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Ecology of Yersinia pestis and the Epidemiology of Plague

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Yersinia pestis: Retrospective and Perspective

Part of the book series: Advances in Experimental Medicine and Biology ((AEMB,volume 918))

Abstract

This chapter summarizes information about the natural foci of plague in the world. We describe the location, main hosts, and vectors of Yersinia pestis. The ecological features of the hosts and vectors of plague are listed, including predators – birds and mammals and their role in the epizootic. The epizootic process in plague and the factors affecting the dynamics of epizootic activity of natural foci of Y. pestis are described in detail. The mathematical models of the epizootic process in plague and predictive models are briefly described. The most comprehensive list of the hosts and vectors of Y. pestis in the world is presented as well.

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Author information

Authors and Affiliations

  1. Stavropol Research Anti-Plague Institute, Stavropol, Russian Federation

    Vladimir M. Dubyanskiy

  2. M. Aikimbayev Kazakh Scientific Center of Quarantine and Zoonotic Diseases, Almaty, Kazakhstan

    Aidyn B. Yeszhanov

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  1. Vladimir M. Dubyanskiy
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Correspondence to Vladimir M. Dubyanskiy .

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Editors and Affiliations

  1. Beijing Institute of Microbiology and Epidemiology, Beijing, China

    Ruifu Yang

  2. State Research Center for Applied Microbiology, Moscow Region, Russia

    Andrey Anisimov

Supplementary Details for “5.1.2 Distribution of Plague Foci”

Supplementary Details for “5.1.2 Distribution of Plague Foci”

5.1.1 5.1.2.1 Africa

5.1.1.1 5.1.2.1.1 North Africa (Group Countries)

  • Natural plague foci in North Africa have hardly been studied, although cases of plague in humans and camels in Libya, Tunisia, and Egypt have been described.

  • Hosts: Acomys cahirinus. The plague microbe was isolated in 1927–1928 [107].

Vectors: Verified data are lacking.

5.1.1.2 5.1.2.1.2 Mauritania, Western Sahara

  • Hosts: Psammomys obesus, with an ecology similar to that of Rhombomys opimus, Xerus erythropus, Mus musculus, Acomys cahirinus, Meriones libycus, Gerbillus pyramidum, G. gerbillus, G. agag, G. nanus. Dipodillus campestris, Pachyuromys duprasi, and Jaculus jaculus [108, 109]

  • Vectors: Xenopsylla ramesis, Synosternus cleopatrae, X. nubica [1]

5.1.1.3 5.1.2.1.3 Algeria

  • Hosts: Rattus rattus, Rattus norvegicus, gerbils of the genus Gerbillus and Meriones

  • Vectors: Xenopsylla cheopis

5.1.1.4 5.1.2.1.4 Democratic Republic of the Congo

  • Plague foci along the border with Tanzania:

  • Landscapes: Savanna

  • Hosts: Mastomys natalensis, Rattus rattus

  • Vectors: Xenopsylla brasiliensis, X. cheopis [30]

5.1.1.5 5.1.2.1.5 Uganda

  • Northern and western provinces:

  • Landscapes: The available literature lacks relevant information.

  • Hosts: Mastomys natalensis, Rattus rattus

  • Vectors: Xenopsylla cheopis, Ctenocephalides spp. [30, 110]

5.1.1.6 5.1.2.1.6 Kenya

  • Landscapes: The available literature lacks relevant information.

  • Hosts: Arvicanthis niloticus, Mastomys natalensis

  • Vectors: Xenopsylla cheopis, X. brasiliensis [30, 111]

5.1.1.7 5.1.2.1.7 Tanzania

  • The slopes of Mount Kilimanjaro and the area southeast of Lake Victoria:

  • Landscapes: 900–2500 m above sea level, forests, pastures

  • Hosts: Rattus rattus alexandrinus, Tatera robusta (Cretzschmar)

  • Vectors: Ctenophthalmus eximius, Nosopsyllus incisus [20, 30, 112, 113]

  • Epizootic area west of Lake Victoria borders plague foci in the Democratic Republic of the Congo:

  • Landscapes: The available literature lacks relevant information.

  • The southeast epizootic area borders the plague focus in Kenya:

  • Landscapes: The available literature lacks relevant information.

  • Hosts: Lemniscomys griselda Thom., Lemniscomys striatus L. [20]

  • Both the area in the vicinity of Lake Rukk and the interstream area between the Greater Rvakh and Ruhundzhi share a focus with that in Malawi and Zambia:

  • Landscapes: The available literature lacks relevant information.

  • Hosts (for all foci in Tanzania): Rattus rattus alexandrines, Tatera robusta (Cretzschmar), Pelomys fallax (Peters), Lemniscomys griselda (Thomas), Lemniscomys striatus [19]

5.1.1.8 5.1.2.1.8 Namibia

  • Foci in the interstream area between Kunene and Okavango, on the Angola border:

  • Landscapes: Desert savanna

  • Hosts: Tatera schinzi, Desmodillus auricularis [30]

  • Landscapes: The available literature lacks relevant information.

  • Foci on the southern half of the Damaraland Plateau and the southern slopes of the Kamala Plateau:

  • Landscapes: The available literature lacks relevant information.

  • Hosts: Tatera schinzi, Desmodillus auricularis

  • Vectors (for all foci in Namibia): X. philoxera, X. brasiliensis, and Dinopsyllus ellobius [30]

  • Foci on the Namaland Plateau

  • Landscapes: Sandy desert

  • Hosts: Tatera schinzi, Desmodillus auricularis [30]

  • Vectors: The available literature lacks relevant information.

5.1.1.9 5.1.2.1.9 Botswana

  • Foci east and west of the Okavango Delta and on the borders of Botswana, Namibia, and southern Angola:

  • Landscapes: Desert

  • Hosts: Tatera schinzi, Desmodillus auricularis [30]

  • Vectors: The available literature lacks relevant information.

  • Foci are also present in the south, just outside South Africa.

5.1.1.10 5.1.2.1.10 Malawi

  • Vectors: The available literature lacks relevant information.

5.1.1.11 5.1.2.1.11 Zambia

  • Focus of plague in eastern Zambia within the plateau; a single focus in Tanzania:

  • Hosts: Tatera valida

  • Vectors: Xenopsylla philoxera, X. hipponax [30]

5.1.1.12 5.1.2.1.12 Zimbabwe

  • Foci in most parts of the country:

  • Landscapes: Desert, savanna

  • Hosts: Tatera leucogaster, Aethomys chrysophilus [30]

  • Vectors: Xenopsylla philoxera, X. brasiliensis [114]

5.1.1.13 5.1.2.1.13 South Africa

5.1.1.14 5.1.2.1.14 Madagascar

  • Focal areas on the high plateau:

  • Landscapes: Madagascar savanna

  • Hosts: Rattus rattus

  • Vectors: Synopsylla fonquernii, Xenopsylla cheopis [30]

5.1.2 5.1.2.2 Southeast Asia

5.1.2.1 5.1.1.2.1 India

  • In northern India, foci on the Indo-Gangetic Plain:

  • Landscapes: The available literature lacks relevant information.

  • Hosts: Tatera indica, Bandicota indica

  • Vectors: Xenopsylla astia, X. cheopis [30]

  • In central India, foci in Madhya Pradesh:

  • Vectors: The available literature lacks relevant information.

  • Hosts: Tatera indica, Bandicota indica

  • Vectors: Xenopsylla astia, X. cheopis [30]

  • Foci in southern India:

  • Vectors: The available literature lacks relevant information.

  • Hosts: Tatera indica, Bandicota indica

  • Vectors: Xenopsylla astia, X. cheopis [30]

5.1.2.2 5.1.1.2.2 Nepal

  • Vectors: The available literature lacks relevant information.

  • Hosts: Rattus rattus

  • Vectors: Xenopsylla cheopis [115]

5.1.2.3 5.1.1.2.3 Burma

  • Foci in the central uplands, between the Irrawaddy and Sittwe Rivers:

  • Vectors: The available literature lacks relevant information.

  • Hosts: Rattus rattus, Bandicota bengalensis

  • Vectors: Xenopsylla cheopis [116]

5.1.2.4 5.1.1.2.4 Thailand

  • Foci on the Khorat Plateau and in the northern and western parts of central Thailand:

  • Vectors: The available literature lacks relevant information.

  • Hosts: Rattus concolor, R. rattus

  • Vectors: Xenopsylla cheopis [30]

5.1.2.5 5.1.1.2.5 Vietnam

  • Landscapes: Coastal lowland plains

  • Hosts: Rattus rattus, R. norvegicus, R. indica

  • Vectors: Xenopsylla cheopis [30]

5.1.2.6 5.1.1.2.6 Indonesia

  • Foci on Java

  • Landscapes: Hillsides, highlands, plateaus with rice fields

  • Hosts: Rattus concolor, R. norvegicus

  • Vectors: Xenopsylla cheopis, Stivalius cognatus [30]

5.1.3 5.1.2.3 Countries of the Former USSR (Part of English Foci Names by Anisimov et al. [117], 2015; Zhou et al. [118])

5.1.3.1 5.1.2.3.1 Pre-Araks

  • Natural foci (07) (it is the registration number of focus according to the Russian system of count) in the low mountains:

  • Landscapes: Deserts, low mountains, midlands

  • Hosts: Meriones vinogradovi, M. persicus, M. tristrami, M. dahli, Allactaga elater, Microtus socialis, Mus musculus [119]

  • Vectors: Xenopsylla conformis conformis, Nosopsyllus iranus iranus, Pulex irritans [15]

5.1.3.2 5.1.2.3.2 Transcaucasian Valleys and Foothills

  • Natural foci in the Kur-Arak lowlands:

  • Landscapes: Dry steppes, semideserts, deserts

  • Hosts: Meriones libycus, M. tristrami, Mus musculus [15]

  • Vectors: Xenopsylla conformis conformis, Nosopsyllus laeviceps laeviceps [15]

5.1.3.3 5.1.2.3.3 Transcaucasian Highland

  • Group of autonomous natural foci (04–06):

  • Landscapes: Mountainous steppe, steppe, subalpine and alpine meadows

  • Hosts: Microtus arvalis, Microtus socialis, Mus musculus [25]

  • Vectors: Callopsylla caspia, Nosopsyllus consimilis [15]

5.1.3.4 5.1.2.3.4 Dagestan Highland

  • Natural focus (39) located at 2,000–3,000 m above sea level:

  • Landscapes: Alpine meadows

  • Hosts: Microtus arvalis, Cricetulus migratorius, Arvicola terrestris [112, 121]

  • Vectors: Callopsylla caspia, Amalaraeus dissimilis dagestanicus [15, 122]

5.1.3.5 5.1.2.3.5 Central Caucasian

  • Natural focus of plague (1) at Mount Elbrus:

  • Landscapes: Mountain steppes, alpine and subalpine meadows

  • Hosts: Spermophilus musicus, Microtus arvalis, Sylvaemus uralensis, Mus musculus [15, 119]

  • Vectors: Citellophilus tesquorum caucasicus , Neopsylla setosa [15, 123]

  • Dagestan valleys and foothills: Natural foci of plague (03) in the Caspian Plain within Dagestan

  • Landscapes: Deserts, semideserts

  • Hosts: Spermophilus pygmaeus, Meriones tamariscinus, Mus musculus [15, 124]

  • Vectors: Neopsylla setosa, Citellophilus tesquorum caucasicus [15]

  • Terek-Sunzha natural focus of plague (02) near the Terek and Sunzha Rivers:

  • Landscapes: Desert steppes

  • Hosts: Spermophilus pygmaeus [15, 119]

  • Vectors: Neopsylla setosa, Citellophilus tesquorum [125]

  • 5.1.2.3.6 Pre-Caspian sandy natural foci (43) in sandy regions near the Caspian Sea in the Republics of Kalmykia, Dagestan, and Stavropol:

  • Landscapes: Semideserts

  • Hosts: Meriones meridianus, Spermophilus pygmaeus [126]

  • Vectors: Nosopsyllus laeviceps, Xenopsylla conformis conformis [127]

  • 5.1.2.3.7 Pre-Caspian northwestern natural foci of plague (14) in the Astrakhan region, Kalmykia, Northwest Caspian steppe:

  • Landscapes: Semideserts

  • Hosts: Spermophilus pygmaeus, Meriones meridianus, Mus musculus [119]

  • Vectors: Citellophilus tesquorum, Neopsilla setosa [1]

  • 5.1.2.3.8 Volga-Ural steppe natural foci of plague (15) in the northern Volga and Ural interstream, from the Caspian lowland to the Common Syrt:

  • Landscapes: Steppe, semideserts

  • Hosts: Spermophilus pygmaeus pygmaeus [128]

  • Vectors: Neopsilla setosa, Citellophilus tesquorum transvolgensis [128]

  • 5.1.2.3.9 Volga-Ural sandy natural foci of plague (16) in the southern part of the Caspian Depression, in the Ural and the Volga interstream

  • Landscapes: Sandy

  • Hosts: M. meridianus, M. tamariscinus, Spermophilus fulvus, Mus musculus

  • Vectors: X. conformis, Nosopsyllus laeviceps [128]

  • 5.1.2.3.10 Trans-UralTrans-Ural natural foci of plague (17) east of the Ural River in West Kazakhstan, Aktobe and Atyrau regions

  • Landscapes: Steppes, semideserts

  • Hosts: S. pygmaeus, Rhombomys opimus [128]

  • Vectors: N. setosa, C. tesquorum [128]

5.1.3.6 5.1.2.3.11 Central Asian Desert

  • 5.1.2.3.11.1 Ural-Emba autonomous plague focus (18) in the Ural-Emba desert between the Ural and Emba Rivers

  • Landscapes: Semideserts, river floodplains

  • Hosts: Rhombomys opimus, M. tamariscinus, M. meridianus [128]

  • Vectors: X. skrjabini, N. laeviceps, C. lamellifer [128]

  • 5.1.2.3.11.2 Pre-Ustyurt autonomous desert plague foci (19) between the Emba River, Ustyurt Plateau, and Caspian Sea

  • Landscapes: Semideserts, deserts

  • Hosts: Rhombomys opimus, M. lybicus, M. meridianus [128]

  • Vectors: X. skrjabini, C. lamellifer, N. laeviceps [128]

  • 5.1.2.3.11.3 Mangyshlak autonomous plague focus (23) in the Mangyshlak Desert and in Mangistau region covering an area of the Buzachi Peninsula that includes the plains and mountains of the Mangyshlak region

  • Landscapes: Deserts

  • Hosts: Rhombomys opimus, M. lybicus

  • Vectors: X. skrjabini, X. nuttalii [128]

  • 5.1.2.3.11.4 Ustyurt autonomous plague focus (20) on the Ustyurt Plateau of the Ustyurt Desert between the Mangyshlak region to the west and the Aral Sea to the east in the territories of the Republics of Kazakhstan and Uzbekistan

  • Landscapes: Deserts

  • Hosts: Rhombomys opimus, C. migratorius [128]

  • Vectors: X. skrjabini, X. nuttalii, X. g. caspica [128]

  • 5.1.2.3.11.5 North-Pre-Aral autonomous plague focus (21) in the northwest pre-Aral region including Large and Small Badger Sands in the North Pre-Aral Desert

  • Landscapes: Deserts

  • Hosts: Rhombomys opimus, M. lybicus, M. meridianus [128]

  • Vectors: X. skrjabini, C. lamellifer, N. laeviceps [128]

  • 5.1.2.3.11.6 Pre-Aral-Karakum autonomous focus (24) in the Pre-Aral Karakum Desert and the Aktobe, Kyzylorda, and Karaganda regions of Kazakhstan northeast of the Aral Sea

  • Landscapes: Sandy, clayey, gravelly deserts

  • Hosts: Rhombomys opimus, M. lybicus, M. meridianus

  • Vectors: X. skrjabini, C. lamellifer, N. laeviceps [128]

  • 5.1.2.3.11.7 Trans-Aral autonomous focus (22) in the Aryskum-Daryalyktakyr Desert and theKyzylorda and Karaganda regions of Kazakhstan

  • Landscapes: Sandy deserts

  • Hosts: Rhombomys opimus, M. lybicus, M. meridianus

  • Vectors: X. gerbilli, X. skrjabini [128]

  • 5.1.2.3.11.8 Karakum autonomous focus (25) in Karakum Desert, Turkmenistan

  • Landscapes: Sandy deserts

  • Hosts: Rhombomys opimus, M. lybicus, M. meridianus

  • Vectors: X. gerbilli, X. hirtipes [1]

  • 5.1.2.3.11.9 Kopet Dag autonomous desert focus (26) in Southwestern Kopet Dag

  • Landscapes: Desertified lowlands

  • Hosts: Rhombomys opimus, M. lybicus

  • Vectors: X. conformis, N. laeviceps [1]

  • 5.1.2.3.11.10 Kyzylkum autonomous focus (27) in the Kyzylkum Desert within Kazakhstan, Uzbekistan, and Turkmenistan’s eastern outskirts

  • Landscapes: Sandy deserts

  • Hosts: Rhombomys opimus, M. lybicus, M. meridianus

  • Vectors: X. gerbilli, X. hirtipes, X. skrjabini [128]

  • 5.1.2.3.11.11 Muyunkum autonomous focus (28) in the northern Muyunkum Desert, in the subzone within the territories of the Zhambyl and South Kazakhstan regions

  • Landscapes: Deserts

  • Hosts: Rhombomys opimus, M. lybicus, M. meridianus

  • Vectors: X. g. minax, X. conformis [128]

  • 5.1.2.3.11.12 Taukum autonomous focus (29) in the Taukum Desert within the territory of Almaty and Zhambyl between the Ili River in the north and the Chu-Ili Mountains in the south

  • Landscapes: Deserts

  • Hosts: Rhombomys opimus, M. lybicus, M. tamariscinus

  • Vectors: X. skrjabini, X. hirtipes, X. g. minax [128]

  • 5.1.2.3.11.13 Pre-Balkhash autonomous focus (30) in the Balkhash Desert within the territory of Almaty in the interfluves of the Ili-Karatal, Aksu, and Lepsy Rivers

  • Landscapes: Deserts

  • Hosts: Rhombomys opimus, M. meridianus

  • Vectors: X. g. minax, X. hirtipes, X. skrjabini [128]

  • 5.1.2.3.11.14 Betpak-Dala autonomous focus (42) in the Betpakdala Desert west of the Sarysu River and probably extending until Balkhash Lake in the east within Zhambyl and South Kazakhstan

  • Landscapes: Deserts

  • Hosts: Rhombomys opimus, M. lybicus

  • Vectors: X. g. minax, X. conformis [128]

  • 5.1.2.3.11.15 In the Pre-Alakol Lowlands, an autonomous focus (45) in the eastern Pre-Alakol and Dzungarian Gate,wwith a difference in height of 400–900 m above sea level

  • Landscapes: Deserts

  • Hosts: Rhombomys opimus, M. meridianus, M. lybicus

  • Vectors: X. g. minax, X. skrjabini, X. conformis [128]

  • 5.1.2.3.11.16 In the Ili Intermountains: an autonomous focus (46) in the Ili valley from the Kapshagai Reservoir in the west to the border with China in the east and from the lowlands of the southern Jungar Ridge in the north to the foothills of Ketmen, Turaigyr, and Trans-Ili Alatau in the south

  • Landscapes: Deserts, desert lowlands

  • Hosts: Rhombomys opimus, M. meridianus

  • Vectors: X. g. minax, X. hirtipes, X. skrjabini [128]

  • 5.1.2.3.12 Talas: a natural focus (40) in Kyrgyzstan on the northern macroslope of the Talas Ridge from Manas City in the west until the Kolba Ridge in the east extending north to the slopes of the Kyrgyz Ridge (including the Kazakh portion)

  • Landscapes: Mountain steppes, meadow steppes, alpine meadows 1600–3700 m above sea level

  • Hosts: Marmota caudata

  • Vectors: Citellophilus lebedevi , Pulex irritans [128]

  • 5.1.2.3.13 Gissar: a natural focus (34) on the northern macroslope of the Gissar Ridge (Tajikistan)

  • Landscapes: Tree and shrub belts, subalpine meadows

  • Hosts: Microtus juldaschi, Marmota caudata [119]

  • Vectors: Callopsylla caspia, Frontopsylla glabravara [17]

  • 5.1.2.3.14 Sarydzhaz natural focus (foci) of plague in the Tian Shan Mountains in the Sarydzhaz Mountains: an autonomous focus (31) within the Almaty region of Kazakhstan and the Issyk-Kul region of Kyrgyzstan on the slopes of Sary-Djaz on the northeastern tip of the Terskey Alatau Ridge

  • Landscapes: Midlands, highland areas

  • Hosts: Marmota baibacina

  • Vectors: Oropsylla silantiewi, Rhadinopsylla liventricosa, Citellophilus lebedevi [128]

  • 5.1.2.3.15 Upper Naryn autonomous focus (32) within Kyrgyzstan

  • Landscapes: Midlands, mountainous areas

  • Hosts: Marmota baibacina

  • Vectors: Oropsylla silantiewi, Rhadinopsylla liventricosa, Citellophilus lebedevi [128]

  • 5.1.2.3.16 Aksaiautonomous focus (33), within Kyrgyzstan

  • Landscapes: Midlands, mountainous areas

  • Hosts: Marmota baibacina

  • Vectors: Oropsylla silantiewi, Rhadinopsylla liventricosa, Citellophilus lebedevi [128]

  • 5.1.2.3.17 Alai natural focus (35) located on the slopes of the Alai Ridge (Kyrgyzstan)

  • Landscapes: Mountain steppes and subalpine and alpine meadows 2800–5000 m above sea level

  • Hosts: Marmota caudata, Alticola argentatus [129]

  • Vectors: Oropsylla silantiewi crassa, Citellophilus lebedevi princeps [129]

  • 5.1.2.3.18 Mountain-Altai(or Saylyugem, Mongolia), focus (36) in the mountain range of Saylyugem, Kurai, South Chu

  • Landscapes: Mountain steppes, alpine meadows 2000–2500 m above sea level

  • Hosts: Ochotona pallasi, O. alpine[1]

  • Vectors: Paradoxopsyllus scorodumovi, Rhadinopsylla dahurica dahurica [1]

  • 5.1.2.3.19 Tuva (Mongun-Taiga) focus (37) in the Republic of Tyva in the south adjacent to the Hanhiro-Turgensk natural focus (Mongolia):

  • Landscapes: Mountain steppes 1650–2550 m above sea level

  • Hosts: Spermophilus undulatus, Ochotona pallasi [119]

  • Vectors: Citellophilus tesquorum altaicus, Amphalius runatus [1]

  • 5.1.2.3.20 Trans-Baikal natural plague focus (38) in the trans-Baikal steppe; (Russia) and foci in Mongolia (Mongolia-Daurian) and China (Barginsky or Hulun-Byrsk focus):

  • Landscapes: Steppe

  • Hosts: Spermophilus dauricus, Marmota sibirica [130]

  • Vectors: Citellophilus tesquorum sungaris, Oropsylla silatiewi [119, 131]

  • 5.1.2.3.21 Dzungarian Highland natural focus (44) of mixed type (vole-ground squirrel-marmot) in the Dzungarian Highlands (Dzungarian Alatau) wwith a presumably larger part of the focus in China on the Boro-Horo Ridge

  • Landscapes: Subalpine and alpine meadows

  • Hosts: Microtus kirgizorum, Spermophilus undulates

  • Vectors: C. tesquorum, Ct. arvalis, C. ullus, C. assimilis, F. elata [120]

5.1.4 5.1.2.4 Mongolia

  • 5.1.2.4.1 In Khan-Khukhey natural focus in the Altai Mountain system on the Khan-Khukhey Ridge

  • Landscapes: Mountain steppes 2500–2800 m above sea level

  • Hosts: Marmota sibirica, Spermophilus undulatus [124, 183]

  • Vectors: Specific fleas

  • 5.1.2.4.2 KhuKh-Serkh-Munkh-Khairkhan natural focus in the central part of the Mongolian Altai

  • Landscapes: Subalpine and alpine meadows

  • Hosts: Marmota baibacina, Marmota sibirica [1]

  • Vectors: Oropsylla silantiewi

  • 5.1.2.4.3 Gobi Altai Mountains natural focus 3000–3500 m above sea level

  • Landscapes: Mountain steppes

  • Hosts: Marmota sibirica, Ochotona pallasi, Spermophilus undulates [133, 134]

  • Vectors: Oropsylla silantiewi, Paramonopsyllus scalonae, Citellophilus tesquorum altaicus [127]

  • 5.1.2.4.4 Changay Mountain Steppe focus in central Mongolia, in the area of the Khangai Highlands

  • Landscapes: Dry plains and mountain steppe 1200–3000 m above sea level

  • Hosts: Marmota sibirica, Spermophilus undulatus [136, 137]

  • Vectors: Oropsylla silantiewi, Citellophilus tesquorum sungaris [1], [137]

  • 5.1.2.4.5 Khentey natural focus south and southeast of the highlands

  • Landscapes: Steppe 1500–2000 m above sea level

  • Hosts: Marmota sibirica, Spermophilus undulates [128, 138].

  • Vectors: Oropsylla silantiewi [1]

  • 5.1.2.4.6 Mongolian Altai Plain natural focus in the Altaic Gobi

  • Landscapes: Gravelly desert

  • Hosts: Rhombomys opimus, Meriones meridianus [128, 139]

  • Vectors: Xenopsylla conformis conformis, X. skrjabini [140]

  • 5.1.2.4.7 In Gurvan Saikhan, a natural focus in the South Gobi Gurvan Saikhan Mountains

  • Landscapes: Desert steppes 2400–2800 m above sea level

  • Hosts: Ochotona pallasi, O. daurica, Meriones unguiculatus [138, 141]

  • Vectors: Amphalius runatus, Ctenophyllus hirticrus [141]

  • 5.1.2.4.8 On the South Gobi Plain, a natural focus probably connected with that of Inner Mongolia

  • Landscapes: Gravelly desert

  • Hosts: Rhombomys opimus, Meriones meridianus [1]

  • Vectors: Xenopsylla skrjabini, X. conformis conformis [1]

5.1.5 5.1.2.5 China

  • 5.1.2.5.1 Plague focus in the Dzungarian Alatau

  • Landscapes: Mountain steppes, subalpine meadows

  • Hosts: Marmota baibacina, Spermophilus undulates [142]

  • Vectors: Oropsylla silantiewi, Citellophilus tesquorum altaicus, C. tesquorum dzetysuensis [131]

  • 5.1.2.5.2 Inner Mongolia, a natural focus in Dzamin-Uden, Erlyan

  • Landscapes: Semideserts, deserts

  • Hosts: Meriones unguiculatus, Ochotona daurica, O. pallasi [131, 136, 143]

  • Vectors: Xenopsylla conformis conformis, Nosopsyllus laeviceps kuzenkovi, Neopsylla pleskei orientalis [131, 140]

  • 5.1.2.5.3 Xilingol Grassland, a natural focus east of Inner Mongolia

  • Landscapes: Steppe, desert 1100–1300 m above sea level

  • Hosts: Lasiopodomys brandtii, Spermophilus dauricus [131]

  • Vectors: Frontopsylla luculenta luculenta, Amphipsalta primaries mitis, Neopsylla pleskei orientalis [128]

  • 5.1.2.5.4 In Sungari-Lyaoh (Manchuria), a natural focus in northeast China

  • Landscapes: Forest, wet steppes

  • Hosts: Spermophilus dauricus, Rattus norvegicus [131]

  • Vectors: Citellophilus tesquorum sungarus, Xenopsylla cheopis [131]

  • 5.1.2.5.5 Gansu-Ningxia Hui (Ordos, Shaanxi): Natural focus in the Loess Plateau

  • Landscapes: Dry steppes 1500–2950 m above sea level

  • Hosts: Spermophilus alashanicus, Ochotona daurica [131]

  • Vectors: Neopsylla abagaitui, Citellophilus tesquorum mongolicus [131]

  • 5.1.2.5.6 Qinghai-Tibet natural focus in the Tibetan Plateau

  • Landscapes: Subalpine and alpine meadows 2700–5450 m above sea level

  • Hosts: Marmota himalayana, Allactaga sibirica [131]

  • Vectors: Oropsylla silantiewi, Callopsylla dolabris [131]

5.1.6 5.1.2.6 Near East

5.1.6.1 5.1.2.6.1 Yemen

  • Hosts: Rattus rattus, Meriones sp.

  • Vectors: The available literature lacks relevant information.

5.1.6.2 5.1.2.6.2 Saudi Arabia

  • Saudi-Yemeni: Natural focus on the desert plains and uplands along the Saudi Arabia-Yemen border:

  • Landscapes: Up to a height of 1000 m above sea level, rocky and sandy deserts, desert savannas, shrub-tree xerophilic communities

  • Possible hosts: Gerbillus nanus, G. cheesmani, Meriones rex, Rattus rattus, Mus musculus, Acomys cahirinus, Jaculus jaculus [144]

  • 5.1.2.6.3 In the Lebanese mountains and on the steppe, foci near the Syrian coast and the mountainous parts of Lebanon and northern Israel

  • Landscapes: Forb-grass mountain steppes

  • Hosts: Gerbils of the genus Meriones

  • 5.1.2.6.4 In the Syrian-Mesopotamian Desert: natural foci in Upper and Lower Mesopotamia, the Syrian Plateau, and Kuwait

  • Landscapes: Desert flood plains of the Tigris and Euphrates Rivers and surrounding desert plateau

  • Hosts: Meriones libycus, Tatera indica, Nesokia indica, Rattus rattus, Rattus norvegicus, Mus musculus. Perhaps: Gerbillus cheesmani, G. dasyurus, G. nanus, Meriones crassus [144]

  • Vectors: Xenopsylla buxtoni, X. astia, Stenoponia tripectinata insperata; plague transmission from rodents to humans via the human flea Pulex irritans [145]

  • 5.1.2.6.5 Natural foci in the semidesert and desert of southern Turkey, northern Syria

  • Landscapes: rocky ephemeral-wormwood deserts

  • Hosts: Mesocricetus brandti, Microtus socialis, Tatera indica, Meriones vinogradovi, M. libycus, M. tristrami, Nesokia indica [146]

  • Vectors: Xenopsylla buxtoni

  • 5.1.2.6.6 Kurds-Iranian mountains and steppe: Natural focus in the western Iranian plateau, southeastern Turkey, northeastern Iraq [1]

  • Landscapes: Mountain plateaus, intermountain valleys 1000–2000 m above sea level, covered with herb-grass steppes

  • Hosts: Meriones persicus, M. libycus, M. vinogradovi, M. tristrami [145], Allactaga elater, Mesocricetus brandti, Ellobius lutescens [147, 148]

  • Vectors: Xenopsylla buxtoni, X. conformis conformis, Nosopsyllus iranus iranus, Stenoponia tripectinata insperata [147, 148], Pulex irritans [149], and others [1]

  • 5.1.2.6.7 Natural focus in the Armenian-Anatolian mountains and steppe, previously considered part of the Kurdish-Iranian focus

  • Landscapes: Steppe, mountain steppe

  • Hosts: Meriones libycus, Spermophilus xanthoprymnus [1, 150]

  • Vectors: Same as those of the Kurdish-Iranian focus

  • 5.1.2.6.8 In the Iran-Afghan Desert lowlands, a natural focus in the central low and northern mountains of Iran, Afghanistan's western plateau

  • Landscapes: Semideserts, deserts of Turan and Southwest Asia, and possibly on the outskirts of the Central Asian desert focus [151]:

  • Hosts: Rhombomys opimus, Meriones libycus, M. meridianus, Mus musculus [1]

  • Vectors: Xenopsylla gerbilli, X. nuttalli, X. conformis conformis, Nosopsyllus laeviceps [1]

  • 5.1.2.6.9 Along the South Afghan flats, a natural focus in the sandy desert of Registan, sands in the lower reaches of the Helmand River, and the clay-stony desert Dasht-Markokhim

  • Hosts: Rhombomys opimus, Meriones persicus, M. libycus, Mus musculus [152]

  • Vectors: Xenopsylla cheopis, X. gerbilli, X. conformis conformis [153]

  • 5.1.2.6.10 Natural foci in the Afghan-Pakistani midland desert-steppe of central, eastern Afghanistan and the western regions of Pakistan

  • Landscapes: Subtropical dry mountain steppes, acacia woodlands

  • Hosts: Meriones libycus, M. persicus, Marmota caudata, Spermophilus fulvus, Mus musculus, Rattus rattus [152]

  • Vectors: Xenopsylla cheopis, X. gerbilli, X. conformis conformis [153]

  • 5.1.2.6.11 Natural focus in the Hindu Kush Highlands [150]

  • Landscapes: Alpine herb-grass meadows

  • Hosts: Marmota caudata, Dryomysnitedula, Microtus juldashi, Meriones persicus, Mus musculus

  • Vectors: Xenopsylla cheopis, Citellophilus lebedevi, Frontopsylla protera

  • A focus in Northern Afghanistan: within the range of Rhombomys opimus [154]

5.1.7 5.1.2.7 North America

  • 5.1.2.7.1 In California and Oregon, foci, between the ridges of the Sierra Nevada and the coast, and the San Joaquin and Sacramento Rivers

  • Landscapes: The available literature lacks relevant information.

  • Hosts: Spermophilus beecheyi

  • Vectors: Diamanus montanus, Hoplopsyllus anomalus [30]

  • 5.1.2.7.2 In Oregon, Idaho, Montana, and Wyoming, in theBlue Cascade Mountains, Bitter Root Ridge, and eastern slopes of Moustache Ridge

  • Hosts: Citellus lateralis, Citellus richardsoni [30]

  • Nevada and Utah: Foci in the basin of the Great Salt Lake

  • Landscapes: Desert

  • Hosts: Neotoma lepida, Cynomys parvidens

  • 5.1.2.7.3 In New Mexico, Arizona, and Utah, foci in the Colorado Plateau and Llano Estacado (entering the territory of Mexico)

  • Landscapes: Deserts and steppes

  • Hosts: Cynomys ludovicianus, C. gunnisoni

  • 5.1.2.7.4 In California, foci in the mountainous area of San Bruno

  • Hosts: Microtus californicus, Peromyscus maniculatus [30]

  • Vectors (for the whole territory): Oropsylla montana, O. idahoensis, Ceratophyllus ciliatus, Orchopeas nepos (Smith et al., 2009)

5.1.8 5.1.2.8 South America

5.1.8.1 5.1.2.8.1 Argentina

  • 5.1.2.8.1.1 Foci in the provinces of Santiago del Estero, Salta, Catamarca

  • Landscapes: Rainforests

  • Hosts: Microcavia australis, Galeamus teloides [30]

  • 5.1.2.8.1.2 Foci in the provinces of Cordoba, San Juan, Mendoza, La Pampa, and Rio Negro, on the eastern slopes of the Andes

  • Landscapes: Alpine steppe

  • Hosts: Microcavia australis, Galeamus teloides

  • Focus in Buenos Aires Province

  • Landscapes: Steppe

  • Hosts: Microcavia australis [30]

  • Vectors: The available literature lacks relevant information.

5.1.8.2 5.1.2.8.2 Bolivia

  • 5.1.2.8.2.1 Foci in the departments of Chuquisaca, Santa Cruz, Tarija, probably connected with the northern foci of Argentina

  • Landscapes: The available literature lacks relevant information.

  • Hosts: Genus Graomys, Hesperomys

  • Vectors: Puligenis biturdus, P. bochisi [30]

5.1.8.3 5.1.2.8.3 Brazil

  • 5.1.2.8.3.1 Enzootic areas in Sinara, Pernambuco, Paraiba, Alagoas, Bahia, Minas Gerais, Teresopolis

  • Landscapes: The available literature lacks relevant information.

  • Hosts: Cavia aperea, C. spixii, Oryzomys subflavus, Zygedon tomispixuna

  • Vectors: Poligenis bohsli, P. tripus [30, 155]

5.1.8.4 5.1.2.8.4 Ecuador

  • 5.1.2.8.4.1 Natural focus in Manabi, western Ecuador

  • Hosts: Sigmodon hispidus

  • Vectors: X. cheopis [30]

  • 5.1.2.8.4.2 Natural foci in Chimborazo and Tungurahua, central Ecuador

  • Landscapes: Alpine steppes

  • Hosts: Sylvilagus brasiliensis

  • Vectors: Pleochaetis dolens, Hoplopsyllus manconis [30]

  • 5.1.2.8.4.3 Natural foci in Loho and El Oro, southern Ecuador

  • Landscapes: The available literature lacks relevant information.

  • Hosts: Sciurus stramineus, Akodon mollis

  • Vectors: Poligenis litardus, Pulex irritans [30]

5.1.8.5 5.1.2.8.5 Peru

  • 5.1.2.8.5.1 In northern Peru: possibly the same focus as that in Ecuadorian Loho and El Oro Provinces [30]

  • Hosts: Sciurus stramineus, Akodon mollis

5.1.8.6 5.1.2.8.6 Venezuela

  • 5.1.2.8.6.1 Nature foci in the states of Miranda and Aragua

  • Hosts: Sigmodon hispidus, Heteromys anomalus [30]

  • Vectors: The available literature lacks relevant information.

5.1.9 5.1.2.9 Hawaiian Islands

  • Possible foci of plague in the districts of Hamakura and Hilo:

  • Landscapes: The available literature lacks relevant information.

  • Hosts: Rattus hawaiiensis

  • Vectors: Xenopsylla vexabilis [30]

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Dubyanskiy, V.M., Yeszhanov, A.B. (2016). Ecology of Yersinia pestis and the Epidemiology of Plague. In: Yang, R., Anisimov, A. (eds) Yersinia pestis: Retrospective and Perspective. Advances in Experimental Medicine and Biology, vol 918. Springer, Dordrecht. https://doi.org/10.1007/978-94-024-0890-4_5

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