes), corsac fox (Vulpes corsac), and the possible "badger dog" (Vormela
peregusna). Please verify the correct species name for the latter.
The Betpakdala desert plague focus belongs to the Central Asian natural plague focus of
Kazakhstan, with a focus code of 42. The total area of the focus is 60,140 km2, and no
epidemic manifestations of plague in a natural focus have been registered. This focus area is
divided into five landscape-epidemiological regions (LER): 42.1. Western Sholak-Espinsky
42.2. Southern Kamkalinsky 42.3. Eastern Akbakaysky 42.4. Central Hilly 42.5. Karakoinsky
The rodent fauna comprises 26 species, with the main carrier of plague in the focus being
the great gerbil (Rhombomys opimus, Licht, 1823). These gerbils are distributed
sporadically, but their individual settlements cover vast areas, with population densities
ranging from 400 to 1,000 gerbils per 1 sq. km on average. In some areas, their numbers
exceed 5,000 animals per 1 sq. km. The red-tailed gerbil populations are more evenly
distributed over a wide area, with similar population densities.
The fauna of fleas parasitizing rodents includes nine species: Xenopsylla gerbilli minax, X.
conformis, Nosopsyllus laeviceps, Citellophilus trispinus, Echidnophaga oschanini,
Mesopsylla hebes, Oropsylla ilovaiskyi, Neopsylla setosa, and Coptopsylla lamellifer. Flea
abundance indices on the great gerbil vary from 0.1 to 5.8, and in burrows, they range from
45 to 365 (Chulak-Espe level). The dominant ectoparasites are X. g. minax and X. conformis,
with populations ranging from 10,000 to 40,000 insects per 1 sq. km. In some areas, X.
skrjabini is also numerous.
The study of rodents in the Betpak Dala desert began in the 1930s and continued through
the 1970s. The first registered epizootic of plague in the Betpakdala focus occurred in 1959,
followed by sporadic occurrences in subsequent years. However, no human cases of plague
were registered during this period. Historical geographical names, such as Karaoba (Kaz.
kara – black; oba – plague), Kosoba (kos - double, couple, shack), and Besoba (bes – five),
indicate the presence of temporary nomadic settlements, pastures, tracts, and wells
associated with plague.
Due to the heterogeneous landscapes, vegetation, rodent settlements, and the unique
characteristics of the epizootic plague process, the Western part of Betpak Dala (42.1.
Western Sholak-Espinsky) was subdivided into three additional landscape-epizootiological
districts (LER) based on long-term observations: Sholakespinsky, Western Betpak Dala, and
Karakoyinsky.
In the study area, we identified four main types of great gerbil settlements: diffuse, band,
island, and solitary.
Figure 1 shows a detailed diagram of the settlement of the main plague carrier in the
western part of the autonomous plague focus of Betpakdala (Fig. 1a), indicating the total
occupied area (Fig. 1b) of the enzootic plague territory of Kazakhstan.
Due to the rapid industrial development of the Betpak Dala desert, global changes in
climatic conditions, and other factors over the past 30 years, the area of the natural focus
has increased by 50.12% (60.14 thousand km2), which has led to the expansion of plague
epizootics among wild rodents, increasing the risk of human infection with the plague (see
Figure 1b).
A large settlement of the great gerbil is located in the Inkuduk tract (the area of the Zhalpak
deposit), where it is represented in a diffuse form (Fig. 1d). The white spots in the picture
represent great gerbil colonies. The density of colonies ranges from 150 to 250 colonies, and
in some places reaches 300 colonies per 1 km2. Another settlement, but of an island type, is
located in the southeast of the described territory (Fig. 1c). Groups of colonies are outlined.
Here, the density ranges from 50 to 200 colonies per 1 km2. Ribbon settlements of the great
gerbil run along the road from the village of Zhuantobe to the village of Kyzemshek and
further to the turn to the Zhalpak deposit. They are found in areas along the Pavlodar-
Shymkent oil pipeline, the Beineu-Bozoy-Shymkent gas pipeline, highways, etc.
Due to its ecological plasticity, the great gerbil can adapt to various natural conditions and
has a significant advantage over other rodent species. The gerbil actively inhabits newly
emerging roads, embankments, and various pipelines, which are often associated with the
construction of new drilling sites and mining enterprises (Sazhnev et al., 2013).

Figure 1: Distribution area, types of settlements of the great gerbil (Rhombomys opimus) in
the western part of the autonomous plague focus Betpakdala of the Central Asian desert
plague focus of Kazakhstan: a – Settlements of the great gerbil in the territory of the Ulytau
and Turkestan regions of Kazakhstan.
b – General border of the Betpak Dala plague focus.
c – Island settlements of the great gerbil.
d – Diffuse settlement of the great gerbil.
Figures c-d are made based on images from the Google Earth Pro application in the public
domain.
Even during the reconstruction of roads, colonies destroyed by heavy road equipment
reappear within 2-3 years, and after a while there are already solid colonies, barely
distinguishable only by the ecological center. The density in ribbon settlements reaches 8 or
more colonies per 1 ha. There are also single colonies on the territory of the LER. They are
so rare that only 1-2 colonies can be found at a distance of 5-7 km. But, as a rule, such
colonies are large, 25-30 m in diameter, with a good ecological center, where at least 5-10,
and sometimes more than 15 animals live. The density of colonies in such settlements is
extremely difficult to determine.
The results of a long-term analysis showed a stable epizootic index, which depended on
many factors, such as the number of carriers and vectors, the epizootic cycle, the
persistence of the pathogen in the inter-epizootic period, and others.
Characteristics of plague microbe strains:
Based on cultural, morphological, biochemical, enzymatic properties, sensitivity to plague
and pseudotuberculosis phages, the strains isolated in the focus belong to the typical sandy
variant of the main subspecies Yersinia pestis subsp. Pestis (Popova and Kutyrev, 2022).
During the analyzed period (2009-2022), a total of 48 strains of Yersinia pestis were isolated
and studied, of which 26 from the great gerbil, 2 from the red-tailed gerbil, 20 from fleas,
and 1 from ticks. The microbial population of the plague microbe has typical properties
characteristic of the populations of the Central Asian desert focus of Kazakhstan. All strains
ferment glycerol, glucose, mannitol, maltose, arabinose and do not ferment rhamnose,
pesticinogenic and do not have denitrifying ability.

All strains produced fraction 1, which demonstrated high virulence for laboratory animals.
The LD50 for guinea pigs was 1.0×10^6 microbial cells (m.c.), and for white mice, it ranged
from 1.6-9.0×10^4 m.c. Furthermore, for white mice, the LD50 of arginine auxotrophs was
1.1 × 10^3 m.c. All strains of the plague microbe exhibited sensitivity to antibacterial drugs,
and no antibiotic-resistant or phage-resistant strains were found in the focus.
A comprehensive analysis of the results obtained by specialists from our scientific center
(Atshabar et al., 2021; Lowell et.al., 2007; Abdrasilova et.al., 2019; Abdel et.al., 2020; Meka-
Mechenko et.al., 2022) and our own molecular genetic studies, including genome-wide
sequencing of Y. pestis strains circulating in the autonomous plague center of Betpakdal in
Kazakhstan, led to their assignment to the Medieval 2.MED1 biovar.
Modern characteristics of the plague focus and differentiation according to the level of
epidemic danger and epizootic activity.
Based on the results of epizootiological monitoring and a comprehensive meta-analysis, the
territory of the plague focus was differentiated according to the level of epidemic danger
and epizootic activity. This assessment considered factors such as the extent and intensity of
the epizootic process of plague among great gerbils and their fleas, as well as the number of
isolated strains of the plague microbe and the registration of seropositive animals for plague
infection during the period 2009-2022 (see Figure 2).
During the study period, it was found that plague epizootics among wild animals in this
focus area were registered on an area of 15,600 km2 (26.0% of the total focus area of
60,140 km2) from 2009 to 2015. Subsequently, an inter-epizootic period began seven years
ago. Plague epizootics occurred in a localized form with low intensity in the southern and
central regions of the autonomous plague focus of Betpakdal in Kazakhstan. In the analyzed
period, a total of 48 strains of Y. pestis were isolated, including 26 from great gerbils, 2 from
red-tailed gerbils, 20 from fleas, and 1 from ticks.
It is well known that when conducting an epizootiological survey of natural plague foci and
their fragments within natural boundaries, the formal territorial principle of dividing the
earth's surface into georeferenced data, or into so-called formalized accounting units
(sectors and primary areas) is used. The area of the autonomous plague focus of Betpakdala
(60,140 km2) is divided into 550 sectors. When differentiating this focus according to the
criteria of levels of epidemic danger and epizootic activity, the breakdown was as follows:
very low – 287 sectors (51.2%), low - 146 sectors (26.5%), average – 75 sectors (13.6%), high
– 40 sectors (7.3%), very high – 2 sectors (0.36%).
An analysis of the population organization of the great gerbil and its neighboring
populations revealed that the peripheral population of this rodent is subject to continuous
variability, influenced by climatic, geographical, and anthropogenic factors. Under the
influence of these factors, a mosaic distribution (diffuse, island, and band settlement) of
populations of the great gerbil is observed in the peripheral zone of the foci. In some places,
the formation of a sparse population density of the main carrier is observed (Abdel et al.,
2021).
Natural sandworm foci of plague in Kazakhstan are characterized by high epizootic activity
and require constant monitoring. The significant persistence of natural plague foci, the
variety of mechanisms and routes of transmission of the infectious agent, the wide
distribution of the pathogen in nature, and the involvement of a large number of warm-
blooded animals and arthropods in its circulation determine the importance of improving
monitoring of these foci (Atshabar et al., 2022).

Figure 2: Results of Meta-Analysis of Epizootic Characteristics and Differentiation of the

Autonomous Focus of Betpak-Dala in Kazakhstan (2009-2022) for Plague:

 a - Total area of the autonomous plague focus and areas with epizootic
manifestations.
 b - Number of sectors with different levels of epidemic danger and epizootic activity.
  c - Results of the study on great gerbils and the number of isolated strains of the
plague microbe.
 d - Results of the study on fleas and the number of isolated strains of the plague
microbe.

Discussion:
Changes in the structure and functioning of natural plague foci in Kazakhstan are
continuously monitored by the Kazakhstan anti-plague service, utilizing GIS technology
tools. This monitoring has led to the development of new strategies and tactics for
epidemiological surveillance of plague and the adoption of new principles for planning and
implementing preventive measures. A crucial aspect of this effort is epizootiological
monitoring with landscape-epizootiological zoning following international standards. This
approach enables the consideration of survey results and the registration of the epizootic
process among wild animals in a differentiated manner, ensuring a unified approach to the
differentiation of the controlled territory.
The epizootic process is a complex, continuous phenomenon involving the emergence and
spread of infectious diseases through the transmission of pathogens from infected animals
to susceptible healthy ones. It is influenced by both natural and anthropogenic factors,
including human activities. For instance, human migrations can significantly impact the
development of the epizootic process, either activating it and increasing its epidemiological
potential or, conversely, isolating a territory or farm to prevent its development (Electronic
Agricultural Knowledge Library, 2023).

It was observed that the trend of expanding the boundaries of the foci occurred
concurrently with increased anthropogenic impact on natural landscapes, climate change
(global warming), and other factors. This observation provides ample reason to anticipate
the further expansion of the great gerbil's range.
From the mid-70s to the early 80s of the last century, industrial development in Western
Betpak Dala (referenced as 'History of Kazakhstan Section XVI, 2023') commenced and
continues to the present day. This development led to the establishment and expansion of
settlements such as Kyzemshek, Taikonyr, and Appak. The construction of the Pavlodar-
Shymkent oil pipeline traversed the entire territory, followed by the development of
deposits for various industrial ore mining enterprises. In December 2010, construction
commenced on the Beineu-Bozoi-Shymkent gas pipeline, designed to transport gas from
western Kazakhstan fields to supply the southern region with natural gas and for export via
the Kazakhstan-China gas pipeline. The total length of all roads, embankments, oil and water
pipelines in Western Betpak Dala now exceeds 600 kilometers. The bulk and ridge soil
underpinning these roads and pipelines are consistently less dense than the surrounding
areas, creating favorable conditions for burrow construction. Betpak Dala is known for the
production of coal, gypsum, uranium, lead and iron ores, quartzites, salt, rock crystal,
marble, and other valuable resources (references: Zhambyl Region Development Program
for 2011-2015, 2014; Kyzylorda Region Development Program for 2011-2015, 2013;
Sarkytkan and Isabekov, 2014; South Kazakhstan Region Development Program for 2016-
2020, 2016).
The territory of Betpak-Dala serves as spring and autumn pastures for cattle. It facilitates the
seasonal movement of livestock from the summer pastures of the semi-desert in the Kazakh
small hills (Sary-Arka steppe) to the winter pastures of the Chui Valley and the Moyinkum
sands, and vice versa. Additionally, there are substantial settlements in the region, including
Kyzemshek (population: 4,230) and Taikonyr (population: 2,000), which belong to the Sozak
district of the South Kazakhstan region.

Currently, approximately a dozen uranium mining enterprises operate on a rotational basis

in Western Betpak Dala, employing over 2,500 people. These rotational camps are
distributed across various regions of Kazakhstan and even neighboring countries, raising
concerns about the potential spread of plague infection if employees fall ill. Additionally,
Western Betpak Dala is home to sizable geological villages like Kyzemshek (population:
4,230) and Taikonyr (population: 2,000). In the eastern part of the region, there are
approximately 23,000 residents living in villages, railway stations, and crossings within the
Zhambyl and Karaganda regions. Enterprises for the extraction of barite, asbestos, marble,
gold, and more are located in the eastern part of the focus. Notably, these settlements are
in close proximity to colonies of great and red-tailed gerbils (references: Zhambyl Region
Development Program for 2011-2015, 2014; Kyzylorda Region Development Program for
2011-2015, 2013; Sarkytkan and Isabekov, 2014; South Kazakhstan Region Development
Program for 2016-2020, 2016).

It's important to note that in recent decades, significant global climate changes have
occurred, impacting not only the atmosphere but also the biosphere, hydrosphere, and
cryosphere. These changes have had noticeable effects on various aspects of human life
(South Kazakhstan Region Development Program for 2016-2020, 2016). According to the
linear trend of air temperature anomalies (against the base period of 1961-1990), there has
been a steady increase in air temperature in Kazakhstan from 1941 to 2015. These global
climate changes have resulted in alterations in the planet's biosphere as a whole (IPCC,
2015).
Scientific research on climate change and water resources in Central Asia has indicated that
climate change impacts will be observed across all regions of the planet, with Central Asia
being no exception. Central Asia has experienced intense climate warming (IPCC, 2015). As
evidence of this trend, Figure 3a presents the results of average annual temperature
estimates obtained from Berkeley Earth and five other research groups that analyze surface
temperature (2017 Global Temperature Report, Berkeley Earth).
This trend toward warmer winters and drier summers in many parts of the region has led to
the retreat of glaciers and permafrost melting in the Pamir and Tien Shan mountains.
Consequently, the risk of heavy precipitation, drought, floods, and mudflows has increased.
It is noteworthy that spring precipitation has decreased in the southern desert regions of
Central Asia while increasing in the northern steppe regions (IPCC, 2015). The northern
borders of the Aryskum-Dariyalyktakyr, Balkhash, and Betpakdal autonomous plague foci
are situated in the northern steppe regions.
The main negative impacts of climate change, assessed for their probability of manifestation
according to the Fifth Assessment Report of the IPCC, include the following: changes in
hydrological systems affecting water resources in terms of their quantity and quality
(medium confidence) are observed; many terrestrial, freshwater, and marine species have
altered their geographical ranges, seasonal activity, migration patterns, abundance, and
interaction with other species (high degree of reliability) (South Kazakhstan Region
Development Program for 2016-2020, 2016; IPCC, 2015).
As a result, the gradual development of rodents, particularly the great gerbil, continues not
only in the Betpakdalinsky autonomous plague foci of Kazakhstan but also in the other two
autonomous plague foci, namely Aryskum-Dariyalyktakyrsky and Balkhashsky.
Thus, these examples demonstrate the presence of numerous industrial enterprises and
zones with stationary buildings or modules within the plague-enzootic territory. Paved roads
and electric lines connect to all these enterprises, and there are numerous access roads and
pipelines, including water pipes, laid around the mining sites themselves.

The total length of roads, oil, and water pipelines is currently more than 600 km in Western
Betpakdal (Electronic Agricultural Knowledge Library, 2023). All members of biocenosis,
especially plant communities and rodents, respond rapidly to changes in indigenous
landscapes. Roads and embankments of water and oil pipelines are particularly attractive to
rodents due to the abundance of ephemeral plants and various shrubs along the
embankments and ditches. Even in dry years, the vegetation on these embankments
remains richer than on the surrounding plain. Additionally, the raised embankments of
roads and various pipelines provide protection from flooding with meltwater and rainwater.
The bulk and upland soil at the base of roads and pipelines is consistently less dense than in
the surrounding areas, making it favorable for the construction of rodent burrows (see
Figure 3b, 3c, and 3d).

Figure 3: Climatic Influences and Anthropogenic Impacts on Natural Plague Foci of

Kazakhstan: a - Results of the assessment of the average annual temperature in Central Asia,
Berkeley Earth, 2017; b - Refining workshop of a uranium mining plant in Western Betpak
Dala, South Kazakhstan Region, 2017; c - Satellite image (Google, 2018) of the working
village of Kyzemshek (Steppe) showing a dense network of roads, pipelines, and great gerbil
colonies in Western Betpak Dala, South Kazakhstan; d - Linear settlements of the great
gerbil along the highway in West Betpak Dala, South Kazakhstan (Turkestan region, 2019).
It's worth noting that the concentration of rodents along highways and other anthropogenic
elevations leads to a higher number of blood-sucking ectoparasites, such as ticks and fleas,
which can act as carriers of various pathogens of infectious diseases. Discussions related to
the transformation of the borders of autonomous plague foci, increasing or decreasing the
area of the main carrier, and the emergence of new focal areas for plague in Kazakhstan
have been held in the academic council of our scientific center and the scientific and
production councils of the country's anti-plague services. Decisions reached during these
discussions have been published in the form of articles in domestic and foreign periodicals
and atlases.
Conclusion: As of January 1, 1990, the total area of the Betpak Dala plague focus in
Kazakhstan was 30,000 km2. The northern part of the focus did not extend beyond the 46th
parallel of Kazakhstan's territory. However, in the context of global warming, anthropogenic
impact, and partial aridization in other regions, the great gerbil population has expanded,
encompassing areas of the Karaganda region beyond the 47th parallel of northern latitude.
Considering the Balkhash zone, this expansion may exceed the mentioned value. Over the
past 30 years, the area inhabited by the great gerbil and, consequently, the plague-enzootic
territory, has increased by 50.12%, equivalent to 60.14 thousand square kilometers. This
expansion has been confirmed through the isolation of the plague-causing agent and the
detection of seropositive animals for plague infection.
An analysis of the population dynamics of the great gerbil and its neighboring populations
reveals that the peripheral settlements of these rodents are subject to permanent
variability, influenced by climatogeographic and anthropogenic factors.

The results were obtained within the framework of the program-targeted financing of the
NTP: 'Development and scientific justification of public health technologies, biological safety
for the impact on the prevention of particularly dangerous infectious diseases' (BR11065207
– IRN program), with funding provided by the Ministry of Health of the Republic of
Kazakhstan. Additionally, we received support from the Science Committee of the Ministry
of Science and Higher Education of the Republic of Kazakhstan for the project titled 'Study of
the genetic diversity of plague strains to create a biorepository and determine the genesis of
enzootic plague nuclei in natural foci of Kazakhstan' (IRN AR09261078)

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Information about the authors

Ziyat Zh. ABDEL, a visiting researcher at the Plague Laboratory of the National scientific
center of especially dangerous infections named after M. Aikimbayev, Candidate of Medical
Sciences, Associate Professor, e-mail: ziyatabdel@gmail.com, https://orcid.org/0000-0002-
2738-6818
Tatyana V. MEKA-MECHENKO, chief Researcher of the Plague Laboratory of the National
scientific center of especially dangerous infections named after M. Aikimbayev, Doctor of
Medical Sciences, Associate Professor, e-mail: tmekamechenko@gmail.com,
https://orcid.org/0000-0002-6322-0065
Zauresh B. ZHUMADILOVA, General Director of the National scientific center of especially
dangerous infections named after M. Aikimbayev, Candidate of Medical Sciences, Professor,
e-mail: zzb.68@mail.ru, https://orcid.org/0000-0002-4161-1751
Nurbol N. SHAKIEV, research associate of the department of epizootology of particularly
dangerous infections with the museum and insectarium of the National scientific center of
especially dangerous infections named after M. Aikimbayev, master of natural sciences, e-
mail: nurbol.shakiy@gmail.com, https://orcid.org/0000-0002-5644-2882
Duman T. YESSIMSEIT, head of the Reference Laboratory of the National scientific center of
especially dangerous infections named after M. Aikimbayev, Master of Veterinary, e-mail:
yessimseit@mail.ru, https://orcid.org/0000-0003-2202-9333
Beck Z. ABDELIYEV, microbiologist at the Reference Laboratory of the National scientific
center of especially dangerous infections named after M. Aikimbayev, PhD student in
molecular biology and genetics, e-mail: abdelbeck@gmail.com, https://orcid.org/0000-
0002-4184-6227
Raikhan S. MUSSAGALIYEVA, epidemiologist of the Department of Advisory and
Methodological Assistance of the National scientific center of especially dangerous
infections named after M. Aikimbayev, Candidate of Medical Sciences, Associate Professor,
e-mail: raikhansafar@gmail.com, https://orcid.org/0000-0002-6838-2338
Aigul A. ABDIRASSILOVA, epidemiologist of the reference laboratory of the M. Aikimbayev
National Scientific Center for Particularly Dangerous Infections, Candidate of Medical
Sciences, e-mail: aigul.abdirassilova@mail.ru, https://orcid.org/0000-0001-7308-2113
Svetlana B. ISSAEVA, Director of the branch the Aral Sea Anti-Plague Station of the National
scientific center of especially dangerous infections named after M. Aikimbayev, Candidate of
Medical Sciences, e-mail: s.isaeva64@mail.ru, https://orcid.org/0000-0001-9587-4445
Veronika P. SADOVSKAYA, head of the GIS technology laboratory National scientific center
of especially dangerous infections named after M. Aikimbayev, e-mail: versad@mail.ru,
https://orcid.org/0000-0001-8389-9362
Zaure Z. SAYAKOVA, chief of the department of epizootology of particularly dangerous
infections with the museum and insectarium of the National scientific center of especially
dangerous infections named after M. Aikimbayev, candidate of biological sciences, e-mail:
zzsayakova@mail.ru, https://orcid.org/0000-0003-1107-6345
Maxim V. KULEMIN, Head of Epizootology Laboratory of the branch the Shymkent Anti-
Plague Station of the National scientific center of especially dangerous infections named
after M.Aikimbayev, e-mail: kmaxim.75@mail.ru; https://orcid.org/0000-0002-8839-9276