The transimission ecology of the echinococcosis on the Tibetan Plateau

doi:10.12140/j.issn.1000-7423.2022.04.017

Abstract

Abstract:

Zoonotic diseases are not only significant risks to humans, but also seriously affect global biodiversity conservation. The eastern Tibetan plateau has the highest echinococcosis prevalence in the world. The complex and stable echinococcosis transmission networks is based on several wild and domestic animal species. In this study, we summarized the epidemiological research advances of echinococcosis in the rural area of the Tibetan plateau. We also discussed the ecological factors and their functional mechanisms for the transmission of echinococcosis including host species community composition and dynamics, host species behaviour ecology, and human disturbance. Based on these discussions, we put forward suggestions for a better integrative strategy for echinococcosis control and the sustainable development of the plateau ecosystem.

Key words: Echinococcosis, Echinococcus, Tibetan Plateau, Host, Wildlife

CLC Number:

R532.32

ZUO Qing-qiu, ZHENG Jia-xin, WANG Gang, WANG Xu, WANG Xiao-ming, WANG Zheng-huan. The transimission ecology of the echinococcosis on the Tibetan Plateau[J]. CHINESE JOURNAL OF PARASITOLOGY AND PARASITIC DISEASES, 2022, 40(4): 524-535.

Figures/Tables 2

感染源	中间宿主	感染率/%	地区	参考文献
棘球蚴（未鉴定物种）	岩羊	6.4 (21/327)	青海	[50]
		6.02 (21/349)	青海	[47]
	藏原羚	6.6 (13/198)	青海	[50]
		6.57 (13/198)	青海	[47]
	猪	5.62 (114/2030)	青海	[47]
	山羊	30.43 (469/1541)	青海	[47]
	蒙原羚	(1/1)	青海	[47]
细粒棘球蚴（广义种）	高原鼠兔	(5/7)	青海	[110]
	牦牛	71.6 (521/728)	青海	[109]
.		78.5 (106/135)	青海	[111]
		21.27 (67/135)	四川	[112]
		50.8 (219/429)	四川	[40]
		19.9 (126/634)	甘肃	[41]
		6.5 (49/757)	甘肃	[113]
		25.6 (33/129)	甘肃	[33]
		9.15 (91/995)	西藏	[114]
	绵羊	82.6 (95/115)	青海	[111]
		66.5 (666/1002)	青海	[109]
		18.24 (191/1047)	西藏	[114]
		11.1 (113/1021)	甘肃	[41]
	双峰驼	24.6 (67/272)	内蒙古	[46]
多房棘球蚴	阿尔泰黄鼠	0.1 (2/2211)	新疆	[115]
	伊犁田鼠	0.8 (7/916)	新疆	[116]
	水	1.6 (1/61)	新疆	[117]
	达乌尔黄鼠	0.2 (3/1500)	宁夏	[118]
	中华鼢鼠	0.3 (1/321)	宁夏	[119]
	布氏田鼠	2.4 (64/2635)	内蒙古	[120]
		6.3 (198/3157)	内蒙古	[121,122]
	青海松田鼠	(1/5)	青海	[43]
		22 (11/50)	青海	[123]
		11.1 (16/144)	四川	[23]
	白尾松田鼠	(3/12)	四川	[40]
	长爪沙鼠	(1/6)	内蒙古	[120]
	柴达木根田鼠	14.7 (5/34)	四川	[51]
		11.4 (5/44)	四川	[23]
	灰仓鼠	2.8 (1/35)	青海	[49]
	藏仓鼠	(1/19)	四川	[51]
	高原鼠兔	9.4 (24/256)	四川	[44]
		5.6 (13/233)	四川	[40]
		3.4 (11/319)	青海	[43]
		15.2 (34/224)	青海	[43]
		4.2 (34/801)	青海	[49]
		1.5 (2/135)	四川	[23]
	灰尾兔	6.7 (5/75)	四川	[44]
		(1/14)	四川	[40]
		(1/19)	青海	[49]
		(1/5)	青海	[43]
	牦牛	0.3 (2/634)	甘肃	[41]
		0.7 (3/429)	四川	[40]
	绵羊	0.3 (3/1021)	甘肃	[41]
石渠棘球蚴	青海松田鼠	0.356 (1/224)	青海	[52]
		1.4 (2/144)	四川	[23]
	长尾仓鼠	(1/1)	四川	[23]
	白尾松田鼠	(1/16)	四川	[23]
	柴达木根田鼠	11.4 (5/44)	四川	[23]
	高原鼠兔	11.3 (27/239)	青海	[126]
		3.7 (5/135)	四川	[23]

References 127

[1]	Taylor LH, Latham SM, Woolhouse ME. Risk factors for human disease emergence[J]. Philos Trans R Soc Lond B Biol Sci, 2001, 356(1411): 983-989. doi: 10.1098/rstb.2001.0888
[2]	Daszak P, Cunningham AA, Hyatt AD. Emerging infectious diseases of wildlife: threats to biodiversity and human health[J]. Science, 2000, 287(5452): 443-449. pmid: 10642539
[3]	Altizer S, Nunn CL, Thrall PH, et al. Social organization and parasite risk in mammals: integrating theory and empirical studies[J]. Annu Rev Ecol Evol Syst, 2003, 34: 517-547. doi: 10.1146/annurev.ecolsys.34.030102.151725
[4]	Bradley CA, Altizer S. Urbanization and the ecology of wildlife diseases[J]. Trends Ecol Evol, 2007, 22(2): 95-102. doi: 10.1016/j.tree.2006.11.001
[5]	Li TY, Qiu JM, Yang W, et al. Echinococcosis in Tibetan populations, western Sichuan Province, China[J]. Emerg Infect Dis, 2005, 11(12): 1866-1873. doi: 10.3201/eid1112.050079
[6]	Clark L, Hall J. Avian influenza in wild birds: status as reservoirs, and risks to humans and agriculture[J]. Ornithol Monogr, 2006(60): 3-29.
[7]	Alwan NA, Burgess RA, Ashworth S, et al. Scientific consensus on the COVID-19 pandemic: we need to act now[J]. Lancet, 2020, 396(10260): e71-e72. doi: 10.1016/S0140-6736(20)32153-X
[8]	Daszak P, Cunningham A. Anthropogenic change, biodiversity loss, and a new agenda for emerging diseases[J]. J Parasitol, 2003, 89: S37-S41.
[9]	Zhou P, Chen N, Zhang RL, et al. Food-borne parasitic zoonoses in China: perspective for control[J]. Trends Parasitol, 2008, 24(4): 190-196. doi: 10.1016/j.pt.2008.01.001
[10]	Wen H, Ding ZX. Atlas of echinococcosis: Chinses-English edition[M]. 2nd ed. Beijing: Science Press, 2008: 2-3. (in Chinese)
	( 温浩, 丁兆勋. 包虫病图谱: 中英文双语版[M]. 2版. 北京: 科学出版社, 2008: 2-3.)
[11]	Eckert J, Deplazes P. Biological, epidemiological, and clinical aspects of echinococcosis, a zoonosis of increasing concern[J]. Clin Microbiol Rev, 2004, 17(1): 107-135. doi: 10.1128/CMR.17.1.107-135.2004
[12]	Thompson RCA, McManus DP. Aetiology: parasite and life-cycles[R]. Geneva: WHO, 2001: 1-17.
[13]	Tang GB. Practical echinococcosis imaging[M]. Beijing: People’s Medical Publishing House, 2013: 8-13. (in Chinese)
	( 唐桂波. 实用包虫病影像学[M]. 北京: 人民卫生出版社, 2013: 8-13.)
[14]	Hao LL, Zhou MZ. A new strategy for echinococcosis control: "Shiqu 238" model[M]. Beijing: Chinese Agriculture Press, 2018: 52-56. (in Chinese)
	( 郝力力, 周明忠. 动物包虫病防控新策略: “石渠238”模式[M]. 北京: 中国农业出版社, 2018: 52-56.)
[15]	Tappe D, Kern P, Frosch M, et al. A hundred years of controversy about the taxonomic status of Echinococcus species[J]. Acta Trop, 2010, 115(3): 167-174. doi: 10.1016/j.actatropica.2010.03.001
[16]	Thompson RCA. Biology and systematics of Echinococcus[J]. Adv Parasitol, 2017, 95: 65-109.
[17]	Pawlowski ZS, Eckert J, Vuitton DA, et al. Echinococcosis in humans: clinical aspects, diagnosis and treatment[R]. Geneva: WHO, 2001: 20-58.
[18]	Wang GQ. Epidemiological survey on echinococcosis in China[M]. Shanghai: Shanghai Scientific & Technical Publishers, 2016: 6-21. (in Chinese)
	( 王国强. 全国包虫病流行情况调查报告[M]. 上海: 上海科学技术出版社, 2016: 6-21.)
[19]	The central people’s government of the people’s republic of China. Action Plan for echinococcosis (2010—2015) [Z/OL]. (2010-12-14) [2021-03-08]. http://www.gov.cn/zwgk/2010-12/14/content_1765485.htm. (in Chinese)
	( 中华人民共和国中央人民政府. 关于印发《防治包虫病行动计划(2010—2015年)》的通知[Z/OL]. (2010-12-14) [2021-03-08]. http://www.gov.cn/zwgk/2010-12/14/content_1765485.htm. )
[20]	Shang ZH, Gibb MJ, Leiber F, et al. The sustainable development of grassland-livestock systems on the Tibetan Plateau: problems, strategies and prospects[J]. Rangel J, 2014, 36(3): 267. doi: 10.1071/RJ14008
[21]	Smith AT, Xie Y. A guide to the mammals of China[M]. Changsha: Hunan Education Publishing House, 2009: 404-411. (in Chinese)
	( Smith AT, 解焱. 中国兽类野外手册[M]. 长沙: 湖南教育出版社, 2009: 404-411.)
[22]	Jiang WB, Liu N, Zhang GT, et al. Specific detection of Echinococcus spp. from the Tibetan fox (Vulpes ferrilata) and the red fox (V. vulpes) using copro-DNA PCR analysis[J]. Parasitol Res, 2012, 111(4): 1531-1539. doi: 10.1007/s00436-012-2993-8
[23]	Wang X, Liu JY, Zuo QQ, et al. Echinococcus multilocularis and Echinococcus shiquicus in a small mammal community on the eastern Tibetan Plateau: host species composition, molecular prevalence, and epidemiological implications[J]. Parasit Vect, 2018, 11(1): 302. doi: 10.1186/s13071-018-2873-x
[24]	Wang H, Chai JJ, Liu FJ, et al. A study on ecological-epidemiology of two hydatid disease in Qinghai[J]. Chin J Parasit Dis Control, 2002(5): 284-287. (in Chinese)
	( 王虎, 柴君杰, 刘凤洁, 等. 青海省包虫病的生态流行病学研究[J]. 中国寄生虫病防治杂志, 2002(5): 284-287.)
[25]	Weng XD, Mu ZQ, Wei X, et al. The effects of dog management on Echinococcus spp. prevalence in villages on the eastern Tibetan Plateau, China[J]. Parasit Vect, 2020, 13(1): 207. doi: 10.1186/s13071-020-04082-6
[26]	Li T, Chen X, Zhen R, et al. Widespread co-endemicity of human cystic and alveolar echinococcosis on the eastern Tibetan Plateau, northwest Sichuan/southeast Qinghai, China[J]. Acta Trop, 2010, 113(3): 248-256. doi: 10.1016/j.actatropica.2009.11.006
[27]	Wu WP, Wang H, Wang Q, et al. A nationwide sampling survey on echinococcosis in China during 2012—2016[J]. Chin J Parasitol Parasit Dis, 2018, 36(1): 1-14. (in Chinese)
	( 伍卫平, 王虎, 王谦, 等. 2012—2016年中国棘球蚴病抽样调查分析[J]. 中国寄生虫学与寄生虫病杂志, 2018, 36(1):1-14.)
[28]	Craig PS, Giraudoux P, Wang ZH, et al. Echinococcosis transmission on the Tibetan Plateau[J]. Adv Parasitol, 2019, 97: 165-246.
[29]	Wang M. Dynamic monitoring of land use change on Tibetan Plateau[D]. Lanzhou: Lanzhou University, 2015: 21-31. (in Chinese)
	( 王敏. 青藏髙原地区土地利用变化动态监测研究[D]. 兰州: 兰州大学, 2015: 21-31.)
[30]	Zhang YL, Liu LS, Wang ZF, et al. Spatial and temporal characteristics of land use and cover changes in the Tibetan Plateau[J]. Chin Sci Bull, 2019, 64(27): 2865-2875. (in Chinese) doi: 10.1360/TB-2019-0046
	( 张镱锂, 刘林山, 王兆锋, 等. 青藏高原土地利用与覆被变化的时空特征[J]. 科学通报, 2019, 64(27): 2865-2875.)
[31]	Eckert J, Schantz PM, Gasser RB, et al. WHO/OIE manual on echinococcosis in humans and animals: a public health problem of global concern[R]. Paris: WHO, 2001: 100-118.
[32]	Ohiolei JA, Xia CY, Li L, et al. Genetic variation of Echinococcus spp. in yaks and sheep in the Tibet Autonomous Region of China based on mitochondrial DNA[J]. Parasit Vect, 2019, 12(1): 608. doi: 10.1186/s13071-019-3857-1
[33]	Wu YT, Li L, Zhu GQ, et al. Mitochondrial genome data confirm that yaks can serve as the intermediate host of Echinococcus canadensis (G10) on the Tibetan Plateau[J]. Parasit Vectors, 2018, 11(1): 166. doi: 10.1186/s13071-018-2684-0
[34]	Shi YL, Wan XL, Wang ZY, et al. First description of Echinococcus ortleppi infection in China[J]. Parasit Vect, 2019, 12(1): 398. doi: 10.1186/s13071-019-3653-y
[35]	Xiao N, Qiu JM, Nakao M, et al. Echinococcus shiquicus n. sp., a taeniid cestode from Tibetan fox and plateau pika in China[J]. Int J Parasitol, 2005, 35(6): 693-701. doi: 10.1016/j.ijpara.2005.01.003
[36]	Musinov M. The epizootiology of echinococcosis in animals in Uzbekistan[J]. Med Parazitol (Mosk), 1998(4): 40-41.
[37]	Craig PS,. Epidemiology of human alveolar echinococcosis in China[J]. Parasitol Int, 2006, 55: S221-S225. doi: 10.1016/j.parint.2005.11.034
[38]	Boufana B, Qiu JM, Chen XW, et al. First report of Echinococcus shiquicus in dogs from eastern Qinghai-Tibet Plateau region, China[J]. Acta Trop, 2013, 127(1): 21-24. doi: 10.1016/j.actatropica.2013.02.019
[39]	Zhang ZX. Qinghai geography[M]. 2nd ed. Beijing: Science Press, 2009: 148-166. (in Chinese)
	( 张忠孝. 青海地理[M]. 2版. 北京: 科学出版社, 2009: 148-166.)
[40]	He JG, Qiu JM, Liu FJ, et al. Epidemiological survey on hydatidosis in tibitan region of western Sichuan Ⅱ. infection situation among domestic and wild animals[J]. Chin J Zoonoses, 2000, 16(5): 62-65. (in Chinese)
	( 何金戈, 邱加闽, 刘凤洁, 等. 四川西部藏区包虫病流行病学研究Ⅱ.牲畜及野生动物两型包虫病感染状况调查[J]. 中国人兽共患病杂志, 2000, 16(5): 62-65.)
[41]	Zhao YM. Epidemiological studies of hydtidosis in the east part of Qinghai-Tibet Plateau (Gannan Tibetan Autonomous Prefecture)[D]. Lanzhou: Gansu Agricultural University, 2008: 25-51. (in Chinese)
	( 赵玉敏. 青藏高原东段(甘南藏族自治州)包虫病的流行病学研究[D]. 兰州: 甘肃农业大学, 2008: 25-51.)
[42]	Lin YG, Lu MK, Hong LX. A review of the current status of Echinococcus and hydatid disease, with notes on some informative achievements in China[J]. Chin J Zoono, 2012, 28(6): 616-627. (in Chinese)
	( 林宇光, 卢明科, 洪凌仙. 回顾棘球绦虫及棘球蚴病的研究现状—着重报道我国的贡献成果[J]. 中国人兽共患病学报, 2012, 28(6): 616-627.)
[43]	Zhao HL. Investigation on infections of alveolar hydatid in small mammals at south Qinghai plateau[J]. J Qinghai Med Coll, 2002, 23(2): 12-14. (in Chinese)
	( 赵海龙. 青海省南部地区小型兽类多房棘球绦虫感染调查[J]. 青海医学院学报, 2002, 23(2): 12-14.)
[44]	Qiu JM, Chen XW, Ren M, et al. Epidemiological study on alveolar hydatid disease in Qinghai-Xizang Plateau[J]. J Pract Parasit Dis, 1995(3): 106-109. (in Chinese)
	( 邱加闽, 陈兴旺, 任敏, 等. 青藏高原泡球蚴病流行病学研究[J]. 实用寄生虫病杂志, 1995(3): 106-109.)
[45]	Romig T, Deplazes P, Jenkins D, et al. Ecology and life cycle patterns of Echinococcus species[J]. Adv Parasitol, 2017, 95: 213-314.
[46]	Tu BX, Zhang WL, Zhang WB, et al. [J]. Investigation of Echinococcus granulosus infection in camels in Alxa Left Banner, Inner Mongolia[J]. End Dis Bull, 18(2): 99. (in Chinese)
	( 图布新, 张文林, 张文彬, 等. 内蒙古阿拉善左旗骆驼中细粒棘球蚴感染的调查[J]. 地方病通报, 2003, 18(2): 99.)
[47]	Cai HX, Wang H, Han XM, et al. Current situation of Echinococcus infection in different hosts in Qinghai Plateau[J]. Chin J Endem, 2012(3): 296-300. (in Chinese)
	( 蔡辉霞, 王虎, 韩秀敏, 等. 青海高原棘球绦虫不同宿主感染情况及意义的研究[J]. 中国地方病学杂志, 2012(3): 296-300.)
[48]	Schaller GB. Wildlife of the Tibetan Steppe[M]. Chicago: University of Chicago Press, 1998.
[49]	Wang H, Zhao HL, Ma SM, et al. Investigation on infections of Echinococcus in animals in Qinghai Plateau[J]. Endem Dis Bull China, 2000, 15(3): 29-33. (in Chinese)
	( 王虎, 赵海龙, 马淑梅, 等. 青海动物棘球绦虫感染调查研究[J]. 地方病通报, 2000, 15(3): 29-33.)
[50]	Guo ZX, He DL, Li YQ, et al. Infection of Echinococcus species in wild animals in Qinghai Province[J]. Chin J Parasitol Parasit Dis, 1994(S1): 145-147. (in Chinese)
	( 郭再宣, 何多龙, 李予青, 等. 青海高原野生动物棘球绦虫感染情况的调查[J]. 中国寄生虫学与寄生虫病杂志, 1994(S1): 145-147.)
[51]	Giraudoux P, Pleydell D, Raoul F, et al. Transmission ecology of Echinococcus multilocularis: what are the ranges of parasite stability among various host communities in China?[J]. Parasitol Int, 2006, 55(Suppl): S237-S246.
[52]	Han XM, Wang H, Qiu JM, et al. Investigation and analysis on the prevalence of alveolar echinococcosis and cystic echinococcosis in Banma County, Qinghai Province[J]. Chin J Zoonoses, 2006, 22(2): 189-190. (in Chinese)
	( 韩秀敏, 王虎, 邱加闽, 等. 青海省班玛县泡型和囊型包虫病流行现状调查分析[J]. 中国人兽共患病学报, 2006, 22(2): 189-190.)
[53]	Giraudoux P, Raoul F, Pleydell D, et al. Drivers of Echinococcus multilocularis transmission in China: small mammal diversity, landscape or climate?[J]. PLoS Negl Trop Dis, 2013, 7(3): e2045.
[54]	Liu SY, Sun Z, Liu Y, et al. A new vole from Xizang, China (Cricetidae ∶ Arvicolinae) and the molecular phylogeny of the genus Neodon[J]. Zootaxa, 2012, 3235(3235): 1-22. doi: 10.11646/zootaxa.3235.1.1
[55]	Liu SY, Jin W, Liu Y, et al. Taxonomic position of Chinese voles of the tribe arvicolini and the description of 2 new species from Xizang, China[J]. J Mammal, 2017, 98(1): 166-182.
[56]	Xiao N, Nakao M, Qiu JM, et al. Dual infection of animal hosts with different Echinococcus species in the eastern Qinghai-Tibet Plateau region of China[J]. Am J Trop Med Hyg, 2006, 75(2): 292-294. doi: 10.4269/ajtmh.2006.75.292
[57]	Boufana B, Umhang G, Qiu JM, et al. Development of three PCR assays for the differentiation between Echinococcus shiquicus, E. granulosus (G1 genotype), and E. multilocularis DNA in the co-endemic region of Qinghai-Tibet Plateau, China[J]. Am J Trop Med Hyg, 2013, 88(4): 795-802. doi: 10.4269/ajtmh.12-0331
[58]	Wang Q, Raoul F, Budke C, et al. Grass height and transmission ecology of Echinococcus multilocularis in Tibetan communities[J]. Chin Med J, 2010, 123: 61-67.
[59]	Smith AT, Zahler P, Hinds A. Conservation Biology in Asia[R]. Kathmandu: Society for Conservation Biology Asia Section and Resources Himalaya, 2006: 285-293.
[60]	Fan N, Zhou W, Wei W, et al. Ecologically-based management of rodent pests[M]. Canberra: Australian: Australian Centre for International Agricultural Research, 1999: 285-304.
[61]	Krebs CJ. Population fluctuations in rodents[M]. Chicago: The University of Chicago Press, 2013: 230-247.
[62]	Liu JK, Zhang YZ, Xin GW. Relationship between numbers and degree of harmfulness of the plateau pika[J]. Acta Zool Sin, 1980, 26(4): 378-385. (in Chinese)
	( 刘季科, 张云占, 辛光武. 高原鼠兔数量与危害程度的关系[J]. 动物学报, 1980, 26(4): 378-385.)
[63]	Wei WH, Zhou WY. Natural enemies of rodents and their protection in Haibei Alpine meadow ecosystem[J]. J Qinghai Enviroment, 1997, 7(4): 145-149. (in Chinese)
	( 魏万红, 周文扬. 海北高寒草甸生态系统中鼠类的天敌动物及其保护[J]. 青海环境, 1997, 7(4): 145-149.)
[64]	Zhao L, Zhou S, Yan DH, et al. Developing trend and controlling countermeasures of pests and mice of Sichuan grassland in 2015[J]. Prataculture Animal Husb, 2015(3): 49-53. (in Chinese)
	( 赵磊, 周俗, 严东海, 等. 2015年四川省草原鼠虫害发生趋势分析及防治对策[J]. 草业与畜牧, 2015(3): 49-53.)
[65]	Zhao L, Yan DH, Zhang XX, et al. Actuality and control technology of prairie rodent pests in Sichuan[J]. Prataculture Animal Husb, 2015(4): 3-7. (in Chinese)
	( 赵磊, 严东海, 张绪校, 等. 四川省草原鼠害现状与防控技术进展[J]. 草业与畜牧, 2015(4): 3-7.)
[66]	Zhang ZB, Zhong WQ, Fan NC. Rats, mice and people: rodent biology and management[M]. Canberra: ACIAR Monograph, 2003: 316-319.
[67]	Liu JY. Glires species composition and habitat utilization pattern in alpine meadow ecosystem[D]. Shanghai: East China Normal University, 2017: 56-68. (in Chinese)
	( 刘佳余. 高寒草甸生态系统啮齿类物种组成和生境利用模式[D]. 上海: 华东师范大学, 2017: 56-68.)
[68]	Mu ZQ. Comparative study on community structure, spatial distribution of burrows and prevalence of echinococcosis of small mammals in wild and human living environment[D]. Shanghai: East China Normal University, 2018: 27-49. (in Chinese)
	( 慕志强. 野外和人居环境啮齿类群落结构、洞穴空间分布及棘球蚴病感染情况对比研究[D]. 上海: 华东师范大学, 2018: 27-49.)
[69]	Zheng SW. Data on the foods of Tibetan sand fox[J]. Acta Theriol Sin, 1985, 5(3): 222-240. (in Chinese)
	( 郑生武. 藏狐的食性资料[J]. 兽类学报, 1985, 5(3): 222-240.)
[70]	Harris RB, Zhou JK, Ji YQ, et al. Evidence that the Tibetan fox is an obligate predator of the plateau pika: conservation implications[J]. J Mammal, 2014, 95(6): 1207-1221. doi: 10.1644/14-MAMM-A-021
[71]	Yang QS, Yan K. Color atlas of mammals of China[M]. Beijing: Science Press, 2007: 27. (in Chinese)
	( 杨奇森, 岩崑. 中国兽类彩色图谱[M]. 北京: 科学出版社, 2007: 27.)
[72]	Gao G, Wang SW. Host animals and the control of plague in China[M]. Lanzhou: Gansu Science & Technology Press, 2012: 89. (in Chinese)
	( 高共, 王升文. 中国鼠疫宿主动物及其防制[M]. 兰州: 甘肃科学技术出版社, 2012: 89.)
[73]	Burlet P, Deplazes P, Hegglin D. Age, season and spatio-temporal factors affecting the prevalence of Echinococcus multilocularis and Taenia taeniaeformis in Arvicola terrestris[J]. Parasit Vect, 2011, 4: 6. doi: 10.1186/1756-3305-4-6
[74]	Tsukada H, Li W, Duo H, et al. A comparison of the diet and fine-scale distribution of sympatric Tibetan and red foxes in Qinghai, PR China[J]. Wildl Biol, 2014, 20(6): 356-361. doi: 10.2981/wlb.00066
[75]	Wang ZH, Wang XM, Ding YZ. The important function of foxes in the transmission of two kinds of echinococcosises[J]. Nat Mag, 2003, 25(5): 255-260. (in Chinese)
	( 王正寰, 王小明, 丁由中. 狐狸在两型包虫病传播途径中所起的重要作用[J]. 自然杂志, 2003, 25(5): 255-260.)
[76]	Guan XH. Prevention and control of cysticercosis and echinococcosis in sheep[J]. Chin J Animal Husb Vet Med, 2019(5): 58. (in Chinese)
	( 关向晖. 羊细颈囊尾蚴病和棘球蚴病的防控方案[J]. 畜牧兽医科技信息, 2019(5): 58.)
[77]	Liccioli S, Giraudoux P, Deplazes P, et al. Wilderness in the ‘city’ revisited: different urbes shape transmission of Echinococcus multilocularis by altering predator and prey communities[J]. Trends Parasitol, 2015, 31(7): 297-305. doi: 10.1016/j.pt.2015.04.007 pmid: 25985897
[78]	Raoul F, Deplazes P, Rieffel D, et al. Predator dietary response to prey density variation and consequences for cestode transmission[J]. Oecologia, 2010, 164(1): 129-139. doi: 10.1007/s00442-010-1647-8
[79]	Raoul F, Hegglin D, Giraudoux P. Trophic ecology, behaviour and host population dynamics in Echinococcus multilocularis transmission[J]. Vet Parasitol, 2015, 213(3/4): 162-171. doi: 10.1016/j.vetpar.2015.07.034
[80]	Liu XQ, Wang XM, Wang ZH, et al. Comparison of the Tibetan fox(Vulpes ferrilata) home range size using methods the fixed kernel estimation and the minimum convex polygon[J]. Acta Theriol Sin, 2010, 30(2): 163-170. (in Chinese)
	( 刘晓庆, 王小明, 王正寰, 等. 固定核空间法和最小凸多边形法估计藏狐家域的比较[J]. 兽类学报, 2010, 30(2): 163-170.)
[81]	Vaniscotte A, Raoul F, Poulle ML, et al. Role of dog behaviour and environmental fecal contamination in transmission of Echinococcus multilocularis in Tibetan communities[J]. Parasitology, 2011, 138(10): 1316-1329. doi: 10.1017/S0031182011000874 pmid: 21854704
[82]	Chineses center for disease control and prevetion. National Program for the Prevention and Control of Echinococcosis and other Key Parasitic Diseases (2016—2020) [Z/OL]. (2016-10-24) [2021-03-08]. http://www.nhc.gov.cn/jkj/s5873/201702/dda5ff-e3f50941a29fb0aba6233bb497.shtml. (in Chinese)
	疾病预防控制局. 关于印发全国包虫病等重点寄生虫病防治规划(2016—2020年)的通知[Z/OL]. (2016-10-24) [2021-03-08]. http://www.nhc.gov.cn/jkj/s5873/201702/dda5ffe3f50-941a29fb0aba6233bb497.shtml. )
[83]	Torgerson PR, Shaikenov BS, Rysmukhambetova AT, et al. Modelling the transmission dynamics of Echinococcus granulosus in dogs in rural Kazakhstan[J]. Parasitology, 2003, 126(5): 417-424. doi: 10.1017/S0031182003002932
[84]	Budke CM, Campos-Ponce M, Qian W, et al. A canine purgation study and risk factor analysis for echinococcosis in a high endemic region of the Tibetan Plateau[J]. Vet Parasitol, 2005, 127(1): 43-49. doi: 10.1016/j.vetpar.2004.08.024
[85]	Budke CM, Qiu JM, Craig PS, et al. Modeling the transmission of Echinococcus granulosus and Echinococcus multilocularis in dogs for a high endemic region of the Tibetan Plateau[J]. Int J Parasitol, 2005, 35(2): 163-170. doi: 10.1016/j.ijpara.2004.10.026
[86]	Zeng JX, Wang ZW, Han YC. On the daily activity rhythm of five small mammals[J]. Acta Theriol Sin, 1981, 1(2): 189-197. (in Chinese)
	( 曾缙祥, 王祖望, 韩永才. 五种小哺乳动物活动节律的初步研究[J]. 兽类学报, 1981, 1(2): 189-197.)
[87]	Zong H, Xia WP. Circadian activity rhythms of plateau pikas, Ochotona curzoniae[J]. Acta Theriol Sin, 1987, 7(3): 211-223. (in Chinese)
	( 宗浩, 夏武平. 高原鼠兔似昼夜活动节律的研究[J]. 兽类学报, 1987, 7(3): 211-223.)
[88]	Wan XR, Liu W, Wang GH, et al. Seasonal changes of the activity patterns of Brandt’s vole (Lasiopodomys brandtii) in the typical steppe in Inner Mongolia[J]. Acta Theriol Sin, 2006, 26(3): 226-234. (in Chinese)
	( 宛新荣, 刘伟, 王广和, 等. 典型草原区布氏田鼠的活动节律及其季节变化[J]. 兽类学报, 2006, 26(3): 226-234.)
[89]	Weng XD. Role of Canids in Transmission of Echinococcus spp. on the eastern Tibetan Plateau, China[D]. Shanghai: East China Normal University, 2020: 87-91.
	( 翁晓东. 青藏高原东部牧区犬科动物在棘球蚴病传播中的作用研究[D]. 上海: 华东师范大学, 2020: 87-91.)
[90]	Miller DJ. Restocking and pastoral development among Tibetan nomads after a severe winter on the Tibetan Plateau in western China[J]. Chin Dev Brief, 1998, 1: 1-9.
[91]	Harris RB. Rangeland degradation on the Qinghai-Tibetan Plateau: a review of the evidence of its magnitude and causes[J]. J Arid Environ, 2010, 74(1): 1-12. doi: 10.1016/j.jaridenv.2009.06.014
[92]	Smith AT, Foggin JM. The plateau pika (Ochotona curzoniae) is a keystone species for biodiversity on the Tibetan Plateau[J]. Animal Conserv, 1999, 2(4): 235-240. doi: 10.1111/j.1469-1795.1999.tb00069.x
[93]	Li XL, Gao J, Brierley G, et al. Rangeland degradation on the Qinghai-Tibet Plateau: implications for rehabilitation[J]. Land Degrad Develop, 2013, 24(1): 72-80. doi: 10.1002/ldr.1108
[94]	Wang Q, Vuitton DA, Qiu JM, et al. Fenced pasture: a possible risk factor for human alveolar echinococcosis in Tibetan pastoralist communities of Sichuan, China[J]. Acta Trop, 2004, 90(3): 285-293. doi: 10.1016/j.actatropica.2004.02.004
[95]	Gongbuzeren, Li YB, Li WJ. China’s rangeland management policy debates: What have we learned?[J]. Rangel Ecol Manag, 2015, 68(4): 305-314. doi: 10.1016/j.rama.2015.05.007
[96]	Lai CH, Smith AT. Keystone status of plateau pikas (Ochotona curzoniae): effect of control on biodiversity of native birds[J]. Biodivers Conserv, 2003, 12(9): 1901-1912. doi: 10.1023/A:1024161409110
[97]	Jiang ZG, Xia WP. The niches of yaks, Tibetan sheep and plateau pikas in the alpine meadow ecosystem[J]. Acta Biol Plateau Sin, 1987, 6: 115-146. (in Chinese)
	( 蒋志刚, 夏武平. 高寒草甸生态系统中牦牛、藏系绵羊和高原鼠兔的生态龛(niche)研究[J]. 高原生物学集刊, 1987, 6: 115-146.)
[98]	Ma B, Wang XM, Liu XQ, et al. GIS analysis of the spatial relationship between plateau pika burrow distribution and vegetation distributional patterns[J]. Biodivers Sci, 2011(1): 71-78. (in Chinese)
	( 马波, 王小明, 刘晓庆, 等. 高原鼠兔洞穴数量与其栖息地植被分布格局的GIS分析[J]. 生物多样性, 2011(1): 71-78.) doi: 10.3724/SP.J.1003.2011.08080
[99]	Wilson MC, Smith AT. The pika and the watershed: the impact of small mammal poisoning on the ecohydrology of the Qinghai-Tibetan Plateau[J]. Ambio, 2015, 44(1): 16-22. doi: 10.1007/s13280-014-0568-x
[100]	Wang LM, Yang K, Xie F, et al. Surveillance result of natural foci of Microtus Fuscus in Sichuan Province from 2000 to 2012[J]. Chin J Control Endem Dis, 2013, 28(5): 335-337. (in Chinese)
	( 汪立茂, 杨孔, 谢飞, 等. 四川省2000-2012年青海田鼠鼠疫自然疫源地监测结果分析[J]. 中国地方病防治杂志, 2013, 28(5): 335-337.)
[101]	Pech RP, Arthur AD, Zhang YM, et al. Population dynamics and responses to management of plateau pikas Ochotona curzoniae[J]. J Appl Ecol, 2007, 44(3): 615-624. doi: 10.1111/j.1365-2664.2007.01287.x
[102]	Qiu JM, Liu FJ, Peter S, et al. Epidemiological study on human hydatidosis in Tibetan region of western Sichuan[J]. Chin J Zoonoses, 2000, 16(2): 77-80. (in Chinese)
	( 邱加闽, 刘凤洁, Peter S, 等. 四川西部藏区包虫病流行病学研究: Ⅰ.囊型包虫病与泡型包虫病人群感染特点与分布趋势[J]. 中国人兽共患病杂志, 2000, 16(2): 77-80.)
[103]	Raoul F, Quéré JP, Rieffel D, et al. Distribution of small mammals in a pastoral landscape of the Tibetan Plateaus (Western Sichuan, China) and relationship with grazing practices[J]. Mammalia, 2006, 70(3/4): 214-225.
[104]	Delattre P, Giraudoux P, Baudry J, et al. Land use patterns and types of common vole (Microtus arvalis) population kinetics[J]. Agric Ecosyst Environ, 1992, 39(3/4): 153-168. doi: 10.1016/0167-8809(92)90051-C
[105]	Giraudoux P, Delattre P, Habert M, et al. Population dynamics of fossorial water vole (Arvicola terrestris scherman): a land use and landscape perspective[J]. Agric Ecosyst Environ, 1997, 66(1): 47-60. doi: 10.1016/S0167-8809(97)80706-2
[106]	Craig PS, Larrieu E. Control of cystic echinococcosis/hydatidosis: 1863—2002[J]. Adv Parasitol, 2006, 61: 443-508.
[107]	He W, Shang JY, Chen F, et al. Regularity of canine echinococcosis in epidemic areas in Sichuan[J]. J Prev Med Inf, 2018, 34(2): 136-139. (in Chinese)
	( 何伟, 尚婧烨, 陈凡, 等. 四川省包虫病流行区犬感染棘球绦虫规律性研究[J]. 预防医学情报杂志, 2018, 34(2): 136-139.)
[108]	Jiang B, Zhou XN, Zhang HB, et al. Slow-release praziquantel for dogs: Presentation of a new formulation for echinococcosis control[J]. Infect Dis Poverty, 2017, 6: 140. doi: 10.1186/s40249-017-0357-4 pmid: 28911334
[109]	Schantz PM, Wang H, Qiu J, et al. Echinococcosis on the Tibetan Plateau: prevalence and risk factors for cystic and alveolar echinococcosis in Tibetan populations in Qinghai Province, China[J]. Parasitology, 2003, 127(S1): S109-S120. doi: 10.1017/S0031182003004165
[110]	Jiang CP. Current epidenic status of echinococcosis in China[J]. Chin J Parasit Dis Control, 1996(4): 290-294. (in Chinese)
	( 蒋次鹏. 我国包虫病流行现况[J]. 中国寄生虫病防治杂志, 1996(4): 290-294.)
[111]	Yu SH, Wang H, Wu XH, et al. Cystic and alveolar echinococcosis: an epidemiological survey in a Tibetan population in southeast Qinghai, China[J]. Jpn J Infect Dis, 2008, 61: 242-246.
[112]	Guo L, Yang AG, Zhang ZZ, et al. An epidemiological survey on livestock hydatid diseases in Sichuan Province[J]. Chin J Vet Med, 2012, 48(2): 25-27. (in Chinese)
	( 郭莉, 阳爱国, 张壮志, 等. 四川省家畜包虫病流行病学调查报告[J]. 中国兽医杂志, 2012, 48(2): 25-27.)
[113]	Li K, Shahzad M, Zhang H, et al. Socio-economic burden of parasitic infections in yaks from 1984 to 2017 on Qinghai Tibetan Plateau of China: a review[J]. Acta Trop, 2018, 183: 103-109. doi: 10.1016/j.actatropica.2018.04.011
[114]	Suo L, Wu WP, Dan Z, et al. Epidemiological survey on hydatid disease in livestock in Tibet Autonomous Region[J]. Chin J Parasitol Parasit Dis, 2018, 36(1): 30-34. (in Chinese)
	( 索郎旺杰, 伍卫平, 旦珍旺久, 等. 西藏自治区家畜棘球蚴感染情况调查[J]. 中国寄生虫学与寄生虫病杂志, 2018, 36(1): 30-34.)
[115]	Lin YG, Hong LX, Yang WC, et al. Observations on the natural rodent hosts of alveolar hydatid cyst of Echinococcus multilocularis in Tacheng region, Xingjiang[J]. Endem Dis Bull, 1993, 8(2): 29-33, 123. (in Chinese)
	( 林宇光, 洪凌仙, 杨文川, 等. 新疆塔城地区多房棘球蚴的鼠类宿主考察[J]. 地方病通报, 1993, 8(2): 29-33, 123.)
[116]	Jiang W, Zheng Q, Zheng Q, et al. First discovery of natural infection of metacestode of Echinococcus multilocularis in Microtus ilaeus in nilka County Xinjiang, China[J]. Endem Dis Bull Chin, 2000(1): 36-37, 96. (in Chinese)
	( 将卫, 郑强, 伊斯拉音. 乌斯曼, 等. 新疆尼勒克县首次发现伊犁田鼠感染多房棘球蚴[J]. 地方病通报, 2000(1): 36-37, 96.)
[117]	Fu C, Israyil O, Jiao W, et al. Report on discovery of natural infection of Echinococcus multilocularis metacestode in water vole (Arvicola terrestris Linnaeus, 1758) from bayimuza area of Emin County, Xinjiang, China[J]. Endem Dis Bull Chin, 2001, 16(4): 39-40. (in Chinese)
	( 傅承, 伊斯拉音·乌斯曼, 焦伟, 等. 新疆额敏县首次发现水感染多房棘球蚴[J]. 地方病通报, 2001, 16(4): 39-40.)
[118]	Li WX, Zhang GC, Lin YG, et al. The occurrence of Echinococcus multilocularis Leuckart, 1863 the natural animal host in China and its morphological study[J]. Acta Zool Sin, 1985, 31(4): 365-371. (in Chinese)
	( 李维新, 张国才, 林宇光, 等. 多房棘球绦虫在我国自然动物宿主的发现及其形态学研究[J]. 动物学报, 1985, 31(4): 365-371.)
[119]	Hong LX, Lin YG. Studies on the development and histopathology of alveolar cestode of Echinococcus multilocularis in human and animal hosts[J]. Endem Dis Bull, 1987, 2(2): 51-61, 86. (in Chinese)
	( 洪凌仙, 林宇光. 多房棘球蚴在动物和人体内的发育与组织病变考察[J]. 地方病通报, 1987, 2(2): 51-61, 86.)
[120]	Tang CT, Cui GW, Qian YC, et al. On the occurrence of echinococcosis multilocularis in Hulunbeier pasture, Nei Mongolian Autonomous Region[J]. Acta Zool Sin, 1988, 34(2): 172-179. (in Chinese)
	( 唐崇惕, 崔贵文, 钱玉春, 等. 内蒙古呼伦贝尔草原多房棘球蚴病病原的调查[J]. 动物学报, 1988, 34(2): 172-179.)
[121]	Tang CT, Shu LM, Tang L, et al. The species and bioepiderniology of alveolar Echinococcus in Xinbaerhu West County, eastern Inner Mongolia[J]. J Xiamen Univ Nat Sci, 2001, 40(2): 503-511. (in Chinese)
	唐崇惕, 舒利民, 唐亮, 等. 内蒙古东部新巴尔虎右旗泡状肝包虫病原种类及流行学调查[J]. 厦门大学学报(自然科学版), 2001, 40(2): 503-511.)
[122]	Tang CT, Tang L, Kang YM, et al. Investigation on the alveolar hydatids in rodents in Ewenke County pasture, eastern Inner Mongolia[J]. Acta Parasitol Med Entomol Sin, 2001, 8(4): 220-226. (in Chinese)
	( 唐崇惕, 唐亮, 康育民, 等. 内蒙古东部鄂温克旗草场鼠类感染泡状棘球蚴情况的调查[J]. 寄生虫与医学昆虫学报, 2001, 8(4): 220-226.)
[123]	Cai QG, Han XM, Yang YH, et al. Lasiopodomys fuscus as an important intermediate host for Echinococcus multilocularis: isolation and phylogenetic identification of the parasite[J]. Infect Dis Poverty, 2018, 7: 27. doi: 10.1186/s40249-018-0409-4
[124]	Wang W, Wu Y, Wu JG, et al. The occurence of Echinococcus multilocularis leukart, 1863 in fox and wolf in Tacheng District, Xinjiang[J]. Endem Dis Bull, 1989, 4(2): 8-11, 89. (in Chinese)
	( 王伟, 吴勇, 吴季高, 等. 新疆塔城地区多房棘球绦虫调查: 国内狼体多房棘球绦虫新记录[J]. 地方病通报, 1989, 4(2): 8-11, 89.)
[125]	Jiang WB. An epidemiologic study on the infection rates of Echinococcus spp. in Tibetan fox population[D]. Shanghai: East China Normal University, 2012: 81-82. (in Chinese)
	( 蒋韦斌. 藏狐群体棘球绦虫感染率的流行病学研究[D]. 上海: 华东师范大学, 2012: 81-82.)
[126]	Han XM, Wang H, Cai HX, et al. Epidemiological survey on echinococcosis in darlag County of Qinghai Province[J]. Chin J Parasitol Parasit Dis, 2009, 27(1): 22-26. (in Chinese)
	( 韩秀敏, 王虎, 蔡辉霞, 等. 青海省达日县棘球蚴病流行病学调查[J]. 中国寄生虫学与寄生虫病杂志, 2009, 27(1): 22-26.)
[127]	Tang CT, Cui GW, Qian YC, et al. Studies on the alveolar Echinococcus species in northward daxingan mountains, Inner Mongolia, China. Ⅰ. Echinococcus multilocularis Leuckart, 1863[J]. Chin J Zoonoses, 2006, 22(12): 1089-1094. (in Chinese)
	( 唐崇惕, 崔贵文, 钱玉春, 等. 我国内蒙古大兴安岭北麓泡状肝包虫种类的研究Ⅰ. 多房棘球绦虫[J]. 中国人兽共患病学报, 2006, 22(12): 1089-1094.)