dr. sarah king
Find out more about my research!
My research focuses on how understanding the behavioral ecology of mammals can inform management of common species and conservation of endangered species.
I am a Research Scientist at the Natural Resource Ecology Laboratory at Colorado State University and Co-Chair of the International Union for the Conservation of Nature (IUCN) Species Survival Commission Equid Specialist Group.
I gained my B.Sc. and Ph.D. from Queen Mary, University of London. I have conducted research on small mammals in the U.S. and led conservation projects in Mongolia, but most of my career has been spent studying the behavior and ecology of wild and feral equids. My Ph.D. was one of the first studies on the behavioral ecology of Przewalski’s horses after their reintroduction to the wild in central Mongolia, and later I was Project Manager of a reintroduction effort of Przewalski’s horses in western Mongolia. For the last 10 years I worked in collaboration with USGS to lead behavioral aspects of research into feral horses and burros in the American west.
Scroll down this page for a description of my research topics, with associated publications
Many of my articles are open access so can be downloaded at no cost. Most can be dowloaded from my ResearchGate page or found on my Google Scholar page if not linked here.
Please contact me for reprints if they are not publicly available.
Behavior and ecology of free-roaming horses and donkeys in the American west
Management of free-roaming horses in the American west is contentious, and can be informed with better knowledge of the behavior, ecology, and demography of free-roaming horses and donkeys in the American west. Behavior plays an essential role in survival and reproductive success as it modulates interactions between individuals and their environment, so ultimately affects the growth and persistence of populations. Exploring interactions between behavior and demography through use of fertility control therefore enables us to understand both how it affects individuals and populations. Further, understanding how animals use the landscape across space and time can enable better management of growing populations to protect the habitat.
This work was conducted in collaboration with USGS Fort Collins Science Center.
Esmaeili, S., Schoenecker, K. & King, S.R.B. 2024. ‘Resource availability and heterogeneity affect space use and resource selection of a feral ungulate’. Ecological Applications. In Press.
King, S.R.B., Cole, M., Barton, C., & Schoenecker, K. 2023 ‘Proximate factors affecting mortality and maternal abandonment of young free-roaming feral horse foals’. Journal of Veterinary Behavior. Online early.
Schoenecker, K., Esmaeili, S. & King, S.R.B. 2023. ‘Seasonal resource selection and movement ecology of free-ranging horses in the Western USA’. Journal of Wildlife Management. 87: e22341.
Bechert, U. S., Turner, J. W., Baker, D. L., Eckery, D. C., Bruemmer, J., Lyman, C. C., Prado, T, King, S.R.B., & Fraker, M. 2022. ‘Fertility control options for management of free-roaming horse populations’. Human-Wildlife Interactions. 16: 179-216.
King, S.R.B., Schoenecker, K., & Cole, M. 2022. ‘Effect of adult male sterilization on the behavior and social associations of a feral polygynous ungulate: the horse’. Applied Animal Behaviour Science. 249: 105598.
Schoenecker, K., King, S.R.B., & Messmer, T.A. 2021. ‘The wildlife profession’s duty in achieving science-based sustainable management of free-roaming equids’. Journal of Wildlife Management. 85(6): 1057-1061.
Telemetry devices: radio collars and tail tags
Individual-based data is invaluable for elucidating fine-scale ecology and enabling animals to be found efficiently for behavioral research. Understanding wild horse and donkey behavior allows management to be put in an ecological context. As radio collars had not been used on free-roaming horses for almost 30 years, we first conducted a pasture study to test their safety. At the same time we developed a novel method for fitting telemetry devices in horse tails.
King, S.R.B. & Schoenecker, K. 2022. ‘Application of tail transmitters for tracking feral horses as an alternative to radio collars. Wildlife Society Bulletin. 46(4): e1338.
Hennig, J., Scasta, J.D., Beck, J., Schoenecker, K., & King, S.R.B. 2020. ‘Systematic Review of Equids and Telemetry Collars: Implications for Deployment and Reporting’. Wildlife Research. 47: 361-371.
Schoenecker, K., King, S.R.B. & Collins, G. 2020. ‘Evaluation of the impacts of radio-marking devices on feral horses and burros in a captive setting’. Human-Wildlife Interactions. 14(1):73-86.
The secrets contained in poop!
Feces is an invaluable resource! And horses leave it everywhere. We used fecal sampling to determine genetic parameters of a population of free-roaming horses (Little Book Cliffs HMA, CO) to explore any genetic effects of long-term population growth suppression with fertility control, finding that it had minimal effect. We also tested whether fecal sampling could be used to estimate population size as a potential alternative to aerial surveys. As well as genetic information, feces can tell you about what animals have eaten, so we additionally explored horse and burro diets and if horses disseminated weeds.
‘Stud piles’ are large fecal piles where stallions repeatedly deposit, yet their function was not well known before I conducted a study on fecal marking in Przewalski’s horse stallions, concluding that these piles act like sign posts alerting stallions to who else was in the area. I then collaborated with colleagues from the Czech Republic to dive deeper into the function of this fascinating behavior in 4 species of wild equid maintained in captivity. This allowed us to analyze fine-scale data from equid species with different social systems.
Esmaeili, S., Schoenecker, K. & King, S.R.B. 2023. ‘Browsers or grazers? New insights into feral burro diet using a non-invasive sampling and plant DNA metabarcoding approach’. Animals. 13: 2683.
Pluháček, J., Tučková, V. & King, S.R.B. 2022. ‘Not just for males: flehmen as a tool for detection of reproductive status and individual recognition across sexes in four African equid species’. Behavioural Processes. 203: 104773.
Pluháček, J., Tučková, V., Šárová, R. & King, S.R.B. 2022. ‘Why wait to mark? Possible reasons behind latency from olfactory exploration to overmarking in four African equid species’. Animal Cognition. Online early.
Schoenecker, K. A., Oyler-McCance, S., Ekernas, L., King, S.R.B. 2021. ‘Using fecal DNA and closed-capture models to estimate feral horse population size’. Journal of Wildlife Management. 85(6): 1150-1161
King, S.R.B., Schoenecker, K. A., Fike, J., & Oyler-McCance, S. 2021. ‘Feral horse space use and genetic characteristics from fecal DNA’. Journal of Wildlife Management. 85(6): 1074-1083
Pluháček, J., Tučková, V., King, S.R.B., & Šárová, R. 2020. ‘Effect of social organisation on interspecific differences in overmarking behaviour of foals in African equids’. Animal Cognition. 23: 131-140
Pluháček, J., Tučková, V. & King, S.R.B. 2019. ‘Overmarking behaviour of zebra males: no scent masking, but a group cohesion function’. Behavioural Ecology & Sociobiology. 73: 136-147.
Pluháček, J., Tučková, V., King, S.R.B., & Šárová, R. 2019. ‘Test of four hypotheses to explain the function of overmarking in foals of four equid species’. Animal Cognition. 22: 231-241.
King, S.R.B. & Schoenecker, K. 2019. ‘Comparison of methods to examine diet of feral horses from non-invasively collected fecal samples’. Rangeland Ecology and Management. 72: 661-666.
King, S.R.B., Schoenecker, K. & Manier, D. 2019. ‘Potential spread of cheatgrass (Bromus tectorum) and other invasive species by feral horses (Equus ferus caballus) in western Colorado’. Rangeland Ecology and Management. 72: 706-710.
King, S.R.B., Schoenecker, K., Fike, J., & Oyler-McCance, S. 2018. ‘Long-term persistence of horse fecal DNA in the environment makes equids particularly good candidates for non-invasive sampling’. Ecology and Evolution. 8: 4053-4064.
Tučková, V., R. Sarova, J. Bartošová, King, S.R.B. & Pluháček, J. 2018. Overmarking by adult females in four equid species: social bonds and group cohesion. Journal of Zoology. 306:180-188.
Przewalski’s horse research
For my doctoral thesis I conducted the first study of the behavioral ecology of Przewalski’s horses in the wild at Hustai National Park. At the time, the species was recently reintroduced to Mongolia after being extinct in the wild for 20 years (kept extant as a species thanks to captive breeding in zoos). My thesis work involved exploring the behavior and movements of Przewalski’s horses to assess how they were adapting to life in the wild.
I subsequently was project manager for Association TAKH, which led a reintroduction to Khomyn Tal in western Mongolia, and had a source population of Przewalski’s in southern France at Le Villaret. We examined demographic information over multiple generations, and how male behavior affects their reproductive fitness in the Le Villaret population to inform conservation of this endangered species, as well as inform welfare of domestic horses.
King, S.R.B. & Gurnell, J. 2019. ‘Associative behaviour in Przewalski’s horses reintroduced into Mongolia’. Nature Conservation Research. 4(Suppl.2): 1-9.
King, S.R.B. 2013 ‘Przewalski’s horses and red wolves: importance of behavioral research for species brought back from the brink of extinction’. Pp:153-158 in: Bekoff, M. (ed). Ignoring Nature No More: The Case for Compassionate Conservation. University of Chicago Press, Chicago, USA.
Bourjade, M., Tatin, L., King, S.R.B. & Feh, C. 2009. ‘Early reproductive success, preceding bachelor ranks and their behavioral correlates in young Przewalski’s stallions’. Ethology, Ecology & Evolution. 21: 1-14.
Tatin, L., King, S.R.B., Munkhtuya, B., Hewison, A.J.M. & Feh, C. 2009. ‘Demography of a socially natural herd of Przewalski’s horses: an example of a small, closed population’. Journal of Zoology. 277: 134-140.
King, S.R.B. & Gurnell, J. 2007. ‘Scent-marking behaviour of stallions: an assessment of function in a reintroduced population of Przewalski horses (Equus ferus przewalskii)’. Journal of Zoology. 272: 30-36.
King, S.R.B. 2006. ‘Extinct in the Wild to Endangered: the history of Przewalski’s horse (Equus ferus przewalskii) and its future conservation’. Mongolian Journal of Biological Sciences. 3: 37-41
King, S.R.B. & Boyd, L. 2006. ‘Behavioural ecology of the takhi’. In: Wit, P. & Bouman, I. The tale of the Przewalski’s horse. Coming home to Mongolia. pp: 124-127. KNNV Publishing, Utrecht, Netherlands.
King, S.R.B. & Gurnell, J. 2005. ‘Habitat use and spatial dynamics of takhi introduced to Hustai National Park, Mongolia’. Biological Conservation. 124: 277-290.
King, S.R.B. 2002. ‘Home range and habitat use of free-ranging Przewalski horses at Hustai National Park, Mongolia.’ Applied Animal Behaviour Science. 78: 103-113
Wild equids and conservation
My work serving IUCN/SSC as a co-chair of the Equid Specialist Group, and work as a project manager for two international conservation-focussed non-profits provide me with the specialization to co-author papers related to mammal and equid conservation. I was also invited to lead one chapter and co-author 2 chapters of a recent seminal book: ‘Wild Equids: Ecology, Management, and Conservation.’
I obtained funding for and co-wrote the first ‘Field Guide to Mammals of Mongolia’, which is now in its third edition.
Esmaeili, S., Mahmoud-Reza, H., Kaczensky, P., Schoenecker, K., King, S.R.B., Shahriari, B., Walzer, C., Goheen, J. 2024. ‘Rainfall reduces the potential for competitive suppression of a globally endangered ungulate by livestock’. Biological Conservation. 292: 110476.
Pacific, M., Cristiano, A., Lumbierres, M. … King, S.R.B. … Wiesel, L. 2023. ‘Drivers of habitat availability for terrestrial mammals’. Global Change Biology. In press.
Esmaeili, S., Jesmer, B. R., Albeke, S. E., Aikens, E. O., King, S.R.B., et al. 2021. ‘Body size and digestive system shape resource selection by ungulates: a cross-taxa test of the Forage Maturation Hypothesis’. Ecology Letters. 24(10):2178-2191.
Bolam, F., Mair, L., Angelico, M., Brooks,T., Burgman, M., Hermes, C., Hoffmann, M., … King, S.R.B., et al. 2020. ‘How many bird and mammal extinctions has recent conservation action prevented?’. Conservation Letters. e12762: 1-11.
McGowan, P., Mair, L., Symes, A., Westrip, J., Wheatley, H., Brook, S., Burton, J., King, S., McShea, W., Moehlman, P., Smith, A., Wheeler, J., & Butchart, S. 2018 ‘Tracking trends in the extinction risk of wild relatives of domesticated species to assess progress against global biodiversity targets’. Conservation Letters. e12588.
Crooks, K.R., Burdett, C.L., Theobald, D.M., King, S.R.B., Di Marco, M., Rondinini, C., & Boitani, L. 2017. ‘Quantification of habitat fragmentation reveals extinction risk in terrestrial mammals’. Proceedings of the National Academy of Sciences. 114: 7635-7640.
King, S.R.B., Asa, C., Pluháček, J., Houpt, K. & Ransom, J. 2016. ‘The behavior of horses, zebras, and asses’. Chapter 3 In: Ransom, J. & Kaczensky, P. (eds). Wild Equids: Ecology, Management, and Conservation. Johns Hopkins University Press.
Schoenecker, K.A, King, S.R.B., Nordquist, M., Dejid, N., & Cao, Q. 2016. ‘Habitat and diet of equids’. Chapter 4 In: Ransom, J. & Kaczensky, P. (eds). Wild Equids: Ecology, Management, and Conservation. Johns Hopkins University Press.
Moehlman, P., King, S.R.B., & Kebede, F. 2016. ‘Conservation of threatened wild equids’. Chapter 12 In: Ransom, J. & Kaczensky, P. (eds). Wild Equids: Ecology, Management, and Conservation. Johns Hopkins University Press.
Batsaikhan, N., Samiya, R., Shar, S. & King, S.R.B. 2014. A Field Guide to the Mammals of Mongolia. Second Edition. 307pp. Zoological Society of London, London.
Olléová, M., Pluháček, J. & King, S.R.B. 2012. ‘Effect of social system on allosuckling and adoption in zebras’. Journal of Zoology.288: 127-134.
King, S.R.B. 2007. ‘Asiatic wild asses in the literature: what do we need to know now?’. In: Stubbe, A., Kaczensky, P., Wesche, K., Samija, R. & Stubbe, M. (eds). Exploration into the Biological Resources of Mongolia, 10. pp: 347-357. Martin-Luther-Universitat Halle Wittenberg, Halle (Saale), Germany.
Squirrels and small mammals
I examined the behavioral ecology of the endangered Mt. Graham red squirrel as a wildlife biologist in the Koprowski lab at the University of Arizona. This squirrel is only found on one mountain in Arizona, so my work focused on exploring its spatial and feeding ecology to inform its conservation.
I was a post-doctoral research associate in the McCain lab at the University of Colorado determining the elevational distribution of small mammals in two ranges of the Rocky Mountains to examine effects of climate change in relation to historical populations. By understanding how species’ elevational and latitudinal movements have changed over time, we can try to predict future changes as the climate warms.
McCain, C., King, S.R.B. & Szewczyk, T. 2021. ‘Unusually large upward shifts in cold-adapted, montane mammals as temperature warms’. Ecology. 102(4): e03300
McCain, C., King, S.R.B., Szewczyk, T., & Beck, J. 2018. ‘Small mammal species richness is directly linked to regional productivity, but decoupled from food resources, abundance, or habitat complexity’. Journal of Biogeography. 45:2533-2545.
King, S.R.B. & McCain, C.M. 2015. ‘Robust discrimination of Reithrodontomys megalotis and R. montanus (Mammalia: Rodentia) from Colorado, using cranial morphology and external characteristics within age classes’. Proceedings of the Biological Society of Washington. 128: 1-10
McCain, C.M. & King, S.R.B. 2014. ‘Body size and activity times mediate mammalian responses to climate change’. Global Change Biology. 20:1760-1769.
King, S.R.B. & Koprowski, J.L. 2009. ‘Human and non-human disturbance of Mt. Graham red squirrels’. In: Sanderson, H.R. & Koprowski, J.L. (eds). The Last Refuge of the Mt. Graham Red Squirrel. Ecology of Endangerment. University of Arizona Press, Tucson, USA.
Koprowski, J.L., King, S.R.B. & Merrick, M. 2007. ‘Expanded home ranges in a peripheral population: space use by endangered Mt. Graham red squirrels’. Endangered Species Research. 3: 105-110.
Isaac, J.L., Jamsranjav, J., Simpson, L. & King, S.R.B. 2006. Small mammals of the Mongolian mountain steppe region near Erdensant: insights from live-trapping and bird pellet remains. Mongolian Journal of Biological Sciences. 3: 43-49.
IUCN Red List assessments and other work
As part of my role as co-chair of the IUCN/SSC Equid Specialist Group and Coordinator of the Equid Red List Authority for IUCN I coordinate and/or co-author Red List assessments for all equid species. I also co-organized the Regional Assessment of the threatened status of various taxa in Mongolia for Zoological Society of London and IUCN.
Gosling, M., Muntifering, J., Kolberg, H., Uiseb, K., & King, S.R.B. 2019. ‘Equus zebra’. Equus zebra. The IUCN Red List of Threatened Species 2010. e.T7960A45171906.
Gosling, M., Muntifering, J., Kolberg, H., Uiseb, K., & King, S.R.B. 2019. ‘Equus zebra ssp. hartmannae’. The IUCN Red List of Threatened Species 2019. e.T7958A45171819.
Rubenstein, D., Low Mackey, B., Davidson, Z.D., & King, S.R.B. 2016. ‘Equus grevyi’. The IUCN Red List of Threatened Species 2016: e.T7950A89624491.
King, S.R.B., & Moehlman, P.D. 2016. ‘Equus quagga’. The IUCN Red List of Threatened Species 2016: e.T41013A45172424.
Hrabar, H., Birss, C., Peinke, D., King, S., Novellie, P., Kerley, G. & Child, M. 2016. A Conservation Assessment of Equus zebra zebra. In: M.F. Child, L. Roxburgh, E. Do Linh San, D. Raimondo, and H. Davies-Mostert (eds), The Red List of Mammals of South Africa, Swaziland and Lesotho. South African National Biodiversity Institute and Endangered Wildlife Trust, South Africa.
Novellie, P., King, S., Munifering, J., Uiseb, K., & Child, M.F. 2016. A Conservation Assessment of Equus zebra hartmannae. In: M.F. Child, L. Roxburgh, E. Do Linh San, D. Raimondo, and H. Davies-Mostert (eds), The Red List of Mammals of South Africa, Swaziland and Lesotho. South African National Biodiversity Institute and Endangered Wildlife Trust, South Africa.
King, S.R.B., Boyd, L., Zimmermann, W. & Kendall, B.E. 2015. ‘Equus ferus’. The IUCN Red List of Threatened Species 2015: e.T41763A97204950.
Boyd, L. & King, S.R.B. 2011. ‘Equus ferus’. In: IUCN 2011. IUCN Red List of Threatened Species. Version 2011.2.
Boyd, L., Zimmermann, W. & King, S.R.B. 2008. Equus ferus ssp. przewalskii. In: IUCN 2008. 2008 IUCN Red List of Threatened Species.
Clark, E.L., Munkhbat, J., Dulamtseren, S., Baillie, J.E.M., Batsaikhan, N., King, S.R.B., Samiya, R. & Stubbe, M. (compilers and editors). 2006. Summary Conservation Action Plans for Mongolian Mammals. Regional Red List Series Vol. 2. Zoological Society of London, London.
Clark, E.L., Ocock, J.F., King, S.R.B. & Baillie, J.E.M. 2006. ‘Proceedings of the Mongolian Biodiversity Databank Workshop:Assessing the Conservation Status of Mongolian Mammals and Fishes. I - Results and Outputs of the Workshop’. Mongolian Journal of Biological Sciences. 3: 3-15.
Clark, E.L., Ocock, J.F., King, S.R.B. & Baillie, J.E.M. 2006. ‘Proceedings of the Mongolian Biodiversity Databank Workshop:Assessing the Conservation Status of Mongolian Mammals and Fishes. II – Mongolian Mammals: Assessment Results and Threats’. Mongolian Journal of Biological Sciences. 3: 17-27.
Ocock, J.F., Clark, E.L., King, S.R.B. & Baillie, J.E.M. 2006. ‘Proceedings of the Mongolian Biodiversity Databank Workshop:Assessing the Conservation Status of Mongolian Mammals and Fishes. III – Mongolian Fishes: Assessment Results and Threats’. Mongolian Journal of Biological Sciences. 3: 29-36.