Skip to Main Content
San Diego Zoo Wildlife Alliance logo
San Diego Zoo Wildlife Alliance Library logo

Przewalski's Horse (Equus ferus przewalskii) Fact Sheet: Bibliography & Resources

Przewalski's Horse (Equus ferus przewalskii)


Agusti J, Anton M. 2005. Mammoths, sabertoths, and hominidis: 65 million years of mammalian evolution in Europe. New York: Columbia University Press.

Asa, C.S., Equid Reproductive Biology IN  Moehlman, P.D. (ed). 2002. Equids: Zebras, Asses, and Horses: Status Survey and Conservation Action Plan. IUCN/SCC Equid Specialist Group, IUCN  : Gland, Switzerland.  Chapter 10, pp 113-117.

Austin NP, Rogers LJ. 2014. Lateralization of agonistic and vigilance responses in Przewalski horses (Equus przewalskii). Applied Animal Behaviour Science. 151:43–50. doi:10.1016/j.applanim.2013.11.011.
AZA. 2023. Species Survival Statistics [data table report]. Chicago (IL): Lincoln Park Zoo/AZA Population Management Center; [accessed 2023 Apr 18].
Benirschke, K. et al (1965) Chromosome complement: difference between Equus caballus and Equus przewalskii Poliakoff.  Science, 148:382-383. 

Bennett, D. & Hoffmann, R.S. (1999). Equus Caballus. Mammalian Species 628, pp. 1-14.

Bourjade M, Thierry B, Maumy M, Petit O. 2009. Decision-making in Przewalski horses (Equus ferus przewalskii) is driven by the ecological contexts of collective movements. Ethology. 115(4):321–330. doi:10.1111/j.1439-0310.2009.01614.x.


Boyd, L. and Houpt, K.A. eds (1994). Przewalski’s Horse : The history and biology of an endangered species. Albany: State University of New York. 

Boyd L, King SRB. 2011. Equus ferus ssp. przewalskii. In: IUCN 2011. IUCN Red lList of Threatened Species. Version 2011.2 Retrieved January 4 2012 from

Bouman, J. (1986). Particulars about the Przewalski Horse.  Foundation for the Preservation and Protection of the Przewalski Horse.

Bowling, A. and A. Ruvinsky 2000. Genetic Aspects of Domestication, Breeds and Their Orgins. In: Bowling, A. and A. Ruvinsky (eds) Genetics of the Horse. CABI International Publishing, UK and USA

Breining, G (2006) The wildest horse; China fulfills a bold dream to restore “P-horses” to the Gobi Desert.  Wildlife Conservation July/August.

Collins CW. 2010. Understanding the reproductive biology of the Przewalski’s horse (Equus ferus przewalskii) [Ph.D. thesis]. [College Park (MD)]: University of Maryland.
Collins CW, Songsasen NS, Vick MM, Wolfe BA, Weiss RB, Keefer CL, Monfort SL. 2012. Abnormal reproductive patterns in Przewalski’s mares are associated with a loss in gene diversity. Biology of Reproduction. 86(2). doi:10.1095/biolreprod.111.092676.
D’Souza-Anjo M, Christensen BW, Brabender K, Zimmermann W, Kass PH, Schwarzenberger F. 2017. A comparison of fecal steroid metabolite concentrations between harem and bachelor stallions in a free-ranging population of Przewalski’s horses (Equus ferus przewalskii). Zoo Biology. 36(2):127–131. doi:10.1002/zoo.21348.
Der Sarkissian C, Ermini L, Schubert M, Yang MA, Librado P, Fumagalli M, Jónsson H, Bar-Gal GK, Albrechtsen A, Vieira FG, et al. 2015. Evolutionary genomics and conservation of the endangered Przewalski’s horse. Current Biology. 25(19):2577–2583. doi:10.1016/j.cub.2015.08.032.
Fedorov NI, Mikhailenko OI, Zharkikh TL, Bakirova RT. 2018. Mapping of vegetation with the geoinformation system and determining of carrying capacity of the pre-Urals steppe area for a newly establishing population of the Przewalski horse Equus ferus przewalskii at the Orenburg State Nature Reserve. IOP Conf Ser: Earth Environ Sci. 107:012100. doi:10.1088/1755-1315/107/1/012100.

Fisher T, Faliano A, Willis K. 2019. Asian wild horse (Equus ferus prezwalskii) 2020 final breeding and transfer plan. Silver Spring: Association of Zoos & Aquariums. [accessed 2020 Dec 01 from].

Forstén, A. 1988. Middle Pleistocene replacement of Stenoid horses by Caballoid horses - ecological implications. Palaeogeography, Palaeoclimatology, Palaeoecology 65:23-33.

Froehlich, D. 2002. Quo vaids eohippus? The systematics and taxonomy of the early Eocene equids (Perissodactyla). Zoological Journal of the Linnean Society 134(2):141-256.

Gashchak S, Paskevych S. 2019. Przewalski`s horse (Equus ferus przewalskii) in the Chornobyl Exclusion Zone after 20 years of introduction. Theriol Ukr. 2019:80–100. doi:10.15407/pts2019.18.080.


Goto H, Ryder OA, Fisher AR, Schultz B, Pond SLK et al. 2011. A massively parallel sequencing approach uncovers ancient origins and high genetic variability of endangered Przewalski's horses. Genome Biology and Evolution. Retrieved January 3, 2012 from doi:10:1093/gbe/ev067

Groves, C.P. (1986)  The taxonomy, distribution and adaptations of recent equids, IN Equids in the Ancient World (eds. Meadow, R.H. and Uerpmann, H-P. ) Wiesbaden: Dr. Ludwig Reichert Verlag. Pp 11-65

Groves, C.P. (1994) Morphology, habitat and taxonomy, In: Przewalski’s Horse:  The History and Biology of an Endangered species (Eds. Boyd, L. and Houpt, K.A.)  Albany : State University of New York Press pp 39-59.

Groves, C.P. (2002) Taxonomy of Living Equidae IN: Moehlman, P.D. (ed). 2002. Equids: Zebras, Asses, and Horses: Status Survey and Conservation Action Plan. IUCN/SCC Equid Specialist Group, IUCN  : Gland, Switzerland.  Chapter 8, pp 94-107

Groves, C.P. and O.A. Ryder 2000. Systematics and phylogeny of the horse. In: Bowling, A. and A. Ruvinsky (eds) Genetics of the Horse. CABI International Publishing, UK and USA.

Hooker, J. 2008. The beginning of the equoid radiation. Zoological Journal of the Linnean Society 112:(1) 29-63.

Kaczensky P, Burnik Šturm M, Sablin MV, Voigt CC, Smith S, Ganbaatar O, Balint B, Walzer C, Spasskaya NN. 2017. Stable isotopes reveal diet shift from pre-extinction to reintroduced Przewalski’s horses. Sci Rep. 7(1):5950. doi:10.1038/s41598-017-05329-6.


King, S.R.B., Boyd, L., Zimmermann, W. and Kendall, B.E. 2015. Equus ferus. The IUCN red list of threatened species [Internet]. Version 2018-2; e.T41763A97204950 [species assessed 2014 Oct 09; page accessed 2018 Dec 18].

Kaleta T, Osija A, Stokowska M, Świderek W. 2017. Preliminary observations of the behaviour in two wild equids: Przewalski horse (Equus ferus przewalskii) and Hartmann’s zebra (Equus zebra hartmannae) kept in socially changed groups at Warsaw Zoo. Annals of Warsaw University of Life Sciences - SGGW - Animal Science. 56(2):249–255. doi:10.22630/AAS.2017.56.2.26.


Klingel, H.  1990.  Horses.  In S. Parker, ed. Grzimeks's Encyclopedia Mammals Volume 4. McGraw-Hill Publishing Company: New York, p568.

Lau AN, Peng L, Goto H, Chemnick L, Ryder OA, Makova KD. 2009. Horse domestication and conservation genetics of Przewalski's horse inferred from sex chromosomal and autosomal sequences. Molecular Biology and Evolution 26(1):199-208.

Li Y, Zhang K, Liu Y, Li K, Hu D, Wronski T. 2019. Community composition and diversity of intestinal microbiota in captive and reintroduced przewalski’s horse (Equus ferus przewalskii). Front Microbiol. 10:1821. doi:10.3389/fmicb.2019.01821.

Moehlman, P. (2005). Endangered Wild Equids. Scientific American, March 2005, pp. 86-93.

Maigrot A-L, Hillmann E, Anne C, Briefer EF. 2017. Vocal expression of emotional valence in Przewalski’s horses (Equus przewalskii). Sci Rep. 7(1):8779. doi:10.1038/s41598-017-09437-1.

Mohr, E. (1971).  The Asiatic Wild Horse Equus przewalskii Poliakoff, 1881,  London:  J.A. Allen & Co. Ltd. 124 pp.

Monfort, S.L., Arthur, N.P. and Wildt, D.E. (1991) monitoring ovarian function and pregnancy by evaluating excretion of urinary oestrogen conjugates in semi-free-ranging Przewalski’s horses (Equus przewalskii)  J Reprod. Fert. 91, 155-164.

National Geographic [Copyright 1996-2018] Przewalski's Horse. Visited 2018 Dec 18.

Naundrup PJ, Svenning J-C. 2015. A geographic assessment of the global scope for rewilding with wild-living horses (Equus ferus). Festa-Bianchet M, editor. PLoS ONE. 10(7):e0132359. doi:10.1371/journal.pone.0132359.

Oakenfull, E.A., H. Lim, O. Ryder 2000. A survey of equid mitochondrial DNA: implications for the evolution, genetic diversity and conservation of Equus. Conservation Genetics 1(4):341-355.

Okenfull, E.A. and Ryder O.A. 2002. Genetics of Equid Species and Subspecies, Chapter 9 In: Moehlman, P.D. (Ed.). Equids: Zebras, Asses, and Horses: Status Survey and Conservation Action Plan. IUCN/SCC Equid Specialist Group, IUCN : Gland, Switzerland Chapter 9, pp. 108-112.

Orlando L, Ginolhac A, Zhang G, Froese D, Albrechtsen A, et al. 2013. Recalibrating Equus evolution using the genome sequence of an early Middle Pleistocene horse. Nature 499:74-78. doi:10.1038/nature12323.

Radcliffe, R.M. and Osofsky, Disease Concerns for wild Equids IN Moehlman, P.D. (ed). 2002. Equids: Zebras, Asses, and Horses: Status Survey and Conservation Action Plan. IUCN/SCC Equid Specialist Group, IUCN: Gland, Switzerland.  Chapter 12, Pp 124-153

Rademacher, U., Langenhorst, T., Simek J., Zimmermann, M., Van der Meer, R. and Voorham, M. (eds.). 2020. Regional collection plan for the EAZA Equid Taxon Advisory Group – first edition. Amsterdam: EAZA Executive Office.

Robert, N., et al. (2005). Pathologic Findings in Reintroduced Przewalski’s Horses in Southerwestern Mongolia. Journal of Zoo and Wildlife Medicine 36, pp. 273-285.

Ryder, O. (1990). Putting the wild horse back into the wild. Przewalski’s Horse Global Conservation Plan. Zoological Society of San Diego, Center for Reproduction of Endangered Species.

Ryder, O. (1993). Przewalski’s horse: Prospects for reintroduction into the wild. International Conservation News 7, pp. 13-15.

Ryder, O. 2009. Rhinoceroses, tapirs, and horses (Perissodactyla). in The Timetree of Life. Hedges, S. and S. Kumar (Eds.) Oxford University Press.

Schiebe, K.M., et al (1998) Water consumption and watering behavior of Przewalski Horses (Equus ferus przewalskii) in a Semi reserve.  Zoo Biology 17:181-192.

Schlichting PE, Dombrovski V, Beasley JC. 2020. Use of abandoned structures by Przewalski’s wild horses and other wildlife in the Chernobyl Exclusion Zone. Mamm Res. 65(1):161–165. doi:10.1007/s13364-019-00451-4.
Sim RR, Joyner PH, Padilla LR, Anikis P, Aitken-Palmer C. 2017. Clinical disease associated with Anaplasma phagocytophilum infection in captive Przewalski’s horses (Equus ferus przewalskii). Journal of Zoo and Wildlife Medicine. 48(2):497–505. doi:10.1638/2016-0247R.1.


Steiner CC, Ryder OA 2011. Moledcular phylogeny and evolution of the Perissodactyla. Zoological Journal of the Linnean Society 163L1289-1303.

Tang L, Xiu Y, Yan L, Cui Y, Ma X, Ente M, Zhang Y, Li K, Zhang D. 2020. Drug efficacy of ivermectin against primary nematodes parasitizing captive Przewalski’s horse (Equus ferus przewalskii) after ten years of annually treatment. Helminthologia. 57(1):57–62. doi:10.2478/helm-2020-0004.
Tarav M, Tokunaga M, Kondo T, Kato-Mori Y, Hoshino B, Dorj U, Hagiwara K. 2017. Problems in the protection of reintroduced Przewalski’s horses (Equus ferus przewalskii) caused by piroplasmosis. Journal of Wildlife Diseases. 53(4):911–915. doi:10.7589/2017-02-024.
Tatin L, King SRB, Munkhtuya B, Hewison AJM, Feh C. 2009. Demography of a socially natural herd of Przewalski’s horses: an example of a small, closed population. Journal of Zoology. 277(2):134–140. doi:10.1111/j.1469-7998.2008.00520.x.


UNEP. 2018. Species+: Equus ferus [online database]. Nairobi, Kenya: UNEP [compiled by UNEP-WCMC, Cambridge, UK]. [accessed 2018 Dec 18]. 

Vaughan, T.A., J.M. Ryan, & N.J. Czaplewski. (2000). Mammalogy.  Philadelphia: Saunders College Publishing.

Vilà, C., Leonard, J.A., Gotherstrom, A., Marklund, S., Sandberg, K., Lide, K., Wayne, R.K. and Ellegren, H. 2001. Widespread Origins of Domestic Horse Lineages. Science 291: 474-477.

Wakefield, S., et al. (2006). Equids: Zebras, Asses and Horses. Status Survey and Conservation Action Plan. Cambridge: IUCN. Chapter 7 pp 82-92

Walzer, C., et al. (2000) Medical aspects of Przewalski horse (Equus przewalskii) reintroduction to the Dsungarian Gobi, Mongolia. in Proceedings of the AAZV and IAAAM Joint Conference. 2000. New Orleans, Louisiana. pp 17-21.

Xia C, Cao J, Zhang H, Gao X, Yang W, Blank D. 2014. Reintroduction of Przewalski’s horse (Equus ferus przewalskii) in Xinjiang, China: the status and experience. Biological Conservation. 177:142–147. doi:10.1016/j.biocon.2014.06.021.

Zimmermann, W. (2005) [Przewalski’s horses on the track to reintroduction – various projects compared], Zeitschrift des Kolner Zoo 48 (4): 183-209. [PDF]

Zimmermann, W. (2011). International Przewalski's Horse Studbook (Equus ferus przewalskii). Cologne Zoo.

ZIMS (Zoological Information Management System). 2020. [accessed 2020 Dec 10].

Additional Internet Resources


On the Lookout

Two Przewalski's horses stand alert, facing viewer

Przewalski's horses at the San Diego Zoo Safari Park.

Image credit: © San Diego Zoo Wildlife Alliance. All rights reserved.

SDZWA Library Links