IWEC2019 Abstract Book FINAL.Pdf

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Book of Abstracts

International Wild Equid Conference 2019

International conference organised by

IUCN/SSC Equid Specialist Group &

Derbianus Conservation &

Prague Zoo &

United States Geological Survey &

Faculty of Tropical AgriSciences (Czech University of Life Sciences Prague)

September 1 - 5, 2019, Prague, Czech Republic

Published by: Czech University of Life Sciences Prague, Kamýcká 129, 165 21, Prague 6, Czech Republic.

Reproduction of this publication for educational, conservation and other non- commercial purposes is authorised without prior written permission from the copyright holder provided the source is fully acknowledged. Reproduction of this publication for sale or other commercial purposes is prohibited without prior written permission of the copyright holder.

The authors of abstracts are fully responsible for the content of their contributions. Abstracts have not undergone proofreading and are reproduced such as supplied by the authors.

Recommended citation: Pluháček J, Ceacero F, Garguláková A, Kubičková S, Komárková M, King SRB, Moehlman P, Schoenecker K, Šimek J, Brandlová K, Gloneková M. (Eds.) 2019: 2nd International Wild Equid Conference. Book of Abstracts. Czech University of Life Sciences Prague. 215 pp.

Photo credits: Pluháček J (2, 6, 8, 114, 202, 215), Kubičková S (cover, 116), Ceacero (12, 30), Bobek M (1), Staněk JV (9). Cover design: Ceacero F. Logo design: Kubičková S.

Number of copies: 160 ISBN 978-80-213-2954-6 www.iwec2019.com

Organising committee: Jan Pluháček – Ostrava Zoo, Czech Republic Sarah R. B. King – Colorado State University, USA Kate Schoenecker – U.S. Geological Survey, USA Patricia Moehlman – IUCN/SSC Equid Specialist Group, Tanzania

Technical organising team: Markéta Gloneková – Derbianus Conservation Martina Komárková – Faculty of Tropical AgriSciences Francisco Ceacero Herrador – Faculty of Tropical AgriSciences Karolína Brandlová – Faculty of Tropical AgriSciences, Derbianus Conservation Jaroslav Šimek – Prague Zoo

Scientific steering committee: Jan Pluháček – Ostrava Zoo, Czech Republic Patricia Moehlman – IUCN/SSC Equid Specialist Group, Tanzania Sarah King – Colorado State University, USA Kate Schoenecker – U.S. Geological Survey, USA Morris Gosling – University of Newcastle, Great Britain Wayne Linklater – California State University, USA Albano Beja-Pereira – University of Porto, Portugal

Volunteers: Lucie Ackermann Blažková Klára Ničová Iva Bernáthová Petra Michalčáková Anna Bernátková Jana Marešova Alena Divišová Aneta Pilíková Tamara Fedorová Irena Schneiderová Katka Gašparová Helena Slepičková Andrea Garguláková Marketa Swiacka Zuzana Holubová Nikola Sýkorová Sabina Jonášová Kateřina Šandlová Klára Kopicová Lucie Šašková Simona Kubičková Kateřina Štochlová

Organisers

Sponsors

Contents

PREFACE 7 USEFUL INFORMATION 10 PROGRAMME 12 PLENARY SPEAKERS 18 PLENARY LECTURES 21 ORAL CONTRIBUTIONS 26 MANELESS ZEBRA SESSION 111 POSTER CONTRIBUTIONS 115 LIST OF PARTICIPANTS 187 AUTHOR INDEX 193

Preface

Dear participants of the conference,

Allow me on behalf of the organizing committee to welcome you to the 2nd Wild Equid International Conference (IWEC) 2019 in Prague. This conference follows a very successful first IWEC that was held in Vienna seven years ago. The IWEC in Vienna was hosted at the University of Veterinary Medicine by Dr. Petra Kaczensky and Dr. Jason Ransom. This second IWEC was initiated by the US Geological Survey feral equid researchers and the International Union for Conservation of Nature Species Survival Commission (IUCN/SSC) Equid Specialist Group (ESG). As a place for this second conference, Prague was suggested as a location accessible to all. This choice complements the long tradition of research and conservation of wild equids in Czechia. It was exactly 60 years ago, in September 1959, that Prague hosted the First International Symposium for Conservation of Przewalski’s Horse. As a consequence of this symposium, Prague Zoo established the International Studbook for Przewalski’s Horse. This initiated a very successful conservation story, resulting in the reintroduction of Przewalski’s horses to Mongolia and China at the end of 20th century and beginning of this century. We would like to follow this success and point out the critical situation of another beautiful wild equid – the maneless zebra (Equus quagga borensis). Similar to Przewalski’s horse in the 1950s, the maneless zebra wild population is currently in a very precarious situation, yet fortunately there is a small but viable captive population, with most individuals kept in Czech zoos. Therefore, the maneless zebra was chosen as the logo of this conference.

This IWEC aims to combine current research on the state of wild equids with conservation efforts. Therefore, the conference starts with a closed meeting of the IUCN/SSC ESG. Subsequent talks and posters were split in to eight symposia: Behavioral ecology; Diseases; Conservation; Population dynamics, viability, and estimates; Genetics and morphology; Modeling for wild and feral equids; Equid reproductive biology and fertility control; and, Feral equid issues and rewilding. In total, 3 plenary talks, 41 oral talks, and 40 posters will be presented at the conference. We are very glad that the conference will be attended by 120 participants representing 30 countries from Africa, Asia, Australia, Europe, and North America. Unfortunately, we have some very sad news: one of our speakers, the eminent Mongolian scientist and coordinator of Asiatic wild asses for the IUCN/SSC ESG, Dr. Badamjav Lkhagvasuren, passed away a few weeks before the start of the conference. His abstract remains included in this book and the talk will be presented by his co-author. The conference is co-organized by the IUCN/SSC ESG, Czech University of Life Science in Prague, Prague Zoo, and the US Geological Survey. My thanks are due to the plenary speakers as well as all conference participants who come to Prague to share their knowledge and experiences. I would like to thank all members of the organizing team, technical organizing team, and volunteers. Without your help the conference could not take place. Welcome to Praha

Jan Pluháček

First International Symposium for Conservation of Przewalski’s Horse in Prague in September 1959.

Useful information

The conference is held in the T-building of the Faculty of Tropical Agrisciences in the campus of the Czech University of Life Sciences Prague. The whole scientific programme, registration desk and exhibitors stands are located there.

The registration desk will be open from Sunday, September 1st, 11:00, during the whole conference according to the schedule in the conference programme. Participants can get any kind of information, register for the excursions or pay additional fees there.

Lunches and coffee breaks take place in the Club C restaurant, neighbouring the T-building at times stated in conference programme and are free for all registered participants. The Gala dinner is served in the Galaxie hotel, about 800 meters from the Campus, see the map.

Do not hesitate to ask the conference organisers and volunteers. You can easily recognise them by their yellow T-shirts with the conference logo.

WiFI

Username: conference Password: iwec2019

Programme

SUNDAY September 1st

11:00 – 18:00 Registration opened at the hall of the T building 12:00 – 18:00 Free entry to the Prague Zoo (for registered participants only) 12:30 – 13:30 Lunch (for IUCN/SSC ESG members only) 13:30 – 17:00 IUCN/SSC ESG – closed meeting 18:00 – 22:00 Icebreaker in Prague Zoo (for registered participants only)

MONDAY September 2nd

8:30 – 9:30 Opening ceremony Jan Pluháček – IUCN/SSC ESG Petr Sklenička – Czech University of Life Sciences Sarah R. B. King, Patricia Moehlman, Kate Schoenecker – IUCN/SSC ESG Miroslav Bobek – Prague Zoo Jiří Volf – Prague symposium for Przewalski’s horse in 1959 Recognition of equid specialists we lost (H. Klingel, C. P. Groves, B. Lkhagvasuren, F.J. Singer)

9:40 – 10:40 Plenary talk: Morris Gosling: Population ecology and conservation of Hartmann’s mountain zebra

10:40 – 11:00 Coffe break

SESSION I. BEHAVIORAL ECOLOGY I. Moderator: Patricia Moehlman 11:00 – 11:20 Martina Burnik Šturm: Dietary behaviour of three sympatric equid species in the Dzungarian Gobi – past and present 11:20 – 11:40 Battogtokh Nasanbat: Mongolian wild ass (Equus hemionus hemionus) water use patterns and interactions with other large mammals in the Gobi Desert 11:40 – 12:00 Amos Bouskila: Space-use patterns of the Asiatic wild ass (Equus hemionus) 12:00 – 12:20 Enkhbayar Jagdag: Movement and space use of Przewalski’s horses in Hustai National Park, Mongolia 12:20 – 12:40 Laura Lagos Abarzuza: Insights into spatial ecology and vegetation use of Galician wild ponies: implications for the conservation of priority wet heathlands and peatlands 12:40 – 13:00 Susane Shultz: Social networks in Carneddau ponies: exploring stability, flexibility and the impact of perturbations

13:00 – 14:00 Lunch

SESSION II. BEHAVIORAL ECOLOGY II. Moderator: Albano Beja-Pereira 14:00 – 14:20 Patricia Moehlman: Comparative behavior and mating systems of African wild ass (Equus africanus) and feral donkeys (Equus asinus) in arid and mesic habitats 14:20 – 14:40 Francesca Parrini: Do zebra shape resource and community level interactions within a grazing guild? DISEASES 14:40 – 15:00 Jocelyn Poissant: Development and application of a non-invasive DNA metabarcoding assay to characterize complex parasitic strongyle infections in equids 15:00 – 15:20 Amy McLean: Survey of serum amyloid A and pathogen frequency in recently captured feral donkeys in Death Valley National Park, California USA

15:20 – 15:40 Luna Gutiérrez Cepeda: The Darwinian adaptation of the equine intestine vascular network to Strongylus vulgaris 15:40 – 16:00 Erin Goodrich: A serosurvey for selected pathogens in feral burros from Death Valley National Park, California, USA

16:00 – 16:20 Coffee break 16:00 – 17:00 POSTER SESSION

SESSION III. BEHAVIORAL ECOLOGY III. Moderator: Wayne Linklater 17:00 – 17:20 Sarah R. B. King: Social system shapes differences in social behaviors of equids 17:20 – 17:40 Ariel Altman: Internal and external effects on grouping of a fission-fusion society of female Asiatic wild asses (Equus hemionus) in the Negev desert, Israel 17:40 – 18:00 Lucy Rees: Anti-predator defence behaviour in feral horses 18:00 – 18:30 MANELESS ZEBRA SESSION

TUESDAY September 3th

9:00 – 10:00 Plenary talk: Wayne Linklater: A future with equids – our culture, values and reconciliation science

SESSION IV. BEHAVIORAL ECOLOGY IV. Moderator: Sarah R. B. King 10:00 – 10:20 Jan Pluháček: Overmarking by adult females in four equid species serves to strengthen the social bonds and to support group cohesion 10:20 – 10:40 Hannah Davie: Relationships between reproductive state and social connections in reintroduced takhi (Przewalski’s horse: Equus ferus przewalskii) family groups in Mongolia 10:40 – 11:00 Dan Rubenstein: The causes and consequences of zebra hybrids

11:00 – 11:20 Coffee break

SESSION V. BEHAVIOURAL ECOLOGY V. Moderator: Jan Pluháček 11:20 – 11:40 Kate Schoeneker: Effect of availability and temporal variability of resources on the distribution and movement of feral burro (Equus asinus) populations

11:40 – 12:00 Redae Teclai Tesfai: Key resource competition threatens viability of the critically endangered African wild ass (Equus africanus) population on Messir Plateau (Eritrea) CONSERVATION 12:00 – 12:20 Yash Veer Bhatnagar: Kiang conservation in India: A landscape approach 12:20 – 12:40 John D. C. Linnell: Reintroduction, persistence and local extinction of central Asian kulan in Turkmenistan and Kazakhstan 12:40 – 13:00 Jaroslav Šimek: Return of the wild horses – reintroduction of Przewalski’s Horses from Europe to Western Mongolia

13:00 – 14:00 Lunch

14:00 – 18:00 Excursion to Prague zoo 19:00 – 00:00 Gala dinner (Hotel Galaxie)

WEDNESDAY September 4th

9:00 – 10:00 Plenary talk: Albano Beja-Pereira: Population genetics of the wild asses

SESSION VI. POPULATION DYNAMICS, POPULATION VIABILITY, AND POPULATION ESTIMATION I. Moderator: Kate Schoenecker 10:00 – 10:20 Sheila Funnell: Warriors using SMART to monitor endangered Grevy's zebra in northern Kenya 10:20 – 10:40 Timothy Kaaria: Population performance of the endangered Grevy's zebra (Equus grevyi) on the Lewa - Borana Landscape, Meru, Kenya 10:40 – 11:00 Peter Gogan: Causes of mortality, survival rates and reproductive patterns in plains zebra in the Serengeti Region, Tanzania

11:00 – 11:20 Coffee break

SESSION VII. POPULATION DYNAMICS, POPULATION VIABILITY, AND POPULATION ESTIMATION II. Moderator: Lee Boyd 11:20 – 11:40 Philip D. Mcloughlin: Social environmental effects on dispersal success and sex differences in survival drives adult sex-ratio bias in feral horses 11:40 – 12:00 Viola Kerekes: Demographical changes in the Hungarian Przewalski horse population during the first 20 years of the project 12:00 – 12:20 Jake Britnell: Can gut health provide a mechanistic link between diet and performance within the Cape mountain zebra (Equus zebra zebra) meta-population?

12:20 – 12:40 John Derek Scasta: Equids and humans: a historical examination of global psycho-ecology influencing emotions 12:40 – 13:00 Bayarbaatar Buuiveibatar: Population status and conservation activities for Asiatic Wild Ass (Equus hemionus) in Mongolia

13:00 – 14:00 Lunch

SESSION VIII. MODELING FOR WILD AND FERAL EQUIDS. Moderator: Morris Gosling 14:00 – 14:20 Saeideh Esmaeili: Does the forage maturation hypothesis predict seasonal movements of equids? 14:20 – 14:40 L. Stefan Ekernas: Modeling population growth and management costs for feral horses in the U.S. 14:40 – 15:00 Jeffrey L. Beck: Safety and application of telemetry collars on wild and free- roaming equids: A global systematic review 15:00 – 15:20 Jacob D. Hennig: Movement syndromes of feral horses inhabiting arid steppe in Western North America

EQUID REPRODUCTIVE BIOLOGY AND FERTILITY CONTROL 15:20 – 15:40 Budhan Pukazhenthi: Application of reproductive technologies for conservation of endangered equids 15:40 – 16:00 Cassandra Nuñez: PZP-induced mare infidelity increases male aggression and stress in the feral horse

16:00 – 16:20 Coffee break

Session IX. EQUID REPRODUCTIVE BIOLOGY AND FERTILITY CONTROL. Moderator: Dan Rubenstein 16:20 – 16:40 Kayla A Grams: Advances in PZP Immunocontraception in two Western wild horse herds GENETICS & MORPHOLOGY 16:40 – 17:00 Sara Oyler-McCance: Facts from feces: the utility of fecal DNA to assist in the conservation and management of wild equids

FERAL EQUID ISSUES AND REWILDING 17:00 – 17:20 Terry A. Messmer: Are free-roaming equids good for western U.S. public lands? 17:20 – 17:40 David Berman: Management of feral horse environmental impact in Australia 17:40 – 18:00 Enrique Alonso Garcia: The rewilding Europe with wild/feral equids and its infrastructural value for equids cognition and social behavioral ecology: the Piornal (Pottokas) and Doñana (Retuertas´ horses) initiatives in Spain

18:15 – 18:45 Closing ceremony

THURSDAY September 5th

8:00 – 18:00 Post conference trip to Dolní Dobřejov (facilities of Prague Zoo for Przewlaski´s horses) + Milovice (feral horse’s rewildling project) 8:00 – 18:00 Post conference trip to Dvůr Králové Zoo (African equids)

Plenary speakers

Morris Gosling

Morris Gosling is an Emeritus Professor in the Institute for Sustainability at the University of Newcastle, UK. He previously worked for the UK Scientific Civil Service, for the Zoological Society of London, and as Professor of Animal Behaviour at the University of Newcastle. He has been studying the population ecology and behaviour of Hartmann's mountain zebra in Namibia, and coordinating the Mountain Zebra Project, since 2004. This work uses an individual-based approach and aims to obtain basic knowledge about individual strategies and population processes and to provide the information needed for evidence-based conservation practice. When in Namibia he is based at the Namibia Nature Foundation in Windhoek.

Wayne Linklater

Wayne Linklater is Associate Professor of Conservation Science at Victoria University of Wellington. He is also a Research Associate with the Centre for African Conservation Ecology at Nelson Mandela University and currently Visiting Scholar in the Department of Environmental Science, Policy & Management at the University of California, Berkeley. His research involves a range of topics from large mammal behaviour, ecology and management (mostly on horses, rhino and elephant), to human-dimensions ecology (like human-wildlife conflict, wildlife crime, and studies of people's pro-environmental behaviour), to critiques of conservation science, culture and policy. He is writing a book about the Kaimanawa Wild Horses and research experience with them for a general audience. Its working title is "Horses of the Last Land".

Albano Beja-Pereira

Albano Beja-Pereira is the Population Genetics and Group Leader at the Center for Research on Biodiversity and Genetic Resources, University of Porto, Portugal (CIBIO-UP). His research topics are on population and conservation genomics of the wild ass-like species (Asian and African). He has been generating genetic data from noninvasive samples from all natural populations of African and Asiatic wild asses to assess population diversity, structuring and connectivity. The resulting data will provide guidance to the management of conservation programs on these species, and provide tools for use on the identification and provenance (at the population level) of biological samples or speciments from museum collections, or apprehended from illegal hunting activities. Albano has also been involved in population and conservation genetics projects on Asiatic wild Ovis and Capra populations.

Plenary lectures

MONDAY 9:40-10:40 Population ecology and conservation of Hartmann’s mountain zebra Morris GOSLING1 1 School of Natural and Environmental Sciences, Newcastle University, UK and Namibia Nature Foundation, Windhoek, Namibia.

Hartmann’s mountain zebra (HMZ) is a sub-species of mountain zebra that occurs mainly in Namibia. The core of it’s range is the mountainous escarpment that runs the length of the country from north to south. There is some genetic structuring of the national population and HMZ differ from Cape mountain zebra, found only in the Republic of South Africa, in a few private alleles. The population was reduced to low levels in the 1980s but has recovered steadily and is now locally abundant. Zebra populations can be impacted by anthropogenic factors, particularly where they compete with livestock, and also by a range of natural factors. A key recommendation of a recent IUCN assessment was the need to better understand the factors limiting HMZ populations throughout Namibia to better inform their conservation management. In this talk I will outline some early results from a study of limiting factors. These data are from long-term research and monitoring at 5 main study areas, selected mainly for variation in rainfall and the occurrence of key predators. The approach is individual-based and uses investigation of key population processes within study sites (such as age specific mortality rates), and a comparative approach between them. The results emphasize the importance of variation in rainfall in a country that is prone to periodic drought and of freedom of movement to patchy, ephemeral grazing. The significance of these findings for the future conservation of the sub-species, and the potential impact of climate change will be considered.

TUESDAY 9:00-10:00 A future with equids – our culture, values and reconciliation science Wayne LINKLATER1 1 California State University – Sacramento, 6000 J. Street, 95819, California, USA

Conservation is not a science. But conservation and science are similar. They are both fundamentally and strongly, although not always transparently, constructed and influenced by our cultures, beliefs and values. For some, this statement will be an anathema to their views about the importance of empirical rationalism and scientific objectivity. Others will view the statement as self-evident and uncontroversial but will think that it needs to be said and thought about more often, especially by scientists. In this talk, I will explore the cultures, values and relationship of conservation and science, with examples from the science and conservation of wild equids. My first premise is that we are on the threshold of a potentially significant schism in conservation. The growing rift is the consequence of resistance by some in the conservation and science communities to calls by others for us to be bio-culturally inclusive and embrace pluralism – a patchwork of different solutions across peoples, space and time. My second premise is that equids and those who study them are at the forefront of this global paradigmatic change in biodiversity conservation and science. Our unique history and relationship with equids and, although numbering just a few species, that they occupy extremes on the bio-cultural spectrum makes this so. Equids are variously regarded as pests or invasive, common and ecologically important or iconic wildlife, as well as critically endangered species. To develop and support these two propositions, I will focus on three case studies: (1) The unique biodiversity of New Zealand has been extirpated or remains highly threatened by introduced mammals. The New Zealand government has announced its intention to eradicate five

introduced mammal predators by 2050. In contrast it has adopted a co-management and co-existence strategy for introduced wild horses. (2) The mega-fauna extinction in Australia was extraordinary and large proportions of its extant vertebrate species are IUCN Red Listed as Threatened or worse. But, if Australia’s immigrant and emigrant populations of those species were included in the Red List, many of them including equids, would be reclassified as Least Concern. (3) Wild horses in North America and Australasia are regarded as invasive species and the subject of intense environmental conflict, but much less controversial elsewhere. Recently, some invasion biologists made accusations of invasive species science denialism at other scientists and scholars who contest the judgement that species, like horses, are ubiquitously harmful. The case studies illustrate how the cultures of conservation and science, especially invasion biology, causes us to under-use ecological science in policy-making, mis-measure global biodiversity, and fuel, rather than resolve, environmental conflicts. I think we need a gestalt-switch. I will make a case for the expansion of Michael Rosenzweig’s concept of reconciliation ecology to include human-human relationships. I’m calling it reconciliation science. Reconciliation science draws more deeply and broadly from the environmental sciences, especially ecology and the social sciences. It will do that without cultural hegemony and prejudice by being actively transparent about, and accepting of, the many human cultures and values that conservation science must serve.

WEDNESDAY 9:00-10:00 Population genetics of the wild asses Albano BEJA-PEREIRA1,2, Rita MONTEIRO1,3 1 Research Centre on Biodiversity and Genetic Resources CIBIO-InBIO, University of Porto, Campus de Vairão 7, 4485-661 Vairão, Portugal 2 Department of Geosciences, Environment and Spatial Planning, Faculty of Sciences, University of Porto, R. Campo Alegre 687, 4169-007 Porto, Portugal 3 Department of Biological and Geographical Sciences, School of Applied Sciences, University of Huddersfield, Queensgate, Huddersfield, United Kingdom

The wild asses are among the least studied species of large mammals. These species occupy a large territory that encompasses a broad part of Asia and the Northeast of Africa, although most of them are scattered and isolated. In this talk, the preliminary results of a large noninvasive population genetics project that aims to assess the population structure, diversity and connectivity will be presented. Particular attention will be given to the E. kiang as the sole equid species that can be found at high-altitude ranges (>2500m) and which its conservation status is the less worrisome among the wild ass species. The analysis of data from uniparental and autosomal genetic markers indicate that despite their different habitats and pressures connectivity between populations geographically close still occur. Moreover, connectivity between different species can also be noticed and the observed diversity levels across populations are relatively high despite the fragmentation and depletion of their habitat that has been increasing in the last century.

Oral contributions

Behavioral Ecology

MONDAY 11:00-11:20

Dietary behaviour of three sympatric equid species in the Dzungarian Gobi – past and present

Martina BURNIK ŠTURM1, Natalia N. SPASSKAYA2, Mikhail V. SABLIN3, Christian C. VOIGT4, Oyunsaikhan GANBAATAR5,6, Steve SMITH7, Petra KACZENSKY1,8

1 Research Institute of Wildlife Ecology, Vetmeduni Vienna, Austria 2 Zoological Museum of Lomonosov Moscow State University, Moscow, Russia 3 Zoological Institute, RAS, St. Petersburg, Russia 4 Leibniz Institute for Zoo and Wildlife Research, Berlin, Germany 5 Great Gobi B SPA Administration, Takhin Tal, Mongolia 6 ITG – International Takhi Group, Richenthal, Switzerland 7 Konrad-Lorenz-Institute of Ethology, Vetmeduni Vienna, Austria 8 Norwegian Institute for Nature Research – NINA, Trondheim, Norway

We studied foraging behavior of three sympatric equid species in the Dzungarian Gobi in southwestern Mongolia: the reintroduced Przewalski’s horse (Equus ferus przewalskii), the khulan (Equus hemionus), and their potential competitor the domestic horse (Equus caballus). Using segmentally cut tail hair, a biological archive that contains chemical information about what the animals have been eating in the form of stable isotopes obtained from food, we draw inferences about their multi-year diet seasonality and isotopic dietary niches. Specifically, we asked: Do they feed as grazers, browsers or mixed-feeders? Do they compete for the same resources? In addition, stable isotope analysis of historic museum samples from the

same area enabled the comparison of pre-extinction and extant reintroduced Przewalski’s horses. A stable isotope mixing model revealed that both extant horse species are predominantly grazers that include browse (>20 %) in their diet only under extreme environmental conditions (e.g. dzud winter). Khulan, on the other hand, are highly seasonal, switching between grass dominated diet in summer and mixed grass-browse diet during winter when food availability is lowest and competition with other wild and domestic grazers highest. Our results suggest that the differences in foraging adaptations facilitate the coexistence of horses and khulan in the same habitat, but the high degree of isotopic niche overlap of both horse species indicates a high potential for pasture competition in winter. The analysis of historic tail hairs revealed a dietary change from pre-extinction to reintroduced Przewalski’s horses, namely from a highly seasonal diet (i.e. grazing in summer and mixed diet in winter) in historic to year-round grazing in extant Przewalski’s horses. We explain this dietary shift in Przewalski’s horses by a release from human predation. Until recently, both the Przewalski’s horses and khulan have been heavily hunted as game and persecuted as pasture competitors. Nowadays both species are fully protected, but whereas the newly returned Przewalski’s horse has become a much appreciated iconic national flagship species, attitudes towards khulan remain ambivalent. This changed positive attitude allows the reintroduced Przewalski’s horses to feed on the scarce, grass dominated pastures alongside local people and their numerous livestock whereas their historic conspecifics (like historic and extant khulan) were forced into less productive habitats dominated by browse during the resource poor winter season.

MONDAY 11:20-11:40

Mongolian wild ass (Equus hemionus hemionus) water use patterns and interactions with other large mammals in the Gobi Desert

Nasanbat BATTOGTOKH1,2, Francisco CEACERO3, Ravchig SAMIYA1

1 National University of Mongolia, School of Arts and Sciences, Ulaanbaatar, Mongolia 2 Mongolian Academy of Sciences, Institute of General and Experimental Biology, Ulaanbaatar, Mongolia 3 Czech University of Life Sciences Prague, Faculty of Tropical AgriSciences, Prague, Czech Republic

The most abundant population of the wild ass or khulan exist in Mongolian Gobi desert and represents more than 80% of the total population in the world. In recent years, researchers have focused on population dynamics, diet and movement patterns. Still, little is known about the rhythms of use of water points by khulan in Gobi desert, as well as interspecific interactions at water points, which is the most scarce essential resource for fauna in such harsh habitat. This study aims to determine how khulans use water sources, detect seasonal, daily and spatial overlaps with other species, and understand the influence of environmental factors (climate and moon phase) in the patterns observed. The ecological information collected about resource partitioning has important value for the conservation of the species inhabiting this extreme and fragile ecosystem. The study was conducted at six natural springs without human influence in the Eastern part of Great Gobi A Strictly Protected Area. Five to eight cameras were deployed on the each water source, depending on their orography (i.e., the number of routes available for the large mammals living in the area to enter to each water source). In total, 38

cameras were working from May 13th, 2015 until May 15th, 2017. We detected 13 species of large mammals. Khulan was the most observed species, with 1416 independent events and mean daily trapping rate of 28.69. Khulan showed a clumped seasonal activity in the area (Rayleigh test; p<0.001) with a peak around the second half of July not overlapping with the activity observed in other large mammals. The average monthly activity of khulan was positively affected by average monthly weather conditions like air temperature, soil temperature and precipitation, but not by the relative humidity. The activity of khulan was also not influenced by the moon phase, showing the lowest lunar phase difference among all the species observed in the area. About the circadian patterns, khulan also showed a clumped distribution with mainly nocturnal activity (Rayleigh test; p<0.001) and overlap was detected with Eurasian lynx and snow leopard (Watson´s U test; p=0.15 and p=0.35). The daily activity showed scarce seasonal variation: the observed frequency of hourly activity showed significant bivariate correlation among seasons (grouped Dec-Feb, Mar-May, Jun-Aug, Sep-Nov; all p<0.001). However, analyzed by month we observed significantly different hourly activity patterns between the winter months (January, February and March) and July, being more nocturnal during winter but crepuscular during July. Finally, there was also a strong spatial segregation between khulan and other large mammals in the area: The frequency of occurrence normalized by effort of khulan at each studied location was only correlated to that of Tolai hare, wild camel and grey wolf (Spearman´s correlation; p<0.001, p=0.031 and p=0.020 respectively). Combining the seasonal, circadian and spatial results obtained, we can conclude that khulan’s natural use of waterholes successfully avoid to overlap with other large mammals, indicating complex patterns of ecological segregation. Further analyses should focus on predation and competition as drivers of the observed segregation patterns.

MONDAY 11:40-12:00

Space-use patterns of the Asiatic wild ass (Equus hemionus)

Amos BOUSKILA1,2, Nina GIOTTO2, Jean-François GERARD3, Alon ZIV1,2, Shirli BAR-DAVID2

1 Department of Life Sciences, Ben-Gurion University of the Negev, Israel 2 Mitrani Department of Desert Ecology, Blaustein Institutes for Desert Research, Ben-Gurion University of the Negev, Israel 3 Comportement et Ecologie de la Faune Sauvage, Institut National de la Recherche Agronomique, Castanet-Tolosan Cedex, France

The movement of animals and their habitat use result from interactions between the spatial distribution of resources and the requirements of the animals in terms of nutrition, water, safety and social interactions. By understanding these requirement, we can improve our ability to prioritize sites to set aside as reserves or even influence the distribution of resources, e.g., through the distribution of water sources. We studied the Asiatic wild ass (Equus hemionus) in the Negev Desert, Israel, using GPS monitoring of five individuals complemented during summer with 274 hours of direct observation. We analysed movement, reflected in displacement and in recursions to specific sites, and aspects of habitat preferences, comparing 50 consecutive days in winter with 50 consecutive days in summer. We characterized habitats by variables that we obtained from geographic information system (GIS) layers and calculated selection coefficients from GPS locations of the marked individuals. We found that summer home ranges tended to be slightly smaller than winter home ranges. Nevertheless, there were marked differences in other variables. While the distance travelled per hour during the day was similar to that in winter, the distances at dawn, at dusk and at night were higher in summer

compared to winter. Combined with the direct observation, the temporal pattern that emerged for the summer was of peak distances moved at dawn with low activity during the day. During the day wild asses often remained at high elevation sites with low NDVI, but these sites are more likely to expose them to higher winds. Movement increased again at dusk and towards night time, when the wild asses occupied mainly wadis with high values of NDVI. Two of the marked males were territorial in summer, while in winter, they were observed switching to multi- male association, occasionally conducting solitary trips to their summer territory. In summer, eight sites fitted the definition of recursion sites. Four of the sites were along a wadi, exhibited high NDVI scores and were used at night. Three of the sites were used during daytime, located on elevated dusty grounds with low NDVI scores. The last site was a water point that was used by all individuals at all times of the day, 1.3 times a day, on average. In winter, 15 main recursion sites were identified. Four of these sites had low to moderate NDVI scores and rather high elevation and they were mainly used during midday hours. This fits our observations of midday winter groups standing on high-elevation dusty plateaus and ridges, occasionally rolling, and intensively interacting socially. In contrast to summer, the main water source was visited by the wild asses in winter only once in 12 days, on average, and animals must be drinking in small ephemeral ponds formed in numerous locations in the wadis. Besides emphasizing the importance of the water source in summer, the space-use pattern that emerged demonstrates the role of high elevation sites, mainly in summer, but used also in winter, presumably for insect control and social interactions.

MONDAY 12:00-12:20

Movement and space use of Przewalski’s horses in Hustai National Park, Mongolia

Enkhbayar JAGDAG1*, Melissa SONGER2, John MCEVOY2, Kate JENKS3, Peter LEIMGRUBER2, Usukhjargal DORJ1, Jared STABACH2, Nandintsetseg DEJID2,4

1 Hustai National Park, Ulaanbaatar, Mongolia 2 Smithsonian Conservation Biology Institute, VA, USA 3 Minnesota Zoo, MN, USA 4 Senckenberg Biodiversity and Climate Research Centre, Frankfurt, Germany

The Przewalski’s horse (Equus ferus przewalski), also called takhi, was previously classified as “Extinct in the wild (EW)” by the International Union for the Conservation of Nature (IUCN) Red List. Through successful reintroduction programs for Przewalski’s horses at various sites, including Mongolia (since 1992) and China (since 2001), the status of Przewalski’s horse was revised to “Critically Endangered (CR)” in 2008 and then is reclassified to Endangered (EN) in 2011. Most free-ranging mature individuals in the wild are found in Hustai National Park (HNP), Mongolia. This suggests that HNP is a critical conservation location for Przewalski’s horse and effective conservation management is a key requirement for the long-term survival of this population. Understanding the movement and space use of Przewalski’s horses in HNP is needed in order to identify priority areas for conservation and improve habitat connectivity, and inform effective conservation management. We deployed global positioning system (GPS) collars on 18 female Przewalski’s horse across distinct family groups in 2016 and 2017. Monitoring periods ranged from 251 to 1,046 days (mean = 657 days), while the number of recorded GPS locations ranged from 5,982 to 24,988 (mean = 15,613).

For each individual, we quantified home range sizes, overlap of the home ranges, and daily movements using the Autocorrelated Kernel Density Estimator (AKDE). We found that on average, an individual Przewalski’s horse in HNP moved 6.2 km in a day and had a mean home range of 72.7 km2 (n = 18). However, there was great variation among individuals (SD = 46.2 km2). The largest home range (194.0 km2) of an individual Przewalski’s horse was more than six times larger than the smallest range (28.6 km2). On average, an individual Przewalski’s horse took 3.0 ± 2.6 days to cross its home range. In addition, we found that the mean percentage overlap of home ranges was 71.0 ± 20.5 %, indicating a high degree of spatial overlap between individual home ranges. All individual home ranges overlapped at the core range of HNP. When forage and water availability was limited during winter, one individual extended its range beyond the core range, while all the other individuals concentrated their space use more tightly within the core range. The high overlap of home ranges indicates that the Przewalski’s horses are primarily using a single core area in HNP. The selected area of concentrated use within home ranges are often denoted as suitable habitat, implying that the selected area is of greater significance to Przewalski’s horses in HNP. Future research is underway to better understand why Przewalski’s horses are limiting their movements to this core area and to assess whether there are management interventions that may encourage them to use broader ranges.

MONDAY 12:20-12:40

Insights into spatial ecology and vegetation use of Galician wild ponies: implications for the conservation of wet heathlands and bogs

Laura LAGOS1, Carlos MUÑOZ-BARCIA1, Jaime FAGÚNDEZ1

1 Biocost Group, Faculty of Biology and Advanced Scientific Research Center (CICA). University of A Coruña, Spain

Traditional populations of native wild ponies have a positive effect on habitats of conservation importance in the Atlantic regions of western Europe as wet heathlands. These are priority habitats of the European Habitats Directive and constitute valuable cultural landscapes. Heathlands are semi-natural habitats that have been shaped by different forms of human use such as mowing, burning and grazing by herbivores. Specifically, the presence of Galician wild ponies in the wet heaths of the north of Galicia (NW Spain) determine vegetation structure and species composition, thus playing a major role in their conservation. Changes in land uses, mainly by the transformation to pastures for livestock and afforestation, as well as abandonment are threatening these habitats. In 2017, we started the project Life in Common Land (LIFE16 NAT/ES/000707) aimed at the conservation of the mosaic of heathlands, blanket bogs and raised bogs on eleven Common Lands in Serra do Xistral, a NATURA 2000 Special Conservation Area located in the North of Galicia, NW Spain. There is a population of 1500-2000 ponies in the project area that extends on about 100 km2. As part of the project, explored the space use and social structure for different bands, differences in management by owners and Commons, as well as diet during a 12-month period. Between June and October 2018 we fitted 28 mares integrated in 15 bands inhabiting 5 different Commons with GPS collars. We conducted surveys every 1-

2 months to find the bands with collared mares and record group size, body condition of mares, behavior of collared mares, survival of foals and other variables related to the dynamic of the group. First 6-month analysis indicates that heathlands and bog constituted a large percentage of the home range of the studied bands (33-87%). Ponies organize themselves into bands of between 7-32 adults. Home range size, estimated through kernel density estimation, varied between107- 317 ha. A low level of overlapping of home ranges (24-30%) was observed in the Commons with a traditional management, where ponies are allowed to free roam and drifted once a year. Ponies show a high consumption of gorse (Ulex spp.), especially in winter, and they also consume Molinia caerulea mainly during the summer. Results on the spatial ecology, social structure and diet of wild pony bands highlight the importance of these herbivores in the conservation of heathlands and bogs.

MONDAY 12:40-13:00

Social networks in Carneddau ponies: exploring stability, flexibility and the impact of perturbations

Susanne SHULTZ1, Jess LEA1,2, Christina STANLEY3, Danielle HINCHCLIFFE1,2, Rachael ANTWIS4

1 School of Earth and Environmental Sciences, University of Manchester 2 North of England Zoological Society, Chester Zoo 3 Department of Biological Sciences, University of Chester 4 School of Earth and Environment, University of Salford

Long-term relationships that underlie many stable mammalian groups often occur between philopatric kin. Although stable groups of nonrelatives appear to be less common, there is increasing evidence that social bonds between non-kin may confer sufficient intrinsic fitness benefits for these groups to persist. Identifying proximate measures of the costs and benefits of group membership and network structure is a missing piece of the puzzle. Moreover, in a management context, we do not understand the consequences of removing key individuals from such populations, but predict that removals have the potential to result in large destabilising impacts to social network structure. Here we use network approaches to evaluate patterns of social stability and flexibility in a free-ranging population of Carneddau mountain ponies. First, we quantified female social stability by applying a three-level framework to a 3-year data set of associations, which tested for stability at the individual, dyadic and subpopulation levels. Despite the relative weakness of these female bonds, we found significant social stability across all levels, as shown by stable association preferences, social networks and individual network positions. Second, we investigated how seasonality impacts on social bond

strength and grouping patterns. We found seasonal fluctuations in female gregariousness, with a peak during the mating season. We therefore propose that significant social stability in female horses is coupled with a degree of flexibility that allows for effects of ecological fluctuations. Third, we evaluate how network structure impacts on gut health and microbiome communities and demonstrate that microbiome communities are more similar between individuals with stronger network ties. Finally, we evaluate the impact of the removal of both young and mature stallions on the social structure of this population. The removal of herd stallions led to widespread changes to the social network structure, signified by a decrease in clustering, tie strength, and an increase in connectivity across the network as more individuals were connected by at least one tie, leading to an increase in individual degree, closeness and the diameter of the network. This suggests that removals can have widespread impacts on population structure and individual physiology that extend far beyond the individuals being removed. These consequences should be taken into consideration when considering removals and translocations. Temporal stability in female relationships suggests that equid social structures are multifaceted: although bonds between stallions and mares are clearly strong, long-term relationships between mares are also important in scaffolding the social network structure. These networks have clear implications for equid fitness: applying this framework across populations can provide key information about how equids structure themselves in response to demographic and environmental challenges.

MONDAY 14:00-14:20

Comparative behavior and mating systems of African wild ass (Equus africanus) and feral donkeys (Equus asinus) in arid and mesic habitats

Patricia D. MOEHLMAN1, Redae Teclai TESFAI2

1 IUCN/SSC Equid Specialist Group, Box 2031, Arusha, Tanzania 2 University of the Witwatersrand, School of Animal, Plant & Environmental Sciences, 1 Jan Smuts Avenue, Braamfontein 2000, Johannesburg, Gauteng, South Africa

The African Wild Ass (Equus africanus) is the world's most endangered equid. It persists in one of the harshest climates and terrains in the world, the Horn of Africa. Research on African wild ass behavior and mating system was done in Eritrea. In Eritrea, individuals were identified and the only stable groups were composed of a female and her offspring and dominant males were territorial. Females do associate with other females and/or males, but even temporary groups are small. Low density and low sociability in African wild ass may be due to low forage quality and availability which limits long term associations. The African wild ass has a resource defense polygynous mating system. Feral donkeys (Equus asinus) were studied in another arid habit, Death Valley, USA. In this locale similar to African wild ass research, individuals were identified and the only stable groups were mothers and their offspring and males were territorial. In Death Valley, feral donkeys had a resource defense polygynous mating system. By comparison, individually identified feral donkeys living in mesic habitats, i.e. Ossabaw Island, USA and Isla Isabella, Galapagos, Ecuador, formed harem groups (female defense polygynous mating system) that were stable or exhibited fusion/fission social behavior. This research provides information on African wild

ass social organization, mating system and daily activity patterns. The four study sites provide comparative data (per sex and age class) on daily activity patterns, inter-individual spacing, social interactions, nursing behavior and mating systems.

MONDAY 14:20-14:40

Do zebra shape resource and community level interactions within a grazing guild? Francesca PARRINI1, Elena MARIOTTI1, Cornelius J. LOUW2, Jason P. MARSHAL1 1 Centre for African Ecology, School of Animal Plant and Environmental Sciences, University of the Witwatersrand, Johannesburg, South Africa. 2 Department of Environmental Sciences, University of South Africa, South Africa.

Abundance and distribution of large herbivore populations depend on, among other things, habitat and forage attributes in an animal’s environment, interactions with others of the same species (intraspecific competition), and interactions with other species (interspecific competition and facilitation). Moreover, individuals alter their environment by using habitat and resources, which in turn influences the environment for other individuals of the same and different species. The balance between resource-level effects and community-level interactions for a particular species ultimately determines distribution patterns and population dynamics for that species and thus its conservation status. Our study aimed at investigating the relative roles of habitat, forage, and community interactions on large herbivore distribution at Telperion and Ezemvelo Nature Reserves, in South Africa, at landscape and feeding patch scale across a wet and a dry season. The Telperion section has had an increasing density of zebra (Equus quagga) which has coincided with a declining hartebeest (Alcephalus buselaphus) population. Trends like these suggest that changes in habitat and/or competition with zebra might contribute to the decline in hartebeest numbers. Generalist grazers such as zebra can handle forage of high biomass and low quality better than more specialist grazers like hartebeest. Thus, the predictions were that the Telperion section would have a

different pattern of resource utilization to the Ezemvelo section, with only zebra favouring the Telperion section where habitat and forage conditions favour generalist grazers. Contributing to those conditions are higher elevation and less frequent wildfires at Telperion, compared to more frequent fires and palatable grasses in the neighbouring Ezemvelo section that would instead favour specialist herbivores that require higher-quality forage. We also expected that the intraspecific and inter-specific spatial distribution of species across seasons would reflect the competition for resources, especially during the dry season. Additionally, factors such as fire frequency, elevation and grass characteristics were expected to interact with competition to influence spatial utilization of herbivores. We found that specialist grazers like hartebeest and black wildebeest (Connochaetes gnou) were mainly limited to the Ezemvelo section, while zebra were found in both sections, confirming that forage available in the Telperion section might be of too low quality for ruminants. The presence of burnt areas did influence spatial utilization of ruminants but not of zebra, but only at the landscape scale. Changes in season triggered changes in grass selection in all species. In the dry season, red hartebeest and black wildebeest switched to feeding patches offering more biomass, blue wildebeest (Connochaetes taurinus) accepted taller grass and zebra used less green feeding patches than in other seasons. Most importantly, the results suggest that the presence of interspecific competition between zebra and ruminants was driving space use at both spatial scales.

Diseases

MONDAY 14:40-15:00

Development and application of a non-invasive DNA metabarcoding assay to characterize complex parasitic strongyle infections in equids

Jocelyn POISSANT1, Stefan GAVRILIUC1, Jennifer BELLAW2, Libby REDMAN1, Russell AVRAMENKO1, Matthew WORKENTINE1, Emily JENKINS3, Martin NIELSEN2, Philip MCLOUGHLIN4, John GILLEARD1

1 Faculty of Veterinary Medicine, University of Calgary, Calgary, Canada 2 Department of Veterinary Science, University of Kentucky, Lexington, USA 3 Western College of Veterinary Medicine, University of Saskatchewan, Saskatoon, Canada 4 Department of Biology, University of Saskatchewan, Saskatoon, Canada

Equids are commonly infected by a large diversity of gastro-intestinal parasitic strongyle nematodes, which damage hosts by sucking blood or intestinal fluids, migrating through tissues, and potentially causing colic and/or diarrhea. While strongyle species infecting equids are known to differ in pathogenicity, non- invasive diagnostics and research on the causes and consequences of infections has so far been limited by our inability to morphologically differentiate eggs from different species and genera. As a consequence, our understanding of the epidemiology, biology and pathogenicity of equine strongyle nematodes remains very limited. We describe the development and application of a novel molecular DNA metabarcoding assay which allows discrimination of most strongyle species infecting horses. When applied to pools of infective third-stage larvae (L3) cultured from fresh feces, the assay provides highly repeatable qualitative and

quantitative assessments of mixed infections. Results from two feral horse populations (Alberta and Sable Island, Canada) and one experimental herd (Kentucky, USA), none of which had been exposed to anthelmintic drugs, will be presented as a proof of concept, including variation across biological replicates and samples collected from the same individuals over a 4 months period. In these populations, the assay identified 31 different strongyle species including three large strongyles (Strongylus vulgaris, S. equinus, and S. edentatus) and 28 cyathostomins, and uncovered significant differences in infection among individuals and populations. Various applications in horses and other equids as well as current limitations will be discussed. This new tool will open up a number of exciting research avenues and applications in wild equid parasitology, including epidemiology and evolutionary ecology research, studying species-specific pathogenicity and drub resistance, and improving diagnostic tools and management of infections in captive and endangered populations.

MONDAY 15:00-15:20

Survey of serum amyloid A and pathogen frequency in recently captured feral donkeys in Death Valley National Park, California USA

Sara JERELE 1, Lauren BRIZGYS2, Samantha MAPES3, Francisco Javier NAVAS GONZALEZ4, Eric DAVIS1, Nicola PUSTERLA3, Amy MCLEAN2*

1 International Animal Welfare Training Institute, School of Veterinary Medicine, Davis, USA 2 Animal Science, University of California Davis, Davis, USA 3 Equine Viral Disease Laboratory, Department of Pathology, Microbiology, and Immunology, School of Veterinary Medicine, Davis, USA 4 Faculty of Veterinary Sciences, Genetics Department, University of Córdoba, Córdoba, USA

Recent removal of feral donkeys from public land has increased concerns for welfare and disease transmission to other equids. Relocation process to new environments and introduction to other equids may cause disease outbreaks. Little is known about the health of feral donkeys prior to relocation. Serum Amyloid A (SAA), is an acute phase protein produced by the liver in response to inflammation from infection or injury and can be tested on site (StableLab, Sligo, Ireland), which can assist in health assessments of feral donkeys. Stall-side test can correlate with the presence of viral infections and results can inform biosecurity measures for handling these animals. Developing an understanding of pathogens and infectious diseases of feral donkeys would both help in improving their welfare and preventing transmission of pathogens to domestic equids. The objective of this study was to measure frequency of pathogens in recently captured donkeys and test for respiratory pathogens and monitoring of disease by testing an inflammatory

marker (Serum Amyloid A). Whole blood and nasal secretion were collected from 85 donkeys (Death Valley National Park); 24 retested after 60 days in long-term holding facility co-mingled with donkeys from multiple locations. Quantitative Real Time PCR (qPCR) using nasal secretions tested for Streptococus equi subspecies equi and zooepidemicus, equine influenza A (EIV), equine rhinitis A and B viruses, and herpesviruses: asinine herpesvirus (AHV)-2, AHV-3, AHV-5, equine herpesvirus (EHV)-1, EHV-4. Positive correlations were found for pathogen DNA and serum amyloid A (SAA), age, and body condition score (BCS). SAA level was correlated with age, foals having a higher SAA; but not BCS, presence of nasal discharge, or behavior. Tests were positive for asinine herpes viruses and Streptococus zooepidemicus. In conclusion, wild donkeys, from remote desert environments, carry viruses and bacteria that may cause disease in donkeys, but do not pose a significant threat to horse populations. Further research on changes in pathogen presence after donkeys are comingled with domestic equids is warranted. Additional research is needed to provide more information on domestic management systems and value of SAA in gauging health of donkeys.

MONDAY 15:20-15:40

The Darwinian adaptation of the equine intestine vascular network to Strongylus vulgaris

Luna GUTIÉRREZ-CEPEDA1,2, Coby BOLGER WILSON,2, Aránzazu MEANA 1

1 Faculty of Veterinary, University Complutense of Madrid, Spain 2 Horse1-Spillers SL, Spain

Strongylus vulgaris is a nematode parasite of the family Stronglylidae that infects members of the genus Equus. Adults are found in the large intestine, particularly in the caecum and are practically ubiquitous in the horse (with a high prevalence of 70-100% in domestic animals). It has also been described in donkeys, mules, zebras, feral horses and wild asses. Data on Spanish prevalence indicate that this parasite can be found in approximately 25% of animals with strongylids. Strongylus vulgaris is the only equine strongyle to undergo larval migration and development in the arterial system of the host, with a predilection for the cranial mesenteric artery system. The 4th stage larvae and young adults (size around 2 cm) are found therein embedded in the thrombotic material and necrotic debris of the endothelium causing various degrees of lesions (verminous arteritis) in the cranial mesenteric artery and its main branches in almost all infected animals. Thromboembolic colic is a typical consequence of the activity and development of these larvae. This problem is related to the liberation of thrombus and obstruction of arterioles or arteries of different sizes. This depends on the area of intestine affected and the flow of collateral blood supply before arterial obstruction and necrosis sets in. It has been hypothesized that the rich vascular network of anastomosis in the arterial supply of the horse intestine wall will overcome almost all interruptions to the blood supply. With this information, we can conject that that

the presence of this parasite favoured the survival of horses with a specific type of circulatory system. The selection pressure exerted by the ubiquitous S. vulgaris infection caused a Darwinian or evolutionary adaptation. The survivors were the genus equus with a richer anastomosis irrigation network. This has been demonstrated by comparing the different animal species digestive systems. The a. mesenterica cranialis is originated at the level of the 1st lumbar vertebra in the horse and immediately enters the cranial mesentery forming the root of the mesentery. In the horse this vessel subdivides at this level while in other domestic mammals it continues caudoventrally so gaining the plate-like meso-jejunum. It supplies all parts of the digestive tract, which are attached to this mesentery. Unlike other domestic animals such as the dog, pig or sheep, all compartments of the hindgut of the horse are irrigated by at least two branches of this arterial system or there are anastomosis between the different branches that ensure efficient supply of these areas even in the presence of S. vulgaris larvae development.

MONDAY 15:40-16:00

A serosurvey for selected pathogens in feral burros from Death Valley National Park, California, USA

Erin GOODRICH1,2, Amy MCLEAN3, Cassandra GUARINO1,2

1 Animal Health Diagnostic Center, College of Veterinary Medicine, Cornell University, Ithaca, NY 2 Department of Population Medicine & Diagnostic Sciences, College of Veterinary Medicine, Cornell University, Ithaca, NY 3 Animal Science, University of California Davis, Davis, California

Recent removal and relocation of feral burros from vast public lands to more concentrated holding pens, training facilities, and offsite adoption locations raises several health and welfare concerns. Very little is known regarding the common pathogens which are circulating within the wild burro population in Death Valley National Park. The aim of this study was to utilize serologic assays to assess previous exposure of burros to some common pathogens that affect equids. In late 2018, serum samples were obtained from 92 burros upon removal from the National Park Services Land. Approximate ages ranged from 4 months to 20 years with a median of 6 years (46 females and 46 males). The serum was analyzed for immunoglobulins against Equine Herpesvirus 1, hemagglutination inhibiting antibodies to Equine Influenza (H3N8), IgG and IgM antibodies to West Nile Virus, and antibodies to three different outer surface protein antigens present on Borrelia burgdorferi (the causative agent of Lyme disease). Only 3% (3/92) of the burros that were tested appeared to be protected against infection with EHV-1, 32% (29/92) demonstrated evidence of exposure to an equine herpes virus but were below the threshold for protection against EHV-1 infection, and the remaining

were completely susceptible to EHV-1 at the time of sampling. Equine Influenza H3N8 titers were all very low, ranging from no antibodies detected at a minimum dilution of 1:4 to titers of 12. All of the burros tested negative for both IgM and IgG antibodies to West Nile Virus. 7% (6/92) of the burros tested positive for at least one of the three antibodies to B. burgdorferi. This study provides useful information to consider when devising vaccination and processing protocols for feral burros removed from Death Valley National Park prior to movement to other geographical regions in the US (where Lyme disease and West Nile Virus may be endemic) and prior to exposure to equids from other areas (who may be shedding EHV-1 or Influenza). It also indicates that further research is needed to learn more about the pathogen exposure of wild equids in other areas, especially prior to relocation. The results of this study indicate that these wild equid populations may be extremely naïve and susceptible to many common equine pathogens upon removal from the wild.

Behavioral Ecology

MONDAY 17:00-17:20

Social system shapes differences in social behaviors of equids

Sarah R. B. KING1, Kathryn A. SCHOENECKER1, 2

1 Natural Resource Ecology Laboratory, Warner College of Natural Resources, Colorado State University, Fort Collins, CO, USA 2 U.S. Geological Survey, Fort Collins Science Center, Fort Collins, CO, USA

African wild asses evolved in highly arid desert ecosystems whereas horses evolved in relatively mesic steppe grasslands. These different environments influenced the development of different social systems (resource defense polygyny and female defense polygyny respectively). Domestic horses retained the female defense polygyny social system of their ancestors, but reports on ass social structure are mixed. Feral asses (known as burros) have rarely been studied, thus behavior and social system differences between horses and burros in a ‘wild’ state are unclear. We used the same methods to examine horses and burros in a free- roaming setting, enabling us to compare the two species directly. Observations took place at two independent horse populations in western Utah (2017-2018), one burro population in Arizona (2017-2018), and one burro population in central Utah (2016-2018). Animals were individually identified by natural markings or human- applied freeze-marks, and located via radio telemetry. Every individual with a telemetry device was observed at least monthly throughout the year to gather data on group composition. Observations on social behavior were conducted March to September annually, resulting in over 2,000 animal-hours of observations on each species. Similar to their wild counterparts, feral horses and burros had different

social groupings: horses were in relatively stable groups of adult females and one male, whereas burros tended to be in groups of females alone, or with one or more males. Horses and burros had similar feeding and standing rates, but burros lay down more, and moved less than horses. Time budgets were similar between species and sexes within each species. Burros exhibited both more affiliative and agonistic behaviors than horses and vocalized more often. In horses, males were responsible for almost all the social behaviors observed, whereas in burros they were conducted more by females. The different evolutionary history and relative adaptation to arid environments of the two species is thus reflected in their social behavior. While female-defense polygyny was evident in horses, there was no evidence of resource defense polygyny in burros, although fission-fusion dynamics similar to other non-harem forming equids was seen. It is possible that burro males have adopted a different breeding strategy. Differences in social networks between the species have implications for how these animals are managed in a feral state and wider applications for the conservation of threatened equid species.

MONDAY 17:20-17:40

Internal and external effects on grouping of a fission-fusion society of female Asiatic Wild Asses (Equus hemionus) in the Negev desert, Israel

Ariel ALTMAN1, Tamar BEN-NUN2, Gili GREENBAUM1, Sharon RENAN2, Shirli BAR-DAVID1, Amos BOUSKILA2, Alan TEMPELTON3

1 Mitrani Department of Desert Ecology, Blaustein Institutes for Desert Research, Ben-Gurion University of the Negev, Israel 2 Department of Life Sciences, Ben-Gurion University of the Negev, Israel 3 Department of Biology, Washington University, St. Louis, USA

In arid environments many equid populations present malleable social properties including frequent changes in group composition and size. One possible evolutionary explanation may involve selection pressures that stem from the patchy and unpredictable resource distribution that characterize arid habitats. Group size is known to affect important fitness determinants such as predation risk, disease transmission, genetic variation and reproduction. In equids that inhabit arid zones, however, little is known on internal and external properties that affect group size. We studied group size variation of female Asiatic wild asses (Equus hemionus) with a fission-fusion social structure. We focused on two subpopulations in the Negev desert, Israel, which differ in size and the environment they inhabit. Internal and external effects were examined including: genetic relatedness, the reproductive state of females, and environmental variables such as temperature, wind speed, distance to water and vegetation indices. We employed sampling technics, both in the lab and in the field, tailored to suit this elusive species, including drone photography, remote sensing, GIS analysis and non-invasive genetic sampling. Fecal samples were collected from observed female groups and amplified by PCR

using microsatellite markers and mitochondrial DNA sequence for the respective genetic relatedness and maternal kinship estimations. Groups of both subpopulations were not found to be more genetically related than expected by chance (Permutation tests: P = 0.18; P = 0.47). The reproductive state of females in the groups (lactating or not) was similar among group members, but it affected group size (ANOVA, P < 0.001). Analysis of environmental variables showed that in both subpopulations vegetation affected group size, yet, additional parameters, such as wind speed, differed between subpopulations. The habitat of the larger subpopulation was windier than that of the smaller subpopulation, yet, the effect of wind on group size was significant only in the small subpopulation. By comparing hourly wind measurements with trapped ectoparasites in situ of both subpopulations, we found correlative support to the observed group sizes which fits fly relief mechanisms reported from other equid systems. This may point to a trade-off between two selection forces simultaneously acting on fly relief mechanisms: equid population size and the environment (wind and flies). Both forces may affect group size in socially fluid populations. Our results underscore factors that are well known in their potential to affect group size, but also some that are less known in this particular context. Yet, for species inhabiting arid zones, refining our understanding of all of these factors is essential for projecting the social-demographic response to climate change, so that appropriate conservation policies can be established.

MONDAY 17:40-18:00

Anti-predator defence behaviour in feral horses

Lucy REES1

1 Asociación Caballos Salvajes, Piornal, Cáceres, Spain.

Although the horse is a prey animal, its behaviour in relation to predator threat has not been described. A population of around 140 feral horses sympatric with puma and jaguar in the llanos of Venezuela was observed during five consecutive breeding seasons. Their defence tactics fell into two classes, precaution and reaction. Precautionary measures comprise: group living; the retrieval of straggling foals by the stallion; avoidance of areas that facilitate predator attack; and vigilance. Reaction to possible visible threat comprises: alarm and orientation; transmission of alarm to other band members and other nearby bands, with a primary response of bunching, or cohesion, of band members and of bands together; assessment of threat by the stallion and other in-band males; if flight developed, synchronicity of reaction and movement; and band reassembly after a stampede. The variation in response was considerable, ranging from assertive investigation by the stallion, through band movements away from threat of differing speeds and duration, to full- blown stampedes involving the whole population. Sounds provoked instant stampedes without assessment. All band or mass movements exhibited the algorithm for self-organizing or emergent behaviour – cohesion, synchrony and collision avoidance – seen in other flocks, herds, shoals and swarms. No fixed leaders or initiators of change were seen during flight; during stampedes, the front-running faster youngsters changed direction if the older horses behind did so. Stallions played a major role in vigilance, foal

protection and assessment of threat. Evidence of the same algorithm during maintenance activities suggests that feral horse bands are self-organising cooperatives without fixed leaders whose adult members assume different roles in raising foals, the band’s most vulnerable members, but unite in collective defence. The final cause of in-band aggression, whose proximate causes vary but do not include in-band resource competition in feral horses, is to establish respect for individual space, thus avoiding collision in flight.

TUESDAY 10:00-10:20

Overmarking by adult females in four equid species serves to strengthen the social bonds and to support group cohesion

Jan PLUHÁČEK1,2 , Vladimíra TUČKOVÁ3, Radka ŠÁROVÁ2, Jitka BARTOŠOVÁ2, Sarah R. B. KING4

1 Ostrava Zoo, Ostrava, Czech Republic 2 Department of Ethology, Institute of Animal Science, Praha - Uhříněves, Czech Republic 3 Department of Zoology, Faculty of Science, University of South Bohemia, České Budějovice, Czech Republic 4 Natural Resource Ecology Laboratory, Warner College of Natural Resources, Colorado State University, Fort Collins, USA

Scent marking by overmarking deposits made by conspecifics has been well studied in male mammals, but this phenomenon remains poorly understood and rarely studied in females. Furthermore, in socially living mammals like most ungulate species many of the established hypotheses cannot be applied, leaving the function of female overmarking unexplained. In this study, we suggest and test a novel hypothesis: maintenance of social bonds. We examined the occurrence of adult female overmarking in equids and test three classical hypotheses which may explain this phenomenon: group cohesion, intrasexual competition, and social hierarchy. We observed all African equid species (Equus africanus, E. grevyi, E. quagga, E. zebra) in five Czech zoos: Brno, Dvůr Králové, Liberec, Ostrava, and Ústí nad Labem. In total, we recorded 4668 eliminations by 130 individuals split in 15 various groups; 327 of these events were overmarked by 69 individual females (out of 74 observed females) representing all species. Adult females overmarked

individuals of each age and sex category, but mostly other females and foals. The rate of overmarking within a dyad of adult females increased with increasing ‘friendship’ of the dyad, thus demonstrating for the first time that overmarking can serve to support social bonds in mammals. In addition, we found a higher overmarking rate among females in the temporary absence of a stallion than in his presence suggesting partial support for the group cohesion hypothesis. On the other hand, we found no evidence supporting hypotheses of intrasexual competition and social hierarchy. Based on our results, we suggest that the use of overmarking to maintain social bonds may be more important in mammals like equids that live in groups of unrelated individuals.

TUESDAY 10:20-10:40

Relationships between reproductive state and social connections in reintroduced takhi (Przewalski’s horse: Equus ferus przewalskii) family groups in Mongolia

Hannah DAVIE1,2, Carol HALL1, Samantha BREMNER-HARRISON1, Claudia FEH2

1 School of Animal, Rural and Environmental Sciences, Nottingham Trent University, Southwell, Nottinghamshire NG25 0HU, UK 2 Association pour le cheval de Przewalski:TAKH, Station Biologique de la Tour du Valat Le Sambuc, 13200 ARLES, France

Although the first phase of takhi (Przewalski’s horse; Equus ferus przewalskii) reintroduction to Mongolia has been successful, the three established populations remain at risk from geographic isolation, resource competition with humans and livestock, and hybridisation with modern domestic horses. Individuals in family groups (or harems or bands) that stay close together and form herds with other families are less apt to be isolated from other takhi and to hybridise with domestic horses. Larger, more cohesive herds are also easier to manage to minimise competition for water and forage with humans and livestock. The aim of our research was to determine which individuals in takhi family groups at the Seer reintroduction project in Khomyn Tal in western Mongolia formed the most social connections, negative and positive, with members of their own family or with members of other families. Using social network analysis, we found a significant difference in social activity relative to age and sex. Within family groups, adults, mostly mares, maintained personal space by displacing other individuals, mostly juveniles. Juveniles, primarily colts, were most involved in within-family play and

social grooming. Family stallions maintained the borders between families within the herd, and stallions and colts engaged in most between-family play and grooming. Our results indicate that early social experience in juveniles and later social tolerance in stallions influenced the cohesiveness of the Seer herd, but within-family social structure was shaped by the social tolerance of mares. Our results demonstrate the varied roles assumed by individuals of differing reproductive state in natural Type I equid social groups and inform adaptive management of takhi populations and groups and herds of other Type I equids, including feral and domestic horses.

TUESDAY 10:40-11:00

The causes and consequences of zebra hybrids

Dan RUBENSTEIN1

1 Princeton University, Department of Ecology and Evolutionary Biology, Princeton, 8544, United States

Equid species in general range widely with species only showing small biogeographic zones of overlap. This is certainly true for zebras, but where they do overlap, hybrids occasionally form. Why they do so and the resulting behavioral, ecological and disease related consequences reveal subtle, but important evolutionary gene-environment interactions. The diversity of behavioral responses exhibited by males and females, niche shifts and the ability to modulate parasite levels will be explored in this talk.

TUESDAY 11:20-11:40

Effect of availability and temporal variability of resources on the distribution and movement of feral burro (Equus asinus) populations

Kathryn A. SCHOENECKER*1, 2, Sarah R. B. KING2, Saeideh ESMAEILI3

1 U.S. Geological Survey, Fort Collins Science Center, Fort Collins, CO 80526, USA 2 Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, CO 80523, USA 3 Department of Zoology and Physiology, University of Wyoming, Laramie, WY 82071, USA

Distribution and habitat use of wild animal populations are the outcome of evolutionary processes in which individuals adapt to resource dynamics in the surrounding landscape. Feral species have not undergone such processes and have been introduced into areas that may be different from where they evolved. Despite this, feral animals can adapt to a wide variety of environmental conditions. Understanding interactions of feral populations with their environment, both biotic and abiotic factors, can improve predictions of behavior of native populations in changing environments. Donkeys (also called wild burros, or feral burros; Equus asinus), were domesticated from African wild ass (E. africanus) and first introduced to North America in 1495. They arrived in large numbers during the nineteenth century gold rush as pack animals and were used in mines and ore- grinding mills. After the mining boom ended many were released and became feral populations. We conducted research on the ecology and demography of feral burros to inform management of populations in the western USA and to contribute to conservation and management of African wild ass, which are the least studied

but most endangered equid species. We determined monthly home range size and movement parameters (step length, movement rate, and total distance traveled) in two feral burro populations between 2016 and 2019 using 30 GPS radio collared females in each population. One population inhabits a high desert ecosystem in Utah with cold snowy winters and hot dry summers. The other inhabits the Sonoran Desert of Arizona, a hot desert climate. We studied effect of habitat variables (ambient temperature, vegetation biomass, distance to water) on monthly home range size (95% fixed kernel density estimate) and movement rate of both populations. We used generalized linear mixed models with individual-year identity as a random effect and habitat variables, quantified within monthly home ranges and along four-hour steps, as fixed effects. Burros in Utah had larger and more variable monthly home ranges, longer steps and total distance traveled, and faster movement rates compared to burros in Arizona. We found total vegetation biomass and temporal variability in biomass and distance to water positively affected home range size in both populations. Higher ambient temperatures were associated with smaller monthly home range sizes. The best supported model using Akaike’s Information Criterion indicated temporal variability in distance to water and vegetation biomass had the strongest influence on burro home range size. In addition, biomass and temperature positively affected step length and movement rate. Overall, less variability in resources, regardless of quantity, resulted in smaller home ranges and more restricted movements in both populations. Management that reduces variability in resources, such as permanent water, could help restrict burro movements. We expect African wild ass home range and movement patterns to be more like burros in Arizona than Utah because the Sonoran Desert is more similar to the Horn of Africa where they evolved. Considering the unpredictable environment, variable resources and human disturbance in current African wild ass range, we suggest conservation and management of the species is highly challenging.

TUESDAY 11:40-12:00 – Student

Key resources competition threatens viability of the critically endangered African wild ass (Equus africanus) population on Messir Plateau (Eritrea)

Redae T. TESFAI1,2, Francesca PARRINI2, Norman OWEN-SMITH 2, Patricia D. MOEHLMAN3

1 Ministry of Agriculture/Forestry and Wildlife Authority, P. O. Box 1048, Asmara, Eritrea. 2 Centre for African Ecology, School of Animal, Plant & Environmental Sciences, University of the Witwatersrand, Wits 2050, South Africa 3 EcoHealth Alliance, 460 West 34th St, New York, NY 10001-2320, USA

Globally, the African wild ass (Equus africanus) is at risk of extinction in the wild. Major threats to its survival are hunting, competition with livestock, and interbreeding with the domestic donkey. An important but rare population exists on the Messir Plateau in the Danakil ecosystem of Eritrea. Long-term data on reproduction and survival rate of African wild asses are limited, making it difficult to predict the population trend. Therefore, we assessed the population's potential viability given certain assumptions. An age-structured matrix model was used to project the population growth rate generated by assumed vital rates. The probability of population persistence over 50 years was estimated allowing for stochastic dynamics, density dependence, influences of rainfall on carrying capacity, and episodic disturbances reducing survival rates. Scenarios used either historical rainfall patterns or potential future rainfall with the same or amplified coefficient of variation. We contrasted two estimates of carrying capacity based either on recent population assessments, or on the potential capacity of the study site. Our models also compared realistic versus optimistic scenarios for age at first

reproduction and potential longevity. The probability of persistence of a population of 18 females was reduced to < 50% in the worst-case scenario. With a potential carrying capacity of 37 females, persistence was projected to be almost 100% under all scenarios. The greatest threat to population viability seems to come from livestock impacts on the resources that limit the African wild ass population size under anticipated climate change scenarios. Faecal samples were analysed and stable carbon isotopes (13C/12C) were compared to assess the dietary overlaps of African wild ass and livestock. Replicates from African wild ass and domestic donkey in pre-rainfall months were compared with rainfall months to assess the relative nutritional levels. The preliminary results indicated that the African wild ass faecal nitrogen content was significantly lower (P < 0.05) than the domestic donkey comparing the same rainfall months. The lower nutrients levels of the African wild ass in rainfall months may be due to the high concentration of cattle in the area. This study provides useful information on how domestic herbivores may threaten the survival of critically endangered herbivores in the very arid ecosystem.

Conservation

TUESDAY 12:00-12:20

Kiang conservation in India: A landscape approach

Yash Veer BHATNAGAR1, Sajid SULTAN2, Abhishek GHOSHAL1

1 Nature Conservation Foundation & Snow Leopard Trust, 1311, “Amritha”, 12th Main, Vijayanagar 1st Stage, Mysore 570 017, India 2 Regional Wildlife Warden, J&K Department of Wildlife Protection, LAHDC Complex, Mini Secretariat Leh, 194101, India

Kiang are an iconic species spread over c. 20,000 km2 of rolling high altitude mountains in four provinces of India. Robust population assessments are not available but estimates over the past three decades point to a stable population of less than 3,000 individuals, most of them being in Ladakh, Jammu & Kashmir. The perception among herder communities and policy makers of the region is that the kiang population is increasing to the extent that they are depleting forage in the pastures important for the thriving cashmere economy. Further, in the past two decades, climate change has allowed crops to be cultivated at elevations even higher than 4,000m, and the issue of crop depredation by kiang, and in turn retaliatory chasing and harassment of kiang by farmer communities is on the rise. These perceived and actual conflicts affect conservation and management of kiang in the region. Additionally, kiang faces emerging threats posed by fencing of important foraging sites by herder communities, growing defence and tourism infrastructure, off-roading often resulting in harassment from long chases by tourists, disturbance from the expanding road network, and depredation by free- ranging dogs. Almost the entire kiang habitat in Ladakh is within the Changthang

Wildlife Sanctuary (c. 13,000km2), with a small portion in the Karakoram Wildlife Sanctuary (c. 4,500km2) and the Hemis National Park (3,300km2). Kiang habitats in rest of the Indian Himalaya in States of Himachal Pradesh, Uttarakhand and Sikkim (c. 3,000 km2) are mostly outside legal protection. In this paper, we propose and discuss comprehensive conservation measures for kiang that include, (i) crop protection - community guarding and appropriate fencing of crop fields; (ii) Moist meadow protection - avoiding fencing of moist meadows to conserve foraging areas of kiang, while providing fodder to herder communities where needed; (iii) forging partnership with government departments mandated for pasture development, soil conservation and infrastructure development to ensure green development measures; (iv) developing partnership with agencies related to defence, tourism and rural development to manage free-ranging dog population and garbage; (v) improving livelihood security of the local nomads - community- based handicrafts and wildlife focused tourism enterprises, generate other diversified sources of earning through innovative value-added livestock products; (vi) targeted research to help management, which includes regular monitoring of kiang populations, identifying and protecting rutting, breeding and nursery sites, monitoring the dynamic livestock density and grazing management. Effective kiang conservation further requires plans and processes for coordinated and integrated efforts with local communities, across development and welfare sectors, but over vast landscapes. Fortunately, in India, participatory landscape level conservation of the snow leopard and its sympatric species is enabled under at least three recent schemes of the Ministry of Environment, Forests and Climate Change, Government of India. In this framework, we discuss inclusive planning and institutional mechanisms for effective and long-term landscape level conservation of kiang.

TUESDAY 12:20-12:40

Reintroduction, persistence and local extinction of central Asian kulan in Turkmenistan and Kazakhstan

John D. C. LINNELL1, Petra KACZENSKY1, Albert SALEMGAREYEV2, Steffen ZUTHER3, Eldar RUSTAMOV4, Shirin KARRYEVA4, Ralph KUEHN5

1 Norwegian Institute for Nature Research, Trondheim, Norway 2 Association for the Conservation of Biodiversity of Kazakhstan, Nur-Sultan, Kazakhstan 3 Frankfurt Zoological Society, Frankfurt, Germany 4 Important Bird Area Project, Ashgabat, Turkmenistan 5 Department of Ecology and Ecosystem Management, Munich Technical University, Munich, Germany

By the middle of the 20th century kulan in central Asia had been eradicated from all of their former distribution except in the Badkhyz reserve in southern Turkmenistan. During the period from the end of World War 2 until the collapse of the Soviet Union kulan from Badkhyz were translocated to multiple sites in Turkmenistan and Kazakhstan. However, there has been almost no information available about the status of these populations since then. Since 2011 we have been conducting wildlife surveys in all areas of Turkmenistan and Kazakhstan where kulan have been known to occur. Most of the populations in Turkmenistan (including Badkhyz) are now functionally extinct, with some very small groups of individuals currently ranging in three fragmented areas. In Kazakhstan, two of the reintroduced populations (Barsa Kelmes and Altyn Emel) appear to be doing well with populations of several hundred animals each. Since 2017, these two populations are now being used as a source for a further reintroduction into the

Torgai steppes of central Kazakhstan (part of the Altyn Dala initiative). This presentation will give details of the status of all populations, present the latest results from genetics and telemetry studies, and the first experiences with the new reintroduction project in Kazakhstan.

TUESDAY 12:40-13:00

Return of the wild horses – reintroduction of Przewalski’s horses from Europe to western Mongolia

Jaroslav ŠIMEK1, Roman VODIČKA1, Jan MAREK1, Barbora DOBIÁŠOVÁ1, Ganbaatar OYUNSAIKHAN2, Miroslav BOBEK1

1 Zoo Praha, U Trojského zámku 120/3, 171 00, Czech Republic 2 Great Gobi B SPA, Takhin Tal, Mongolia

The Przewalski horse was extirpated from the wild at the end of 1960’s and survived only thanks to the captive breeding. In late 1980s’/early 1990s’ the first transports of captive bred horses were organized back to their homeland in China and Mongolia. In 1992, the reintroduction project was initiated in the area of the Great Gobi B SPA in southwestern Mongolia. Within 13 years, a total of 89 horses were transported to the area. After initial stagnation, the promisingly growing population was hit by severe loss due to heavy winter 2009/2010. At that time, the need for another support arose. Within a scope of long-term Takhi project of International Takhi Group, Prague Zoo as its member organized the new set of transports of Przewalski horses from Europe to Mongolia under the project name ‘Return of the Wild Horses’. From 2011 to 2018, 31 horses were released in Mongolia. The horses for the reintroduction are selected in cooperation with European breeding program (Przewalski Horse EEP) and many criteria are taken into account (e.g. age, sex, genetics, pregnancy status, hooves condition, veterinary status, body condition, response to stress). The horses from various facilities across Europe are collected at Dolni Dobrejov, Prague Zoo’s breeding and acclimatization facility, where the final set of veterinary testing is performed. From more candidates, the best four are selected for the shipment in a given year. The aerial

transport is performed by the military plane of the Czech Army, just for operational cost. The horses are transported in crates specially designed and accustomed for a long trip to minimise the risks. Each horse is loaded to the crate at the moment of standing up from anaesthesia and remains under long-acting neuroleptics during the transport and few days after releasing into acclimatization enclosure. The stay in the crates is around 30 hours; the trip consists of a few hours on road, 16 hours on plane (with two stopovers) and another few hours on road in Mongolia. The horses are accompanied by experienced veterinarian and head keeper and are regularly fed and watered. Upon release from the crates, the horses stay in the acclimatization enclosure under supervision of Great Gobi B rangers over the first winter. Next spring, the horses are treated against piroplasmosis and released into the wild (soft release), in companion of local horse(s). Out of 31 horses, there were two deaths within the acclimatization period in the fence, one suspected colic and one possible consequence of Babesia sp. infection. Within the first year after the release, one loss was reported. At least 17 mares (out of 22 possible) foaled at least once or were recorded pregnant. There is currently 25 living offspring of transported horses and 2 foals in the second generation. The project is financially supported mainly by the City of Prague, Czech Ministry of Environment, and various zoos and breeding institutions (Cologne Zoo, Domaine des Grottes de Han, Gaia Zoo Kerkrade, Heinz Sielmann Stiftung, Leipzig Zoo, München Zoo, Taipei Zoo, Tierpark Berlin, Wildpark Grünau).

Population dynamics, viability, and estimation WEDNESDAY 10:00-10:20

Warriors using SMART to monitor endangered Grevy's zebra in northern Kenya

Belinda LOW1, Sheila FUNNELL1, Justine CORDINGLEY1, Annsarah WANGUI1, Joshua LABARAKWE1

1 Grevy's Zebra Trust, Mukoma Road, 15351-00509 Nairobi, Kenya

Understanding the trends and distribution of wildlife populations can be challenging, especially where the focal species is difficult to observe and is distributed across a range that covers many thousands of square kilometres. This can be expounded by a limited research budget and lack of personnel that are capable of using traditional research tools to collect data. This can lead to a lack of understanding about the species and lead to serious challenges in detecting and reversing population declines. One of the most important yet least protected populations of Grevy’s zebra is found in Laisamis, northern Kenya. Major threats to Grevy’s zebra in Laisamis include limited access to water, loss of grazing habitat and sporadic poaching. As a response to these threats, GZT recruited 10 Warriors from five communities in Marsabit County, Kenya. Using the Spatial Monitoring and Reporting Tool (SMART) and CyberTracker application on their android phones, the Warriors collect data on Grevy’s zebra location, numbers, age and reproductive status and Grevy’s zebra associations with different types of habitat, wildlife, livestock and settlements. The Warriors also use GPS-enabled cameras to document their sightings of Grevy’s zebra. We will show the

conservation benefits of recruiting and training local community members to collect data on this endangered species both for understanding the species population trends and distribution and also for fostering positive local community attitudes towards the species. Descriptive analyses of the Warriors data reveal wet and dry season distribution hotspots for the species and the localities favoured by lactating females and their foals. The results reveal the disappearance of the species from previous hotspots, and the importance of previously unidentified areas to the survival of the species during the dry season. Here, we advocate for the use of modern technology to help local communities monitor wildlife with more precision and efficiency and show that the Grevy’s Zebra Trust Warrior Program can generate valuable insights for Grevy’s zebra conservation at local and national scales and actively promote conservation of the species through positive behavioural changes from the local community towards the species.

WEDNESDAY 10:20-10:40

Population performance of the endangered Grevy’s zebra (Equus grevyi) on the Lewa – Borana landscape, Meru, Kenya

Timothy KAARIA1, David KIMITI1

1 Lewa Wildlife Conservancy, Meru, Kenya

The world’s population of the endangered Grevy’s zebra stands at less than 3,000 individuals. This is after a drastic decline from over 15,000 in 1970’s mainly due to habitat degradation and poaching for meat and medicinal purposes. Currently, over 90% of the population inhabits northern Kenya and the remaining few in Ethiopia. Although current estimates suggest a stable and relatively self-sustaining population, continuing habitat degradation and loss continue to impede the long- term recovery of Grevy’s zebra in Kenya. From 2015 to 2017, we took photographs, GPS position, and age and sex of individual Grevy’s zebra monthly and then identified individuals using the Image Based Ecological Information System (IBEIS) software (now being developed as part of Wildbook). IBEIS utilized pattern matching algorithms to analyse Grevy’s zebra images and uniquely identify individual animals by the stripe patterns on their coat. We also collared different prides of lions with VHF-GPS enabled collars to monitor their movements as well as track them to understand their diet by analyzing the scat collected. This supplemented opportunistic sightings by field rangers. A total of 210 foals were born between 2015 and 2017 with male to female sex ratio of 1:1. Additionally, close to 40 % of foals graduated yearly from the vulnerable age class of 0-12 months with similar male to female sex ratio. We further sub-divided the foal age class into three age sub-classes of 0-3 months, 3-6 months, and 6-12 months to discern the most vulnerable age sub-class. Our results show that the

chance of a foal surviving increased significantly as it graduated from one age sub- class to another, with 0-3 month being the most vulnerable (χ2 = 42.343, df = 2, p = 0.0001). In 2015 and 2016, Jacob’s Selectivity Index was 0.3, showing slight predation preference on Gervy’s zebra, but reduced to 0.0 in 2017, indicating neither preference nor avoidance. These results indicate the impact predation has had on suppressing Grevy’s zebra foal survival and overall growth rates on the Lewa-Borana Landscape and underscore the need for unique identification of individual animals to allow for more accurate demographic estimates.

WEDNESDAY 10:40-11:00

Causes of mortality, survival rates and reproductive patterns in plains zebra in the Serengeti Region, Tanzania

Peter J. P. GOGAN1, Ronald O. SKOOG2,3

1 U.S. Geological Survey, Fort Collins Science Center, Fort Collins, Colorado 80526, USA 2 Caeser Kleberg Research Program in Wildlife Ecology, Texas A&M University, College Station, Texas 77843, USA 3 Deceased

Numbers of zebra in the Serengeti region have remained remarkably constant over the past 50 years while the abundance of coexisting wildebeest and Thompson’s gazelle populations have changed markedly. However, there has been little understanding of factors limiting zebra population growth. We present historical data on zebra demographics to address potentially limiting population growth. We identify causes of mortality among Serengeti zebra based upon examination of animals dying of natural causes encountered in the field. We assess zebra age structure and survival rates derived from the sample of carcasses of zebra dying of natural mortality supplemented with a sample of fresh skulls collected in the field. Both samples are potentially skewed toward older age classes, attributable to scavengers consuming completely the remains of younger animals. Additionally, we necropsied zebra culled along Serengeti National Park’s eastern boundary and estimated zebra age structure, survival rates and reproductive rates after taking in to account potential shooter biases in selection of culled animals. We compare survivorship estimates from the sample of natural mortality to estimates of survival obtained from the culled sample. These data are being evaluated for suitability of

use in an integrated population model to advance an understanding of Serengeti zebra demographics and to identify factors potentially limiting zebra numbers. Data were gathered while the authors were affiliated with the Serengeti Research Institute, Tanzania National Parks and with cooperation of the Tanzania Game Department. We are grateful to these organizations for logistical support.

WEDNESDAY 11:20-11:40

Social environmental effects on dispersal success and sex differences in survival drives adult sex-ratio bias in feral horses

Philip D. MCLOUGHLIN1, Jocelyn POISSANT2, Charlotte REGAN1

1 Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, SK S7N 5E2, Canada 2 Department of Ecosystem and Public Health, Faculty of Veterinary Medicine, University of Calgary, 3280 Hospital Dr. NW, Calgary, AB T2N 4Z6, Canada

Adult sex ratio has substantial impacts on individual behaviour and life history and hence population dynamics. We used data from the long-term, individual based study of feral horses on Sable Island, Canada, to shed light on the potential causes of bias in the adult sex ratio of wild animal populations. Over the past decade, as density increased and now fluctuates around 500 horses, it has become apparent that this population is becoming increasingly male biased, particularly in the adult portion of the population. This phenomenon has been observed in other feral horse populations; however, little is known of its causes and consequences. We quantified the temporal and spatial patterns in pre-adult and adult sex ratios, determined the demographic causes of adult sex-ratio bias, and investigated the role of the social environment in mediating sex-differences in demographic parameters. We found that the sex ratio was even in foals, yearlings, and subadults, but was significantly male-biased in adults (4+ years), with some portions of the island being more biased than others. Survival analyses indicated that sex- ratio bias was driven by sex differences in the survival of sub-adults and adults, with females having lower survival probability in both cases. Survival differences of adult males versus adult females appears to be driven by the reproductive costs

experienced by females. While trading off survival for reproduction has been observed in feral horses before, we also observed that survival differences of subadult males and females was driven by a sex difference in the survival of dispersers. Upon further investigation, we found that the survival of dispersing females was influenced by characteristics of both pre- and post-dispersal bands, as well as by the number of bachelor males in the pre-dispersal area. In contrast, the survival of male dispersers was not significantly influenced by any of these factors. This suggests that local social environment and friction it entails is an important factor influencing the survival of sub-adult females in this population, and that male sex-ratio bias may be self-perpetuating due to the effects of male presence on band structure and the survival of female dispersers. Further, it suggests that even where both sexes disperse, differences in the dispersal process may lead to sex- ratio variation. The extent to which these dynamics generalize to other feral horse populations and possibly reflects an evolutionary stable strategy is an outstanding question. These dynamics also have implications to the direct and indirect competitive ability of feral horses in comparison to sympatric ungulates and conservation of the latter.

WEDNESDAY 11:40-12:00

Demographic changes in the Hungarian Przewalski horse population: the first 20 years

Viola KEREKES1, Istvan SANDOR2, Katalin OZOGANY3, Zoltán BARTA4

1 Hortobagy National Park Directorate, Hungary 2 Budapest Zoo and Botanical Garden, Hungary 3 Department of Evolutionary Zoology and Human Biology, Debrecen University, Hungary 4 MTA-DE Behavioural Ecology Research Group, Department of Evolutionary Zoology and Human Biology, Debrecen University, Hungary

The Hortobagy National Park established a semi reserve to maintain the special ecosystem of Hortobagy grassland with large herbivores and minimal human intervention. Within the framework of the Pentezug project (initiated in 1997, by Cologne Zoo, Germany, and Hortobágy National Park, Hungary) twenty three Przewalski horses (Equus ferus przewalskii) from different Zoological Gardens were set free until 2007 in a large semi-reserve called Pentezug in the Hortobagy, the Hungarian Steppe. The number of horses has grown quickly until 2013, when different strategies of population control, most frequently immunocontraception treatment of selected females, had to be implemented. Since then the population growth rate has been slowing down. Twenty years after the initiation of the project, approximately 300 individuals were grazing freely in Pentezug. The demographic changes in this horse population can be summarized as follows. First, the females give birth for their first foals later than at the beginning of the project (at the age of 3-4 years instead of 2), most likely due to higher density of horses. Second, the gene diversity remained stable (around 0.8) over the years, which can be the

consequence of natural social system of horses. For example, young individuals always leave their natal group. In the closed population, however, the inbreeding coefficient started to grow slowly. Third, horse harem groups united into a large herd, with bachelor groups positioned at the edges of the herd. Based on our observations we concluded that not only population monitoring but population control is also critical for running projects of Przewalski horses for a long time.

WEDNESDAY 12:00-12:20 – Student

Can gut health provide a mechanistic link between diet and performance within the Cape mountain zebra (Equus zebra zebra) meta-population?

Jake BRITNELL1,2, Corey LEE1, Rachael ANTWIS3, Sureshnee PILLAY4, Jennifer GIANDHARI4, Tulio de OLIVERIA4, Graham KERLEY5, Susanne SHULTZ1

1 School of Earth and Environmental Sciences, University of Manchester, United Kingdom 2 Chester Zoo, United Kingdom 3 School of Earth and Life Sciences, University of Salford, United Kingdom 4 KwaZulu-Natal Research Innovation and Sequencing Platform (KRISP), University of Kwa-Zula Natal, South Africa 5 Centre for African Conservation Ecology, Department of Zoology, Nelson Mandela University, South Africa

Human-induced environmental change has, at an unprecedented rate, led to fragmentation and conversion of historic habitat. These changes have resulted in huge declines in population number and increased the extirpation risk of many populations. Furthermore, contemporary species ranges may be biased to marginal parts of their fundamental/historic niche which may predispose species to poor resilience through comparatively low survival and reproductive rates. However, the physiological aspects of marginality, leading to poor survival or reproductive rates under stressful conditions, are not well understood. Physiological factors such as gut health have important implications for the nutritional status, health and fitness of individuals within populations. Gut health is a multifactorial trait which includes the microbiome and parasitological communities of the gut as well as the diet of

the animals. All three of these factors are assumed to interact; such that diet may lead to variations in microbial communities which may in turn impact parasite burdens or vice versa. Gut health therefore may be an important link between biotic and abiotic conditions and an animals’ ability to persist in that environment. However, multiple aspects of gut health have rarely been analyzed within the conservation ecology framework. Here, we test how the factors of gut health interact and how these impact the resilience and performance of Cape mountain zebra (Equus zebra zebra) across a cline of habitat qualities throughout the metapopulation. We analyzed %grassiness and parasite load across 9 populations to investigate the impact of vegetation, as a proxy of diet, on parasite burden. Furthermore, we investigated the microbiome composition from 47 individuals across 5 of these populations to evaluate the impact and interactions of variable habitat quality, microbiome and parasite communities on the performance of these Cape mountain zebra populations. Zebra occupying low-quality habitat (low grass abundance investigated via point transects) had higher parasite egg count (investigated via faecal egg count). Firstly, we predict that zebra occupying low- quality habitat will have reduced diversity of gut microbial communities, analysed via 16S ribosomal RNA gene sequencing. Secondly, we predict that populations with lower gut microbial diversity will have comparatively high faecal egg counts revealing an interaction between microbial composition and parasite susceptibility. Thirdly, we predict populations of lower gut microbiome diversity and high parasite burden will correlate with records of poor performance, thus providing a mechanistic link between environmental factors and performance. Our research will help to uncover the interactions between different aspects of gut health and how these relate to population resilience, density and performance with implications for conservation management.

WEDNESDAY 12:20-12:40

Equids and humans: a historical examination of global psycho-ecology influencing emotions

John Derek SCASTA1

1 University of Wyoming, Laramie, Wyoming, USA

Horses are iconic megafauna that elicit strong emotions from humans globally. From the often burgeoning feral horse herds, to the introduced herds for rewilding purposes, horses are a popular topic that stimulate lively debates about animal welfare and biological conservation. In some instances, the emotions humans express for horses can also hinder administrative actions. While human emotions are important for motivation enhancing awareness and action, emotions may not always align with data. For the emotions surrounding feral horse management, I review the history of horse domestication, the contemporary understanding of the psychology of human-horse relationships, and the status of feral horse ecology and management. Domestication of horses approximately 5500 years ago in Eurasia coupled humans with horses in a symbiotic relationship where the two functioned almost as one. This domestication process enabled the spread of human genes and food production. Fundamentally, human reliance on horses became a working relationship founded on trust and respect that facilitated bonding through the overcoming of adversity that likely deepened human-horse emotional connections – a coevolutionary process for which there are few analogous examples. While human reliance on horses has waned in developed countries recently, it has been humans that have largely facilitated the spread of horses through space and time. Contextualized with contemporary data, feral horse populations in the United States and Australia have increased dramatically. Consequently, horses have had

negative effects on native ecosystems in some areas, and degradation has led to emaciated horses in some areas. The public often expresses opposition to horse management, likely out of an innate emotional reaction. However, the emotional co-evolutionary history of humans and horses, coupled with cognitive appraisal of contemporary data particularly as it pertains to horse welfare and populations, offers an opportunity for emotional regulation and de-escalation for this contentious issue to optimize horse welfare.

In memoriam

WEDNESDAY 12:40-13:00

Population status and conservation activities for Asiatic Wild Ass (Equus hemionus) in Mongolia

Badamjav LKHAGVASUREN1, Bayarbaatar BUUVEIBAATAR2

1 Principal Scientist, Institute of General & Experimental Biology, MAS 2 Science Advisor, WCS Mongolia Presenting author: Bayarbaatar Buuveibaatar

We discuss here current population status, population viability as well as estimation efforts and conservation activities for the Asiatic Wild Ass (Equus hemionus) in Mongolia.

The Southern Gobi of Mongolia is an iconic ungulate stronghold that supports the world's largest populations of Asiatic wild ass (or khulan – Equus hemionus). A growing human population, intensifying exploitation of natural resources, and the development of infrastructure in the region place increasing pressure on khulan and its habitat. During 2012–2015, we studied factors influencing the distribution of khulan in the Southern Gobi to better inform management. We built Generalized Linear Mixed Models (GLMMs) to predict the location of suitable habitat for khulan using environmental and human-associated factors. These models were validated using independent telemetry data. The GLMMs suggest that the probability of khulan presence decreased with increasing human influence and increased in areas with intermediate values of elevation and Normalized Difference Vegetation Index. Notably, human-associated factors were more important than environmental variables in explaining the distribution of the khulan. Habitat

models predicted between 45 and 55% of the study area to be suitable for khulan during 2012–2015. The distribution models had good predictive power, as nearly 90% of khulan telemetry locations from separate data set was found within the predicted preferred areas. Our approach quantifies the key drivers of khulan distribution and our findings are useful for policy makers, managers, and industry to plan mitigation measures to reduce the impacts of development.

Wild asses are highly mobile, in the Mongolian Gobi collared animals covered average daily straight line distances of 11.9 km (Kaczensky et al. 2006) or cumulative distances of 21.8 km (Kaczensky unpubl. data) within 24 hours. Wild ass movements do not seem to follow easily predictable patterns (classical migration), but are rather nomadic. The latter is likely the result of the unpredictability in the availability of pasture and water in space and time (Kaczensky et al. 2008, Kaczensky et al. 2011a, von Wehrden et al. 2012). In the past, when wild asses still occurred on mountain steppe and steppe habitats, they made seasonal migrations between summer ranges on the productive steppe habitats in the north to winter ranges in the drier, but less snowy areas in the south (Bannikov 1981).

Asiatic wild asses in Mongolia have huge annual ranges, the size of which increase from east to west, likely as a result of increasing unpredictability of pasture and water availability. In the Dzungarian Gobi annual ranges averaged 5860 km², in the Transaltai Gobi they ranged between 14,695–16,907 km², but in the SE Gobi ranged from 18,186 to 69,988 km2 (Kaczensky et al. 2011a, Batsaikhan et al. 2014).

Modelling for wild and feral equids

WEDNESDAY 14:00-14:20

Does the forage maturation hypothesis predict seasonal movements of equids?

Saeideh ESMAEILI1, Brett R. JESMER2, Shannon E. ALBEKE3, Jerod A. MERKLE1, Briana ABRAHMS4, Lkhagvasuren BADAMJAV5, Buuveibaatar BAYARBAATAR6, Jeffrey L. BECK7, Randall. B. BOONE8, Simon CHAMAILLE-JAMMES9, Paul C. CROSS10, Usukhjargal DORJ11, Ilya R. FISCHHOFF12, Adam T. FORD13, Kate JENKS14, Mahmoud-Reza HEMAMI15, Jacob. D. HENNIG5, Petra KACZENSKY16, Sarah R. B. KING17, John MCEVOY18, Jorg MELZHEIMER19, Thomas. MUELLER20, Dejid NANDINTSETSEG20, Kirk OLSON21, Daniel I. RUBENSTEIN22, Hall SAWYER23, John. D. SCASTA7, Kathryn A. SCHOENECKER17, 24, Melissa SONGER18, Jared A. STABACH18, Seth STAPLETON14, Theresa STRATMANN20, Siva R. SUNDARESAN25, Ito Y. TAKEHIKO26, Jacob R. GOHEEN1.

1 Department of Zoology and Physiology, University of Wyoming, Laramie, USA 2 Max Planck-Yale Center for Biodiversity Movement and Global Change, Department of Ecology and Evolutionary Biology, Yale University, New Haven, CT, USA 3 Wyoming Geographic Information Science Center, University of Wyoming, Laramie WY, USA 4 Southwest Fisheries Science Center, National Oceanic and Atmospheric Administration, Monterey, CA, USA

5 Mammalian Ecology Laboratory, Institute of Biology, Mongolian Academy of Sciences, Ulaanbaatar, Mongolia 6 Mongolia Program, Wildlife Conservation Society, Amar Street-3, “Internom” Bookstore Building, 3rd Floor, Ulaanbaatar, Mongolia 7 Department of Ecosystem Science and Management, University of Wyoming, Laramie WY, USA 8 Department of Ecosystem Science and Sustainability and the Natural Resources Ecology Laboratory, Colorado State University, Fort Collins, CO, USA 9 Researcher, Centre d' Ecologie Fonctionnelle et Evolutive, Montpellier, France. 10 U.S. Geological Survey, Northern Rocky Mountain Science Center, Bozeman, MT, USA 11 The National University of Mongolia and Mongolian Ornithological Society, Ulaanbaatar, Mongolia. P.O. Box 537, Mongolia 12 Cary Institute of Ecosystem Studies, Millbrook, NY, USA 13 Department of Biology, the University of British Columbia, Okanagan BC, Canada 14 Minnesota Zoo, 13000 Zoo Boulevard, Apple Valley, MN 55124, USA 15 Department of Natural Resources, Isfahan University of Technology, Isfahan, Iran 16 Department of Terrestrial Biodiversity, Norwegian Institute for Nature Research, Trondheim, Norway 17 Natural Resource Ecology Laboratory, Warner College of Natural Resources, Colorado State University, Fort Collins, CO, USA 18 Conservation Ecology Center, Smithsonian Conservation Biology Institute, National Zoological Park, Front Royal, VA, USA 19 Department of Evolutionary Ecology, Leibniz Institute for Zoo and Wildlife Research, Berlin, Germany

20 Senckenberg Biodiversity and Climate Research Centre, Frankfurt (Main), Germany 21 Fauna & Flora International, Jupiter House, 4th Floor, Station Road, Cambridge, CB1 2JD, United Kingdom 22 Department of Ecology and Evolutionary Biology, Princeton University, Princeton, NJ, USA 23 Western Ecosystems Technology, Inc., Laramie, WY, USA 24 US Geological Survey, Fort Collins Science Center, Fort Collins, CO, USA 25 Jackson Hole Conservation Alliance, Jackson WY, USA 26 Arid Land Research Center, Tottori University, Tottori, Japan

The Forage Maturation Hypothesis (FMH) predicts that ungulates should optimize their energy intake by selecting high-quality forage of intermediate biomass, which in turn translates to seasonal movements and drives global distributions. Although the FMH was developed to explain aggregation and seasonal movements of ungulates in general, it has been tested primarily on ruminants from temperate regions. Therefore, it remains unclear whether and how the FMH applies more broadly to non-ruminant ungulates. Because of differences in their digestive physiology, we expected that wild and feral equids would prioritize (1) forage biomass over forage quality to meet their nutritional needs; and (2) free water to compensate for their physiological limitations. We used a step selection framework to test for the effect of temporal patterns of forage biomass, forage quality, and distance to free water on movements and resource selection of eleven populations of wild equids, representing six (out of nine) extant species of equids. In contrast to previous works on ruminants, no population of equid selected for high-quality forage (as indexed by instantaneous rate of green-up). Instead, forage biomass and distance to free water consistently drove movements in equid

populations, regardless of geographic locale or body size of species. Although it provides a useful framework for understanding the distribution of ungulates, our work suggests that the FMH should be updated to accommodate alternative predictions specific to the digestive physiology of particular taxa. Migration (in response to predictable patterns of vegetative growth across landscapes) and nomadism (in response to vegetative growth that is temporally unpredictable across landscapes) evolved in ungulates to enhance their exposure to abundant vegetation, high-quality vegetation, or both, but both movement strategies are threatened by human activities. To better protect declining ungulate populations, it is increasingly urgent that we quantify the environmental drivers that underlie long-distance movements throughout the globe.

WEDNESDAY 14:20-14:40

Modeling population growth and management costs for feral horses in the U.S.

L. Stefan EKERNAS1, Kathryn A. SCHOENECKER1, 2

1 United States Geological Survey, Fort Collins, CO, USA 2 Ecosystem Science and Sustainability, Colorado State University, Fort Collins, CO USA

Feral horse populations (Equus ferus caballus) in the United States are growing at approximately 20% annually and present unique management challenges. Feral horses are impacting habitat and negatively affecting other wildlife across the West. Overabundant ungulate populations are typically managed with hunting, but lethal removals for federally-managed horses are prohibited under current U.S. regulations. Other potential management options include temporary female contraceptives, permanent sterilization via surgery, and non-lethal removals in which horses are made available for public adoption and unadopted horses are permanently cared for by the federal government. This is a very costly enterprise. Evaluating potential management options requires information on both horse population outcomes and operational costs. Here, we present results from age- structured matrix population models that incorporate operational costs of different management actions. At current abundances – 72,000 horses on-range at beginning of 2019 – contraceptives are prohibitively difficult and costly to implement on a nationwide or regional basis. Permanent sterilization can stabilize population growth at λ = 1.0, but even the most aggressive sterilization scenarios cannot meaningfully decrease population size of these long-lived animals at management- relevant timescales. Non-lethal removal is the only available tool that can reduce

population size at management-relevant timescales. However, non-lethal removals incur extremely high costs for the duration of horse life expectancy (~25 years) and thus hamper long-term solutions under realistic budget scenarios. The most cost- effective solution is a combination of removals and permanent sterilization to achieve and maintain target population sizes (Appropriate Management Level, “AML,” per the Bureau of Land Management). At AML, permanent sterilization decreases operating costs by ~80% compared to removals alone. Compared to business-as-usual scenarios of using only removals to achieve AML nationwide, incorporating permanent surgical sterilization would save over $1 billion during the next 10 years for the management of feral equids. In addition, our model can be used for conservation planning in rewilding projects to evaluate and guide management actions aimed at maintaining habitat health.

WEDNESDAY 14:40-15:00

Safety and application of telemetry collars on wild and free-roaming equids: A global systematic review

Jeffrey L. BECK1, Jacob D. HENNIG1, J. Derek SCASTA1, Kathryn A. SHOENECKER2, Sarah R. B. KING3

1 Department of Ecosystem Science and Management, University of Wyoming, Laramie, Wyoming, USA 2 United States Geological Survey, Fort Collins Science Center, Fort Collins, Colorado USA 3 Natural Resource Ecology Laboratory, Department of Ecosystem Science and Sustainability, Colorado State University, Fort Collins, Colorado USA

Models using information from individual animals equipped with GPS and VHF transmitters have advanced our understanding of wild and free-roaming vertebrates globally. However, this technology has rarely been used to study free-roaming horses (Equus ferus caballus) and burros (E. asinus) in the United States. Safety concerns about attaching telemetry collars to these equids have been a major impediment for adopting such technology. Such justified concerns arose from a telemetry study in the late-1980s when multiple collared horses suffered a variety of collar-induced injures. In the ensuing quarter of a century, advancements in collar design may have alleviated these issues, yet telemetry studies on free- roaming equids in the United States still face resistance from engaged citizens. On the other hand, telemetry collars have been utilized to study other wild or free- roaming equids around the globe. Thus, we conducted a global systematic review of studies reporting data from telemetered equids. Our goals were to 1) determine the number of equids fitted with VHF or GPS collars, 2) record the number of

individuals that died or suffered injuries while fitted with collars, 3) document the number of complications that were explicitly attributed to the collar, and 4) summarize knowledge gained from use of GPS technology for all equid species. We then conducted a comparative review for 3 wild North American ungulates: elk (Cervus elaphus), mule deer (Odocoileus hemionus), and pronghorn (Antilocapra americana), to assess if equids face an increased chance of mortality or injury while fitted with telemetry collars. Our review revealed 1,089 wild or free-roaming equids have been fitted with VHF or GPS tracking collars across 48 studies from 1979–2017 that contained unique telemetry data. Of these individuals, 87 (8.0%) were reported to have perished, with only 1 (0.09%) mortality explicitly linked to the collar. Comparatively, our abbreviated review found 4,761 elk, mule deer, and pronghorn were equipped with tracking collars in an equal number of studies from 1981–2017. Of these individuals, 1,095 (23.0%) died, though no mortalities were reported to be directly related to the collar. We also synthesized findings from GPS-collar studies for all equid species to better understand how technology has increased understanding of equid ecology and improved management and conservation. Results suggest wild and free-roaming equids do not experience an increased mortality risk compared to other ungulates, but studies have not provided sufficient information to assess relative risk of experiencing collar-related injuries. We recommend explicit reporting and discussion of collar impacts in future publications of all animal species to improve understanding of how telemetry collars impact equids and other ungulates.

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WEDNESDAY 15:00-15:20 – Student

Movement syndromes of feral horses inhabiting arid steppe in Western North America

Jacob D. HENNIG1, J. Derek SCASTA1, Jeffrey L. BECK1

1 Department of Ecosystem Science and Management, University of Wyoming, Laramie, Wyoming USA

Movement patterns of feral horses (Equus ferus caballus) are not well understood, but are important to better conserve habitats that these animals use. Movement syndromes are classified by ecologists as migratory, nomadic, and range resident. Which syndrome a population exhibits is typically correlated to spatiotemporal dynamics of resources within the population’s range. In arid systems, vegetation and water resources are often variable and unpredictable across space and time; therefore, herbivores inhabiting these systems are often nomadic, displaying no defined patterns in their movements. Indeed, some wild equid populations inhabiting arid systems exhibit nomadism, such as the Asiatic wild asses (E. hemionus) in the Gobi Desert; however, others, like sympatric Przewalski’s horses (E. f. przewalskii), display a range resident syndrome with home ranges centered on water sources. Feral horses in the United States predominantly inhabit arid systems but their movement syndromes have yet to be characterized. To understand how these introduced equids move in response to resource dynamics, we placed GPS collars on 20 adult mares in the Adobe Town Herd Management Area (ATHMA) in southern Wyoming, USA from 2017-2019. We used dynamic Brownian Bridge movement models to estimate 95% annual home range sizes and used net squared displacement to characterize the movement syndrome of each horse. We then quantified the spatiotemporal variability and unpredictability of

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vegetation and water resources within the ATHMA to understand if horses moved as expected given resource dynamics. Like Przewalski’s horses in the Gobi Desert, horses in the ATHMA predominantly exhibited a range resident movement syndrome with relatively large annual home ranges (x̅ = 63.2 km2, SD = 37.6). Both vegetation production and surface water availability varied across space; however, surface water varied little across time and its availability was more predictable than vegetation production (CV = 0.01 and 0.43, respectively). Our research suggests that feral horses that inhabit arid systems in the western United States have defined home ranges centered on locations offering predictable sources of water, instead of moving nomadically to track spatiotemporal dynamics of forage production.

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Equid reproductive biology

WEDNESDAY 15:20-15:40

Application of reproductive technologies for conservation of endangered equids

Budhan S. PUKAZHENTHI1

1 Center for Species Survival, Smithsonian Conservation Biology Institute, Front Royal, VA 22630, USA

Many of the world’s wild equids are threatened throughout their natural ranges. While much is known about wild equid ecology, less is known of their fundamental biology, information that is critical to establishing self-sustaining populations in captivity as a hedge against extinction. This presentation will review 1) the importance of integrating animal management and specialized animal handling infrastructure to promote research on wild equids, 2) recent studies on the reproductive biology of three endangered equids - the Przewalski’s horse (Equus ferus przewalskii), Persian onager (Equus hemionus onager), and Grevy’s zebra (Equus grevyii), and 3) successes achieved with semen collection, cryopreservation and artificial insemination. Reproductive assessments via longitudinal monitoring of reproductive hormone metabolites in urine, and rectal ultrasonography in a hydraulic handling device have revealed that both Przewalski’s horse and Persian onagers are seasonally polyestrous (March – September and June – October, respectively) with a mean estrous cycle length of ~25 d. In Przewalski’s horse and Persian onager, follicles grow at ~1.2 mm and ~2.1 mm/day, and attain a preovulatory size of >38 mm and >25 mm, with gestation lasting 320-340 d and 320-350 d, respectively. Ovulation can be induced using exogenous gonadotropins,

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including deslorelin, human chorionic gonadotropin, and recombinant equine luteinizing hormone. With respect to male reproduction, spermic ejaculates can successfully be collected using electroejaculation in all three species with varying degrees of success of obtaining spermic ejaculates (Przewalski’s horse, >90%; Persian onager, >60%; and Grevy’s zebra, <30%). Ejaculates from all three species contain >2 x 109 total sperm (range, 1 – 5 x 109) with >60% total sperm motility. Spermatozoa can be cryopreserved using a skim milk based extender containing methyl- and/or dimethylformamide as cryoprotectant (Przewalski’s horse, Grevy’s zebra) or lactose EDTA extender containing glycerol (Persian onager) with >60% total sperm motility after thawing. A frozen repository of spermatozoa from a large number of Przewalski’s horse and Persian onagers has been established at the Smithsonian Conservation Biology Institute. Pregnancies have been achieved via artificial insemination in the Przewalski’s horse (fresh semen) and Persian onager (fresh-chilled and frozen-thawed sperm). However, additional research is needed to 1) understand the reproductive biology (eg., follicular dynamics, artificial insemination) of female Grevy’s zebra, 2) improve pregnancy rates via artificial insemination using fresh or cryopreserved sperm in the Przewalski’s horse and Persian onager, and 3) optimize semen collection techniques in the Persian onager and Grevy’s zebra. Overall, advances to date demonstrate the feasibility of using assisted reproductive technologies to improve genetic management and conservation of wild equids.

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WEDNESDAY 15:40-16:00

PZP-induced mare infidelity increases male aggression and stress in the feral horse

Cassandra M.V. NUÑEZ1,2, Maggie M. JONES1

1 Iowa State University, United States of America 2 University of Memphis, United States of America

In social species like the feral horse (Equus caballus), changes in individual behavior are likely to impact associated animals. On Shackleford Banks, North Carolina, USA, mares treated with the contraceptive agent porcine zona pellucida (PZP) demonstrate decreased fidelity to their band stallions, engaging in group changing behavior more often than untreated mares; however, the potential impact of these changes on male behavior and physiology remains largely unexplored. We compared the frequency of male-male contests and subsequent cortisol levels for males that experienced varying levels of mare group changing behavior. We found that males experiencing more group changes (either into or out of their bands) fought more often and exhibited elevated cortisol levels. These results are important as they 1) demonstrate the potential for PZP-induced changes in recipients to affect non-recipient behavior and stress physiology and 2) increase our understanding of how contraceptive management may impact non-target animals.

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WEDNESDAY 16:20-16:40

Advances in PZP Immunocontraception in two western wild horse herds

Kayla GRAMS1, Allen T. RUTBERG2, John W. TURNER Jr.3

1 The Science and Conservation Center, Billings, MT, USA 2 Center for Animals and Public Policy, Cummings School of Veterinary Medicine at Tufts University, North Grafton, MA, USA 3 Department of Physiology and Pharmacology, University of Toledo, Toledo, OH, USA

The use of immunocontraceptives on individual female wild horses is well described in published data, notably at Assateague Island National Seashore, Maryland, USA. Several wild horse herds in the western US have also been using the porcine zona pellucida (PZP) immunocontraceptive vaccine as a management tool for years. If immunocontraceptives are to be used effectively as a management tool on western wild horse populations, factors influencing population dynamics in contracepted populations and the challenges of application must be understood. A contraceptive longer acting than the Assateague prototype, which required annual boosters, has been sought for years. Here we present results on population dynamics in herds treated with PZP preparations at two locations, Cedar Mountains Herd Management Area (CM), UT, and Sand Wash Basin HMA (SWB), CO. At both sites we monitored foaling, population size, mortality, group associations, body condition, immigration and emigration. At SWB, 79% of known wild horse mares >2 years old received initial hand-treatments of PZP-22 (PZP emulsion plus controlled-release pellets) during a gather in October 2008. Approximately 65% of SWB mares >2 years old were boosted by dart in autumn 2010 with either PZP-22 or native PZP emulsion. At CM, approximately 52% of known wild horse mares

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>2 years old were gathered and received initial treatments of PZP-22 in December 2008. CM horses were gathered again in February 2012, and approximately 73% of known mares were treated with PZP; untreated CM mares received initial treatments of PZP-22, and previously treated mares received boosters of either PZP-22 or native PZP emulsion. At both SWB and CM, foal and adult mortality were low, and some horses moved across HMA boundaries. Efficacy of initial PZP-22 treatments was comparatively low, but single boosters of either PZP-22 or native PZP emulsion extended contraceptive effectiveness for at least three years at both sites. Foaling rates among all SWB mares >2 years old averaged 46% post- treatment (2010-2013) versus 78% pre-treatment (2009), and population growth averaged 16% per year (2009-2013). At CM, foaling rates among all mares >2 years old averaged 47% post-treatment (2010-2015) versus 62% pre-treatment (2009). Following the 2008 treatments, the CM population grew an average of 18% per year (2009-2012). Because a high proportion of CM mares was treated in 2012 and the boosters were effective and long-lasting, population growth averaged only 12% per year from 2012-2015, with a low of 5.8% in 2013. Our experience suggests that applying current contraceptive tools to large HMA’s can achieve dramatic reductions in population growth if >70% of mares present in the population can be maintained under contraceptive treatment. Given that PZP boosters were effective and long-lasting, planning for multiple treatments will be crucial.

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Genetics & Morphology

WEDNESDAY 16:40-17:00

Facts from feces: the utility of fecal DNA to assist in the conservation and management of wild equids

Sara OYLER-MCCANCE1, Kathryn SCHOENECKER1,2, Sarah R. B. KING2, Jennifer FIKE1, Lars EKERNAS1

1 U.S. Geological Survey, Fort Collins Science Center, Fort Collins, CO, USA 2 Natural Resource Ecology Laboratory, Warner College of Natural Resources, Colorado State University, Fort Collins, CO, USA

For several decades, genetic techniques have been used to address a wide variety of wildlife conservation and management questions. More recently, a push toward non-invasive sampling techniques has led researchers to examine the utility of using DNA from feces to answer such questions. Here, we assess the utility of using DNA extracted from feces of feral horses and burros to address several different management questions. We discuss our research on the degradation of DNA in feral horse feces, how to best sample and store feces, and how long samples can persist in the environment and still be useful for microsatellite amplification. We found that DNA from feral horse feces degrades slowly in the environment, with most samples still amplifying across all 8 loci tested after exposure for 2 months. This is of great benefit to equid researchers as the constraint of finding fresh samples is removed from field methods. Further, we found that the cheapest and easiest sample collection method, paper bags, produced better amplification results than collection of feces stored in vials of ethanol. We also discuss other applications of our equid fecal DNA research including

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estimating population size, determining sex, examining diet, documenting spatial movements, and investigating paternity and relatedness. We highlight here the promise of using fecal DNA to collect a variety of information that is highly relevant to management and conservation of feral horses and burros. We offer that such techniques may be widely applicable to other equid species, particularly those living in arid or semi-arid ecosystems.

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Feral equid issues and rewilding

WEDNESDAY 17:00-17:20

Are free-roaming equids good for western U. S. public lands?

Terry A. MESSMER1

1 Jack H. Berryman Institute, Utah State University, 5230 Old Main Hill, Logan, Utah 84322-5230

The Wild Free-Roaming Horse and Burro Act (WFRHBA) gave the Bureau of Land Management (BLM) and the U. S. Forest Service (USFS) the statutory obligation to manage and protect free-roaming equids (i. e., wild horse and burros; Equus ferus and E. asinus, respectively) in designated management areas. While all free-roaming equids in the U. S. are feral, only the subset designated by the WFRHBA have the legal protections of 'wild horses and burros.' These include descendants of unclaimed, unbranded, free-roaming horses and burros present on BLM and USFS lands in 1971. The intent of the WFRHBA was to ensure healthy populations of feral equids on certain public lands, which are defined as ‘wild horses and burros,’ in ecological balance with other multiple-uses on public lands. These designated public lands are referred to as herd management areas (HMAs). The BLM and USFS have the statutory responsibility for deciding how the HMA should be managed along with the agency’s multiple-use mandate, which includes wildlife, livestock, wilderness, and recreation considerations. Ecological balance was originally set by the BLM in 1978 at 26,715 feral equids inhabiting 12 million hectares in HMAs designated on public land across 10 western US states. As of 1 March 2018, the BLM estimated that there were 81,951 free-roaming equids BLM- administered HMAs. This is more than 3 times the ecological balance. This figure

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does not include an estimated 14,000-18,000 new foals or reflect removals since that date. To achieve herd sizes to sustain ecological balance the BLM would now need to remove more than 60,000 animals from designated HMAs. Overabundant free-roaming equids are impacting the overall health of the US western public rangelands by degrading ecosystem functions and reducing the forage and water available for domestic and native wildlife species. Although the WFRHBA provided the BLM and USFS management options, congressional budget riders and agency policy directives precluded the euthanasia and unrestricted sale of healthy wild horses or burros. The intense emotions of the electorate regarding that tool has indirectly impeded biologically sound management herd reduction practices. The consequences of unchecked annual free-roaming equid growth rates will ultimately impact all western U. S. public land stakeholders. As free-ranging populations increase, the ecological impacts to the arid and unpredictable western U. S. arid rangelands will be exacerbated, and the costs of caring for unadopted horses will continue to compete with other high priority public needs. Most Americans generally have positive perceptions about science and individuals with greater science literacy are more likely to report positive attitudes toward science. This argument has been used to support the science deficit model of communication; those who do not accept the science, simply do not understand it; therefore, all they need are more facts. Whether measured in terms of general educational attainment, science educational attainment, or science literacy, may increase rather than decrease polarization on issues linked to beliefs. These patterns suggest that in the case for the biological management of free-roaming equids, more scientific knowledge may facilitate increased advocacy in individuals defending positions motivated by non-scientific concerns.

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WEDNESDAY 17:20-17:40

Management of feral horse environmental impact in Australia

David BERMAN1

1 University of Southern Queensland, 454 Wellcamp Westbrook Rd, 4350, Westbrook, Australia

Feral horses tend to become overabundant in Australia and there is concern for the negative impact they have on the environment. This paper describes the work conducted to measure the environmental impact of feral horses in various parts of Australia. In 1986 a natural experiment was conducted in central Australia where horses were obviously overabundant. They were walking up to 50 km from drinking water to find food. Many were perishing due to lack of food or water. A valley with both horses and cattle and a valley with horses but no cattle were surveyed. The differences/similarities between the impact of cattle and feral horses on soil, vegetation and wildlife were determined. This work assisted in justifying removal of all horses from Finke Gorge National Park. However, the results in arid Australia do not necessarily apply to more moist areas. Work conducted in the Australian Alps in the late 1980s showed the impact of feral horses on soil and vegetation along the paths that they create. They were shown to influence only a very small proportion of the area. These results may not be relevant today due to increases in the number of feral horses since then. Further work to determine the relationship between feral horse density and impact is necessary in the Australian Alps. In north east New South Wales work was conducted in the Guy Fawkes River National Park. This work involved fencing horses out of exclosures to produce areas with no grazing, as well as measuring changes that occurred after horse density was reduced in some surrounding areas. While this work was a well

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designed manipulative and controlled experiment the results appear to have not influenced management. Research needs to be conducted in conjunction with management and with involvement of community interest groups to ensure results are accepted and used to direct management. There have been a number of recent studies that have attempted to measure the damage caused by feral horses in the Australian Alps. These studies are all correlative and cannot conclusively attribute degradation to feral horses. The impact on the health of ecosystems, habitats and species need to be better measured. The abundance/diversity/population health of native plants or animals need to be measured where feral horses are and compared to measurements in similar sites where feral horses have been removed or their density reduced. It is a common belief in Australia that feral horses will only have negative environmental impact because native plants and animals evolved in the absence of ungulates. However, the fossil evidence indicates that for many millions of years there were mega-fauna, some with large hard toenails, grazing, browsing and trampling native plants and presumably wallowing in mud, compacting soil and creating paths in Australia. Native plants and animals may not only tolerate the impact of feral horses, some at least, may benefit from their presence. Better controlled, manipulative experiments are required to quantify both the negative and positive impact of feral horses.

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WEDNESDAY 17:40-18:00

The rewilding Europe with wild/feral equids and its infrastructural value for equids cognition and social behavioral ecology: The Piornal (Pottokas) and Doñana (Retuertas´ horses) initiatives in Spain

Enrique ALONSO GARCIA1,2, Lucy REES3, Coby BOLGER2, 4

1 Council of State, Kingdom of Spain 2 Animal Studies Research Group, Franklin Institute-University of Alcalá, Spain 3 "Lucy Rees" Wild Horses Association, Spain 4 Horse1, Equine Nutrition Centre

Wild and semi-wild (feral) equids´ populations, as a biodiversity conservation goal, is not acknowledged to exist in Europe (DG Environment, 2018 official statement), although member States can exceptionally establish nature reserves for these equids under EU Agricultural Policy (DG SANTE 2019). Data will be shown on the very few that exist. Almost none is managed in a way that equids´ natural behavior can be expressed and monitored. Reiterative removal of newborn foals for business reasons (sales) or the lack of flexibility in identification requirements (microchipping) are the main problem. The Piornal site of scientific interest, in Cáceres, Extremadura, and the Retuertas site in Doñana National Park are rare places, where the social interactions of entirely wild Pottokas and Retuertas have a solid tradition of at 10 and 30 years. They are also old autochthonous natural breeds. Pottokas have been estimated to be at least 2,000 -year-old (inferred from mitochondrial DNA sequences). Analysis of mitochondrial DNA sequences is still pending in the case of the Retuertas; although, new funding has been provided in order to test their connections to historically and genetically non-mixed mustangs; it will also offer the opportunity to test whether the Retuertas can be traced to the

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4,800- to 3,900-year-old Iberian specimens, as it is the case with Lusitano C (the only group that contains maternal lineages also present in wild Iberian horses from the Early Neolithic period). In the case of Pottokas research has shown why wild horses’ social behavior is widely misunderstood as a dominance hierarchy in which aggression maintains authority. There is solid evidence gathered during ten years that demonstrates that social behavior is not one of dominance but mainly a self- organizing collective movement, based on synchronization and respect of mutual space, avoidance of inbreeding and response to different individuals´ initiatives; the stallions´ behavior been related only to their role as protectors, and bodyguards in exchange for fathering foals, whose growth is the responsibility of the mother. There is no blind following of leaders, but synchrony. Finally, from this learning, partially based in new TIC and innovative satellite imaging techs, some aspects of relevance for applied animal welfare science a) are being translated into very practical ways of undertaking human-horse interactions (nutrition, equestrianism, equine-therapy...), fostering new knowledge on horse cognition and ethology; and b) are bringing science-based changes to the traditional understanding that wild equids are a threat to biodiversity rich ecosystems rather than a potentially ecosystem-rewilding and climate change mitigating keynote species. Current tests on land use and vegetation cover control in Piornal as a Natura 2000 site and holistic pastures management in Doñana National Park are potential success stories.

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Maneless zebra session

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MONDAY 18:00-18:30

Introduction to maneless zebra session

Jan PLUHÁČEK1,2, Jan ROBOVSKÝ3,4, Luděk ČULÍK5, Barbora DOBIÁŠOVÁ6, Tomáš JIRÁSEK7, Lubomír MELICHAR4

1 Ostrava Zoo, Ostrava, Czech Republic 2 Department of Ethology, Institute of Animal Science, Praha - Uhříněves, Czech Republic 3 Department of Zoology, Faculty of Science, University of South Bohemia, České Budějovice, Czech Republic 4 Liberec Zoo, Liberec, Czech Republic 5 Dvůr Králové Zoo, Dvůr Králové, Czech Republic 6 Prague Zoo, Prague, Czech Republic 7 Plzeň Zoo, Plzeň, Czech Republic

Maneless zebra (Equus quagga borensis) represents the most northern subspecies of plains zebra. Originally described by Lönnberg (1921), it was not recognised as a valid subspecies in many reviews of plains zebra taxonomy during 20th century, but predominantly without any comparisons or reasons. Most of them did not consider the 75% rule for subspecies recognition. Applying this rule would means that if adults of borensis exhibit reduced mane in 100%, adults of others of other subspecies (e.g. E. q. boehmi) should exhibit mane reduction in less than 75% of adults (which is the case without any doubts).

On the other hand, several other studies in 1960s emphasized the uniqueness of the taxa and provided good arguments for supporting its subspecific status. In addition, some maneless specimens were mislabelled by invalid names as E. q. jallae and E. q. cuninghamei. In 1959 new synonym for maneless zebra was suggested: E. b.

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isabella. In 21st century, the situation changed and most studies recognized maneless zebra as a valid subspecies or even the most distinguished one. These studies were based on differences in morphology, behavioural ecology, as well as genomic data.

Maneless zebra differs from other subspecies (and all other equid species) by the fixed absence of mane in adults of both sexes (females have mane developed but reduced), larger body size than E. q. boehmi, the skull which is more sexually dimorphic than in other subspecies of plains zebra with relatively short diastema. Further, unlike other subspecies of plains zebra, maneless zebra is absent the infundibulum on lower incisors. Moreover, lower occurrence of infanticide was recorded in this subspecies, and genomic data recognised it as the most distinct population across plains zebra.

Despite all these facts, this subspecies was neglected by many institutions, therefore no conservation action has ever been made. Formerly, the subspecies was distributed in large savannah areas in South Sudan, North Kenya and North Uganda. Nowadays, they survive in Kidepo National Park, North Uganda,. Recently (July 2019), some of us observed two herds counting 15 individuals of maneless zebras in Pian Upe Wildlife Reserve, Northeast Uganda. Both these unique and autochtonous populations do not include more than 250 individuals (Robert Aruho and Samuel Loware, pers. comm. 2019). Thus, conservation action and establishment of another back-up population is urgently needed.

All other maneless zebras outside of range countries came from Czechoslovak imports in 1969-1975 led by former director of Dvůr Králové Zoo Josef Vágner. Unfortunately, a lot of animals (either imported or born in zoos) were sent to institutions where they were mixed with other plains zebra and lost. Thus, the pure population remained stable in size of around 50 individuals. Since 1995 monitoring of the current European population started. The population came from 10 founders 119

originating in Uganda and more than 80 % of pedigree is known. However, at the beginning of the last decade population declined dramatically as several traditional holding institutions stopped keeping the subspecies in their collections. Currently, the European population (being the only captive population) consists of 39 individuals kept in 12 European zoos, most of them located in three zoos in Czechia (Dvůr Králové, Liberec, Prague). However, the foaling rate is quite low with only 3 foals reported during last 12 months.

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Poster contributions

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Behavioral Ecology

POSTER 1

Courtship patterns in semiferal horses (Equus caballus)

Martina KOMÁRKOVÁ1, Adéla BOUŠKOVÁ2, Francisco CEACERO1

1 Faculty of Tropical AgriSciences, Czech University of Life Sciences Prague, Czech Republic 2 Faculty of Zoology, Charles University in Prague, Czech Republic

Courtship in horses (Equus caballus), polygynous mammals with harem type of social organization, naturally involves wide spectrum of behaviour shown within several days of estrus. Contrary to that, under domestic conditions courtship almost does not occur, although it may have a major impact on the reproductive success of the couple.

Our study addressed mostly the individual specifity of the courtship in semi-feral and domestic horses mating freely, specifically Exmoor pony and Merens horse breed. A total of 45 individuals in five harem herds were observed for 240 hours from February to June 2018. All the courtship and mating events were recorded; we measured the length of interaction, time of the first mounting and mating and specific behaviours of the male and female separately: initiator and terminator of the interaction, vocalisation, urination, flehmen, clitorial winking, tail lifting, following, licking, sniffing, biting, stamping, nose-nose contact, threats and attacks, reaction to mounting, erection, ejaculation, way of termination). We expected highly repeatable patterns in courtship behaviour within particular couples, but the results showed the opposite: a high variability was observed for most couples (ICC ranging from 0.235 to 0.832). 5 couples showed low

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repeatability, 8 showed fair repeatability, 3 showed good repeatability and any showed excellent repeatability. Moreover, the observed repeatability did not correlate with individual characteristics of the mare or the stallion, i.e. age, parity, experience or postpartum estrus. Similarly, these factors did not affect the length of the interaction. The probability of mounting was higher when the stallion started the interaction (15.3% vs. 0% when the mare started; χ2=3.807, p=0.051). The typical courtship initiated by the stallion included erection, sniffing and short naso- nasal contact, and mare reacted with low occurrence of stamping and neighing and also indifference. On the contrary, when mare was the initiator, no indiference was perforemed, but more neighing and stamping and naso-nasal contact, which, however, never finished on mating. About the terminator of the interaction, it was mostly the mare when mounting happened (25% vs. 0% terminated by the stallion; χ2=11.957, p=0.003). Mares also used to stop the interaction when threatening towards the stallion was observed (84.2 %, χ2 = 18.492, p <0.001). Our results suggest that the courtship should be allowed in domestic breeding, so horses can naturally display a variable range of behaviours. It is therefore necessary to allow the couple to have more sexual interactions, precopulation contact, and typical courtship behavioural elements, and to provide the mare a sufficient freedom of movement. These results may be also beneficial for the breeding programs of other endangered equids (Przewalski‘s horse, zebras).

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POSTER 2 – Student

Multi-level structure in feral horse society: Evidence from the aerial observation from drones

Tamao MAEDA1, Sakiho OCHI1, Monamie RINGHOFER2, Satoshi HIRATA1, Shinya YAMAMOTO2

1 Wildlife Research Center, Kyoto University, Kyoto, Japan 2 Institute for Advanced Study, Kyoto University, Kyoto, Japan

Some mammalian species of different taxonomic groups, including humans, have developed social structures with nested levels of organization. This multilevel society is one of the most complex social systems in animals. Most of the previous studies provide only descriptive data, but qualitative definition and description of multi-level society is essential to understand their function and evolutional processes. Equine groups are one of the taxa that have nested social structure, and there are some studies on several species such as plains zebras and Przewalski's horses, but not on domestic horses (Equus caballus). In this study, we aimed to show the evidence of multi-level society and define unit groups using positional data of individuals. We took aerial photos of feral horse herd, which is a large group consisting of 123 adults and subadults, in Serra D’Arga Portugal in 30 minutes interval using drones, identified all the individuals and collected their position data. The distribution of the inter-individual distances was binomial, which suggest the presence of unit groups in the herd because first peak represents the inter-individual distances inside the unit groups, and the second one is those of between the unit groups. Their home range were largely overlapped and the area of convex hull of these groups were significantly smaller than each home range, which suggests harems and bachelors aggregates to form a herd. Those results

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overall suggest the multi-level structure of horse society. We also found that we could use the valley of the inter-individual distances as the threshold of intra-unit association, and using this method we defined 21 harems and 2 bachelor groups as unit groups. Moreover, this herd had a structure that large harems were likely to be in the center, while bachelors were in the peripheral zone, and small harems were located somewhere between that. The presence of this stable spatial pattern strongly indicates the multilevel structures of feral horse society.

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POSTER 3

Insights into social behaviour and adaptation to the environs of free-ranging Asiatic wild asses by photographic documents

Gertrud NEUMANN-DENZAU1, Helmut DENZAU1

1 Panker, Germany

Walking along with Asiatic wild asses in their natural habitats we were able to observe and document many aspects of their behaviour since decades. The question 'How do they manage to survive?' has accompanied us on all our travels. Seasonal challenges, predation, human-wildlife conflicts (including poaching) will be dealt with in this presentation. First of all one has to be able to identify individual animals to understand the course of events. So we'll explain our major identification criteria for these equids (like size and shape of chestnuts, extend of white colouration of jaw angles, and others) with pictures for comparison. In several photographic sequences we’ll show aggressive as well as friendly and mutual interactions. When chased the pursuer will be kept away by kicked out hind legs. But severe fights among stallions are fought in close combat by biting, and might take a bad turn, if no one gives up. The full cycle of life from defence of territories, mating, foaling, and passing away will be depicted. Just once in more than 30 years we were lucky to witness a birth. That was in India's Little Runn of Kutch. Grooming and playing are rarely seen among wild asses, and are obviously just occasionally performed between certain reconciled individuals. It's not an overall behaviour. Described as a nightmare when it occurs in a zoo, we feared for the worst when a sleeping young foal got grabbed by a territorial stallion. In this case we witnessed a Happy End. We can prove with pictures, that the invasive thorn bush Prosopis juliflora, devilised as a pest for invading semi-arid and arid landscapes, has not only negative impacts on the native

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vegetation, but quite a relevance as food source for the wild asses, which feed on its leaves and seed pods.

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POSTER 4 – Student

How vocalizations affect social behavior in free ranging horse groups (Equus caballus)

Sakiho OCHI1, Satoshi HIRATA1

1 Wildlife Research Center, Kyoto University, Kyoto, Japan

For a better understanding of animal behavior or society, various studies on animal vocalizations have been conducted, and that is also important for animal welfare from the perspective of potentially monitoring their emotional states or social relationships. It is known that horses (Equus caballus) frequently communicate by using vocalizations, and previous studies show that vocalization has a big role in forming horse groups in some situations; for example, adult mares make calls when they are separated from other group members or their offspring. However, few studies discuss how those vocalizations are used for maintaining their social relationships thoroughout their daily life as a member of a group. In the present study, we aim to understand the function of vocalization in free ranging horse groups, and also aim to investigate whether they also use their vocalization to communicate with human caretakers. To achieve those goals, we recorded horses’ vocalizations from 58 individuals from five groups, in four contexts: 1) separation from a familiar individual(s), 2) demanding food from humans, 3) aggression to another individual, 4) free ranging. We attached small microphones to all individuals’ heads in the target group to identify which individual made the vocalization, and recorded their vocalizations for ten hours per day. We also observed their social behavior to evaluated their social ranking by applying an average dominance index. From those data, we investigate the spectral difference

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between vocalizations, by using the Hidden Markov Model. In this presentation, we will present the preliminary results of those analysis.

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POSTER 5

Movement patterns and leadership in a multilevel social group of Przewalski horses Katalin OZOGÁNY1, Viola KEREKES1,2, Máté NAGY3,4,5, Zoltán BARTA6 1 Department of Evolutionary Zoology and Human Biology, University of Debrecen, Debrecen, Hungary 2 Hortobagy National Park Directorate, Hungary 3 Department of Collective Behaviour, Max Planck Institute for Ornithology, Konstanz, Germany 4 Department of Biology, University of Konstanz, Konstanz, Germany 5 MTA-ELTE Statistical and Biological Physics Research Group, Hungarian Academy of Sciences, Budapest, Hungary 6 MTA-DE Behavioural Ecology Research Group, Department of Evolutionary Zoology and Human Biology, University of Debrecen, Debrecen, Hungary

In a moving animal group collective movement patterns arise as a result of the interactions between the individuals, and hence the movement patterns are often modulated by social relations and reflect the structure of the society the animals are living in. Social relations of individuals may also influence the dynamics of leading during movements. This is even more interesting in a complex multilevel society, where the motion of the large group, as well as its smaller sub-groups can be studied. To investigate the relationship between social structure, movement patterns and leadership, we observed the collective motion of free-ranging Przewalski horses in Hortobágy National Park, Hungary. These horses are especially suitable to such a study because in this reserve they live in a complex multilevel society: harems of various size and bachelor groups unite into a large herd. In this study we used novel methods, to obtain continuous and simultaneous

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position data at high frequency for a large number of wild animals in natural environment. Our observational method is based on aerial videos of two drones, one provides high resolution motion data for all animals in the herd, while the other ensures individual recognition. In this way, we collected simultaneous trajectory data on the daily movements of around 250 individuals of known identity. Using these trajectory data, we are studying the motion patterns and leadership dynamics inside the harems and the whole herd. Since all the horses are recognized and their life- and harem history is recorded regularly for the past 20 years, motion data can be related to many individual and group characteristics. High-accuracy tracking of many individuals became possible only in the recent years, therefore studies regarding the collective movements of large groups are scarce yet. As one of the first studies in a multilevel social group, our results on collective movements and leadership will contribute significantly to the understanding of collective behaviour.

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POSTER 6 – Student

Does size matter? Examining the possible mechanisms and adaptive significance of single and multi-male groups in horse societies

Pandora PINTO1, Satoshi HIRATA1

1 Wildlife Research Center, Kyoto University, Japan

Horse groups with single and multiple stallions occur simultaneously throughout the world in different feral horses’ populations. However, little is known on why such groups with more than one male exist, considering that stallions naturally fight to monopolize the females. Body size is often an important determinant of male fighting ability and reproductive success, thus it’s possible that males in single-stallion groups are larger than males in multi-stallion or bachelor groups, making them better competitors or more dominant. We observed feral horses, more specifically the Garrano horses that live in Northern Portugal, and examined the relationship between body size, sex, group composition (single-stallion, multi- stallion or bachelor group) and number of females in the group, by measuring individual body length and height with a non-invasive laser distance meter. Preliminary results showed no significant differences in body length and height for all parameters assessed, suggesting that other factors may be more important to explain the existence of single and multi-stallion groups in horse societies. We plan to further test the reliability of remote body size measurements in the field, through comparison with known body size of domestic individuals.

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POSTER 7

Why are zebras striped

Daniel RUBENSTEIN1

1 Princeton University, Department of Ecology and Evolutionary Biology, Princeton, 8544, United States

Few diurnal mammals are so boldly colored as zebras. Their black and white stripes are their signatures. Many hypotheses have been presented to explain why they have evolved. The major ones are: 1) that they facilitate social cohesion; 2) that they dazzle and confuse large attacking predators; 3) that they deter flies from landing and biting; and 4) that they enhance temperature regulation. Evidence for all but avoiding biting flies is scanty and even then the mechanism responsible for deterring biting flies is still unknown. The poster will examine theory and evidence for each of these hypotheses and suggest that multiple explanations are possible, with some being flip sides of the same coin.

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POSTER 8

Herding mechanisms to maintain the cohesion of a harem group: two interaction phases during herding

Monamie RINGHOFER1*, Clark Kendrick C. GO2, Sota INOUE3, Renata S. MENDONÇA1, Satoshi HIRATA3, Takatomi KUBO2, Kazushi IKEDA2, Shinya YAMAMOTO1

1 Institute for Advanced Study, Kyoto University, Kyoto, Japan 2 Nara Institute of Science and Technology, Nara, Japan 3 Wildlife Research Center, Kyoto University, Kyoto, Japan

In animal groups, individual interactions achieve coordinated movements to maintain cohesion. In horse harem groups, herding is a behaviour that the males chase the females from behind; it is considered to assist with group cohesiveness. However, the mechanisms by which the individuals move to maintain group cohesion is unknown. We applied new and non-invasive methods of drone filming and video tracking to observe horse movements in the field with high temporal and spatial resolution. We tracked all group members and drew trajectories during herding events. Females did not start to move simultaneously, but instead started to move in sequence. We analysed the movements of females during herding events and found two phases of interactions based on their timing of the movement initiation. The females that moved first were those nearest the herding male and they started to move when the herding male approached at a distance of 7.96±0.95 BL (mean±SE, n = 10). Movement initiation of the second and subsequent females was determined by distance from the nearest moving female, rather than by distance from the herding male (GLMM with binomial error structure with logit- link, conducted with likelihood ratio test: Z = 1.000, P = 0.313 for the distance

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from the herding male, and Z = －2.464, P = 0.003 for the distance from the moving female, n = 37). These interactions are unique among animal group movements and may represent a mechanism for herding to maintain group cohesion. This might be due to the long-term stable relationships within horse harem group, and the strong social bonds between females. This study showed the combination of drone filming and video tracking is a useful method for analysing the movements of animals simultaneously in high resolution.

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POSTER 9 – Student

Paternal role of a harem male: Analysis of stallion-foal interactions

Kateřina ŠANDLOVÁ1, Martina KOMÁRKOVÁ2, Francisco CEACERO2

1 Department of Zoology, Faculty of Science, University of South Bohemia in České Budějovice, Branišovská 1760, 370 05, České Budějovice, Czech Republic 2 Department of Animal Science and Food Processing, Faculty of Tropical AgriSciences, Czech University of Life Sciences, Kamýcká 129, 165 00, Prague 6 - Suchdol, Czech Republic

Social behaviour of horses (Equus caballus) has been studied extensively and thoroughly. However, the relationship between stallions and their offspring remains still unclear. The aim of this study was to describe and analyse the interactions between harem male and foals. Two herds of Exmoor ponies living under semi-feral conditions were studied each consisted of a stallion, 14 mares and their offspring. The observation lasted for 6 months for both harems, totally 357 hours of observation collected. During the sessions, aggressive, friendly, play and snapping behaviour of all group members was recorded by “ad libitum sampling” method with the special focus on interactions between stallion and foals. According to our results, the stallion is the most frequent adult recipient of foal’s interactions (p < 0.001 for friendly behaviour and snapping, p = 0.01 for play). Moreover, the male is also the most tolerant adult individual compared to the behaviour of the foal’s mothers and other mares and as the only adult, the stallion occasionally plays with his offspring. In this study the common friendly contact between stallion and foals was proven. The benefits from this relationship are questions for our next research. Yet, the stallion’s presence could be beneficial for physical and psychical development of the foals, especially the colts, or stallion

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could act as a behavioural model for appropriate social development of the young. On the other hand, in the common breeding practise, fertile males are usually kept separately, and the foals live without contact with the stallion.

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Conservation

POSTER 10

Does livestock leave room for wildlife movements and distribution in multi- use landscapes?

Saeideh ESMAEILI1*, Mahmoud-Reza HEMAMI2, Brett R. JESMER3, Petra KACZENSKY4, Chris WALZER5, Jacob R. GOHEEN1

1 Department of Zoology and Physiology, University of Wyoming, Laramie, WY, USA 2 Department of Natural Resources, Isfahan University of Technology, Isfahan, Iran 3 Department of Ecology and Evolutionary Biology, Yale University, New Haven, CT, USA 4 Department of Terrestrial Ecology, Norwegian Institute for Nature Research, Trondheim, Norway 5 Wildlife Conservation Society, Bronx, NY, USA

Human activities can alter when and where long-distance movements (i.e. migration and nomadism) occur, and may even threaten the persistence of migration routes. This is exacerbated when humans and their livestock track the same shifting distributions of resources as wildlife (for example, by tracking rainfall and subsequent vegetation growth). To alleviate such conflicts in multi-use landscapes, it therefore is important to quantify human-wildlife interactions. The onager (Equus hemionus onager), otherwise known as the Persian wild ass, is an endangered subspecies of Asiatic wild ass (which is itself listed as Threatened by the IUCN). Currently, onagers are restricted almost entirely to Qatrouiyeh National

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Park (QNP; a 310 km2 protected area in which livestock grazing is illegal) and the surrounding Bahram-e-Goor Protected Area (BPA; a 4090 km2 multi-use landscape) in south-central Iran, in which an estimated 750 individuals occur. Since 2008, national protections have led to increased populations within QNP, but not the broader BPA. Because of increasing livestock production throughout BPA, onagers have been “compressed” into QNP. We investigated (1) how onager movements were influenced by humans and their livestock; and (2) whether the distribution of onagers followed an “ideal free” strategy, through which individuals within QNP and within BPA equalized exposure to high-quality foods. To test the hypothesis that onagers were displaced by livestock, we calculated home range overlap between onagers and livestock. To test if onagers were distributed in an ideal free manner, we compared levels of crude protein and crude fiber in fresh fecal samples (n=320) within QNP and outside QNP but within BPA. Annual home ranges of onagers (n=18 animal-years) were larger than the whole of QNP, and human-onager conflicts were rooted in areas where onager home ranges overlapped with livestock movements and farms within BPA. Over two-thirds of telemetered onagers were nomadic, and the remaining individuals were identified as residents; onager movements were confined to areas in which use by livestock was minimal. The percent of crude protein in onager diets within QNP and BPA did not differ significantly, despite densities in QNP that were nearly eight times those in BPA. However, onagers within BPA exhibited significantly lower levels of crude fiber in their diets, likely indicating the influence of high-quality subsidies (crops) from neighboring farmlands. In sum, the distribution of onagers conformed to an “ideal free strategy”, although livestock production and other human activities have likely reduced carrying capacity and population growth within BPA.

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POSTER 11

EEP Przewalski horse – overview of working procedures and latest developments

Benjamin IBLER1

1 AG Zoologischer Garten Köln, Riehler Straße 173, D-50735 Köln, Germany

European ex-situ species programme (EEP) for the Przewalski horse (Equus przewalski) is managed at Cologne Zoo since 1985 in addition to the international studbook at Praha Zoo. In 2018 the author took over the co-ordination. During that period a lot of achievements have been made. The Przewalski horse is one of the stars of combined ex -situ and in-situ conservation. EEP and international studbook are the central point to bundle as many activities as possible. Beside the traditional zoo population, a lot of semireservates in European countries have been established. Such semireservates were developed for the protection of open- landscapes with their native fauna and flora with the Przewalski as a flagship species. The greatest semireservate is Hortobagy in Hungary to supported financially by Cologne Zoo since its foundation in 1997. The transports to Mongolia are run by Praha Zoo, transports to China are in the planning phase as well. In the practical work many describing analyses of the population are necessary: structures, age-distributions and analyses of relationships. This are the basis for the detailed working procedures and instruments of the EEP. Here a short introduction over instruments, interdisciplinary possibilities and many activities as a part of applied conservation is given making the Przewalski horse to a story of success of nature conservation.

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POSTER 12 – Student

Space use patterns of Konik horses in conservation grazing

Lilla LOVÁSZ1, Valentin AMRHEIN1

1 University of Basel, Department of Environmental Sciences, Zoology

Robust horse races, such as the Konik, are increasingly introduced in nature protection areas to counteract succession and enhance biodiversity. Both these duties are largely influenced by movement patterns and thus space utilization of horses. We investigated space use of a group of Koniks recently introduced in a 32 ha alluvial area in a French nature reserve, the Petite Camargue Alsacienne. Horses graze the area year-round, without supplementary feeding and together with a herd of Highland cattle. The movements of Konik horses and Highland cattle have been followed with GPS collars. We present preliminary results on seasonal changes in utilization distribution of horses and its correlation to the movements of the cattle herd.

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POSTER 13 – Student

Habitat overlap of African wild ass (Equus africanus Fitzinger, 1857) with livestock’s in Afar Region, Ethiopia

Redwan MOHAMMED1*, Fanuel KEBEDE2, Patricia D. MOEHLMAN3, Afework BEKELE4, Nicholas E. YOUNG5, Paul H. EVANGELISTA6

1 Faculty of Natural Sciences, Department of Biology, Samara University, P. O. Box 132, Samara Afar, Ethiopia 2 Ethiopian Wildlife Conservation Authority, P.O. Box 386, Addis Ababa, Ethiopia 3 IUCN/SSC Equid Specialist Group, P.O. Box 2031, Arusha, Tanzania 4 Department of Biology, Addis Ababa University, P.O. Box 1176, Addis Ababa, Ethiopia 5 Natural Resource Ecology Laboratory, Colorado State University, 1499 Campus Delivery, Fort Collins, Colorado 80523 USA 6 Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, CO, 80523-1499, U.S.A

Extensive habitat encroachment by humans and their livestock’s for water and forage in Afar depression specially in highly water scarce season is causing serious problems on the existence of African wild ass (Equus africanus, Fitzinger, 1857) which is the most Critically Endangered equid in the world as listed in IUCN’s red list category and other wildlife in Afar region, Ethiopia. Wildlife crises are a result of increasing human activities, particularly competition for space and resources. As a result, wildlife populations are becoming fragmented and restricted to islands of natural habitat. Conflicts between human and wildlife are emerging as a major conservation challenge worldwide. A major cause for human impact on wildlife is increasing human population encroachment on accessible wildlife habitats.

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Wildlife conservationists need to know the level of overlap between African wild ass and livestock’s as well as other wildlife in the area to take a proper measure on the existing problems to save this precious wild equid from extinction. The techniques used in this investigation are Remote Sensing and NDVI to know the level of overlap and extent of encroachment. Seasonal temperature levels, distance from water, distance from human settlement and elevation were used as predictor variables. The area has bimodal seasonal condition showing very dry and relative wet seasons. Continuous field surveys were carried out twice in dry and twice in wet seasons for consecutive two years. In addition to habitat overlap, the other critical issue is poaching for food and traditional medicinal purpose. The results of this investigation will be very important on showing the extent of overlap between African wild ass and livestock’s as well as other wildlife and forward remedial conservation strategies for wildlife managers on the conservation of African wild ass and other wildlife in the area.

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Population dynamics, viability, estimation

POSTER 14 – Student

Feral horse population ecology in a multi-prey, multi-predator system

Paul BOYCE1, Philip MCLOUGHLIN1

1 University of Saskatchewan, Department of Biology, Saskatchewan, Canada

The Alberta foothills represent a multi-prey, multi-predator system situated east of the Rocky Mountains. A population of feral horses has survived in this region since the l9th century, amongst native ungulates such as elk and moose, and a suite of large predators including wolves, grizzly bears, black bears, and mountain lions. Annual aerial counts indicate that the population has been rapidly expanding in recent years, likely in connection with logging activity, leading to potential impacts on both native and domestic ungulates including free-ranging cattle. This system presents a unique research opportunity into the ecology and potential management of feral horses in a complex predator-prey system. In particular, we aim to understand the role of the horse as both a direct competitor of alternate prey, and as an indirect competitor through apparent competition as mediated by predators. Dietary studies of predators in the foothills have shown that feral horses are preyed upon, though nothing is known about survival and reproduction in this population, nor the impact of the large addition of biomass to the landscape for both competing ungulates and predators. We are using both GPS radio collars and a trail camera grid system to assess density, survival, and reproduction in this population of feral horses, while also assessing concurrent densities of other prey species and predators. Using a combination of capture-mark-recapture tehniques and resource

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selection functions, we aim to assess the impacts of predator populations on feral horses and the interactions between horses and other prey populations.

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POSTER 15

Where donkeys run free? The case of Equus asinus in Jericoacoara National Park

Cara CLANCY1, Mariana GAMIERO2

1 The Donkey Sanctuary (UK) 2 University of Sao Paulo (Brazil)

For over 400 years, donkeys (Equus asinus) have thrived in a remote sand dune ecosystem in northeast Brazil, known today as Jericoacoara National Park. Initially brought over by Portuguese conquistadors in the mid-1500s to support Brazilwood extraction, sugar production and salt mining, the northeastern donkey has played a central role in the development of Brazil. In Jericoacoara, they have maintained a dynamic population over the years, despite regional development and a variable climate, and are the only donkeys currently known to inhabit this kind of ecosystem in Brazil. In 1984, the area (8,416 hectares) was declared an Environmental Protection Area and became a National Park in 2002, famed for its unique ecology, migrating birds and rare wildlife. It is also a top tourist hotspot in Brazil, with approximately half a million visitors per year, attracted by its beaches and water sports, as well as the herds of free-roaming donkeys. However, the future of these semi-wild animals is uncertain, as is the future of donkeys all over Brazil as thousands are slaughtered every year for their skins (The Donkey Sanctuary, 2017). Under Brazilian law, domestic animals are not permitted into national parks and so discussions are underway to remove the donkeys from the park. Yet many local people argue that they are part of the landscape, both culturally and ecologically. The population at Jericoacoara is unique: self- sustaining, with no supplemental feeding; experiencing similar selective pressures

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to those of ‘wild’ equids. This paper examines the complexity of the domestic/wild interface through the case of donkeys in Jericoacoara National Park, including the environmental politics at play and the tensions between conservation, welfare and cultural heritage.

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POSTER 16

Feral horses’ (Equus ferus caballus) behavior toward dying and dead conspecifics

Renata S. MENDONÇA 1,2, Monamie RINGHOFER1, Pandora PINTO3, Sota INOUE3, Satoshi HIRATA3

1 Institute for Advanced Study, Kyoto University, Kyoto, Japan 2 Centre for Functional Ecology - Science for People & the Planet, Department of Life Sciences, University of Coimbra, Coimbra, Portugal 3 Wildlife Research Center, Kyoto University, Kyoto, Japan

In the rapidly expanding field of comparative thanatology, reports from a wide range of taxa suggest that some aspects of a concept of death may be shared by many non-human species. In horses, there are only a few anecdotal reports on behaviors toward dead conspecifics, mostly concerning domestic individuals. Here, we describe the case of a 2-month-old, free-ranging male foal that died around 12 h after being found severely injured due to a presumed wolf attack, focusing on other individuals’ reactions to the dying foal. We also placed camera traps near horse carcasses to investigate reactions by other horses. Although largely opportunistic, our observations may help clarify 1) mother-foal attachment and other factors affecting the mother’s behavior, 2) more generally, horses’ perception of and responses toward dying kin and non-kin, and 3) responses to dead conspecifics. After the foal fell and was unable to move, only his mother remained nearby, despite attempts by the male to herd her back to the group. However, she eventually abandoned the foal and re-joined her group after being confronted and harassed by two bachelor males. The mother’s final abandonment of her dying foal seems likely to have arisen from a combination of social and environmental

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factors. Non-kin individuals – especially two young adult females – also showed affiliative behaviors toward the injured foal, even though adults usually behave agonistically toward extra-group foals. Our observations on reactions toward horses’ carcasses, using camera traps and direct observation of behavior showed that horses seem to avoid dead conspecifics. These events suggest that horses react to injured individuals with whom they share close bonds (mother-foal) or even looser bonds (e.g., non-kin adults-foal), and they pay more attention to individuals when they are still alive than to dead ones. Recording individual reactions to dead and dying conspecifics in naturalistic settings will enhance our knowledge about death-related behaviors in horses, allowing comparisons with other species that have been more thoroughly studied, to understand the evolutionary basis of these behaviors.

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POSTER 17

Population dynamic and demography of reintroducing Przewalski’s horse Equus ferus przewalskii Groves, 1986 in Hustai National Park, Mongolia

Usukhjargal DORJ1, Enkhbayar JAGDAG1 and Dashpurev TSERENDELEG1

1 Hustai National Park, Mongolia, Building number: 40/2, II khoroo, II khoroolol, Bayangol district, 16051, Ulaanbaatar, Mongolia

The Przewalski’s horse Equus ferus przewalskii Groves, 1986 (PH) is considered to be the last and only remaining wild horse species in the world although recent genetic research suggests that the PH may originate from early domesticated horses (Gaunitz et al., 2018). PH went extinct in the wild by 1969 (Dovchin, 1969) and was reintroduced back into the wild in Mongolia, China and Kazakhstan in the 1990s (Kaczensky et al., 2016). From 1992 to 2000, 84 PH were re-introduced to Hustai National Park (HNP) in Mongolia. Since then there have been 1032 foals born in HNP. The average survival rate of foals to their first year is 53.5%, second year is 80%, and survival to adulthood is 90-100%. Females start to give birth at age 3. Average birth rate is 12.5% at age 3; 55.4% at age 4; 68.1% at age 5-15 and 42.6% at age above 16. The record for lifespan is 28 years held by a mare, “Altan”, born in De Ooij Nature Reserve in the Netherlands in 1990 and transported to HNP in 1992 who died in 2018. Even oldest female gave birth record is belonged to her. She gave birth normally when she was 24 years old. Over 1032 births, twin birth or twin fetus have never been recorded. The growth rate of the PH population during the period of active transportation from 1992 to 2000 was 32% and during the period without new transportation from 2002 up to 2018 was 7.2%. Since the first reintroductions in 1992, there have been annual population declines in only three years; 2001, 2010 and 2017 and all other years increased. The population declines

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of -9.9% in 2001 and -10.1% in 2010 were due to natural disasters known as Dzud (severe winters). The greatest loss was recorded in 2017 (-13.7%), suggested density dependence population regulation system. PH movement behaviour in HNP has been largely sedentary with only limited use of areas outside the centre of the park. In recent years home range of PH are shrinking density is increasing. Causes of mortality differed by age and sex. The predominant cause of mortality in foals and yearlings was predation by grey wolves. Recently, deaths of yearlings from blood parasites have tended to increase. Stallions usually died of bad condition and injuries sustained when fighting for mares. Parturition problems are the most common cause of death among mares. At present there are 335 PH in HNP with 34 breeding harems and more than 80 bachelors competing for access to the mares. Six of the original transported PH (1.8%) remain in the population and the rest were born in HNP (98.2%). Adult males make up 37.7% of the population, females 37.3%, subadult males 9.7%, subadult females 10.4% and foals 13.0%.

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POSTER 18

Population dynamics of a reintroduced population of Przewalski’s horse Equus ferus przewalskii in Khomiin Tal, Khar Us Nuur National Park, Mongolia, compared to its source population, other reintroductions and feral horses

Florian DROUARD1,2, Byamba MUNKHTUYA2 and Claudia FEH2,3

1 Khomiin Taliin Takhiin NGO, Ulan Bataar, Mongolia 2 Association pour le cheval de Przewalski: TAKH, Le Villaret, France 3 Tour du Valat Research Institute, Le Sambuc, 13200 Arles, France

Globally, large animals are the most endangered species, mammals being of particular concern. To identify factors that influence fecundity and mortality in these populations is therefore paramount. In free living populations, direct evidence is often difficult to collect due to observational limitations. The endangered Przewalski’s horse (Equus ferus przewalskii) may have survived for centuries due to the cultural attitude of mostly Mongolian herders, and has come back from the brink of extinction thanks to zoos, followed by international reintroduction efforts. Here we compare population dynamics of one of the 3 reintroduction projects in Mongolia, in Khomiin Tal, Khar Us Nuur National Park, with its source population in Le Villaret, France, where predators and infectious disease transmission through domestic horses were absent. We continue the comparison to data available from other reintroductions as well as feral horses, over a roughly 15 years’ time-frame.

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POSTER 19

Assay validations for a wild equid population: measuring physiological responses to environmental stress in the Carneddau Welsh Mountain Pony

Danielle HINCHCLIFFE1,2, Susanne SHULTZ1

1 School of Earth and Environmental Sciences, University of Manchester, UK 2 Chester Zoo, Upton-By-Chester, UK

Human activity is changing the world at an unprecedented rate. Physiological biomarkers can provide insight into how animals respond to environmental change, predict long-term impacts of environmental change and interpret the consequences of conservation interventions. Physiological biomarkers, however, remain virtually untapped as tools for understanding large-scale macro-ecological and evolutionary processes of population dynamics. Here we use non-invasive faecal biomarkers to assess physiological responses in wild equid populations to address large-scale conservation questions. However, given the potential biomarkers have to provide useful information on wild populations where we know very little if not nothing about the individuals or their backgrounds, it is paramount that the biomarkers and assays used are analytically and biologically validated for that species. The Carneddau Welsh mountain ponies (Equus ferus caballos) are free-ranging and subject to minimal interventions. They are not provisioned and do not receive veterinary interventions. They do under-go an annual roundup, which presents a controlled stressful environment in which to validate biomarkers of physiological stress. First, we use the round-up as a biological validation to compare three different ‘stress’ assays: we compare two enzyme immunoassays that measure faecal glucocorticoid metabolites and we also explore the use of oxidative stress as a alternative marker for investigating stress in a population of free-ranging equids.

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Our results demonstrate that the use of hormone assays must be carefully validated, as observed patterns of change may not necessary reflect the underlying physiological processes we are trying to measure. We emphasise the need for thorough validations to ensure that results are robust, repeatable and accurately reflect biological patterns of interest that could be of relevance and/or importance to species management. Second, we use these three biomarkers, along with reproductive hormones to explore seasonal patterns of physiological responses across a population. If appropriately validated, a combination of steroid stress markers, oxidative stress and reproductive hormones can provide a holistic assessment of population health and variation in population performance. Ultimately, these non-invasive techniques can be applied to a range of species (both ex situ and in situ) to investigate whether conservation interventions are effective and assess causes of variation in population performance. Furthermore, they can provide data for species distribution models in order to help predict future responses to environmental change, and ultimately the long-term viability of the species. However, it is fundamental that care is taken when identifying biomarkers and the assays to use for faecal studies, as this remains a relatively new area within Conservation Physiology with huge potential to revolutionise how we monitor wild populations of conservation interest and biological importance.

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POSTER 20 – Student

Differences in spatial positioning of individuals among horse groups

Sota INOUE1, Shinya YAMAMOTO2, Monamie RINGHOFER2, Renata MENDONÇA3, Satoshi HIRATA1

1 Wildlife Research Center, Kyoto University 2 Institute for Advanced Study, Kyoto University 3 Primate Research Institute, Kyoto University

Animal behavior studies have revealed that individual health, social relationships, and spatial positioning interact to influence group dynamics. However, there are no studies assessing whether the spatial positioning of individuals is consistent across group of the same species and in similar environments. For this study, I assessed whether the spatial positioning of individual horses (Equus caballus) differed among groups with the same number of horses in similar environmental conditions. Three feral, horse groups were followed in Serra D’Arga, Portugal. For each group, approximately 65 drone images were analyzed to examine the nearest neighbor distance between individuals and level of dispersion. Results show that the nearest neighbor distance between individuals was similar for all three groups (2-3 body lengths). Additionally, the level of dispersion was different between the groups. In conclusion, horses may have an innate behavior to maintain a certain distance from nearest neighbors, i.e. personal space, while the variation in levels of dispersion between all individuals indicates that group assemblages are influenced by individual characteristics.

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POSTER 21

Space use of feral horses from non-invasively collected fecal samples

Sarah R. B. KING1, Kathryn A. SCHOENECKER1,2, Jennifer FIKE2, Sara OYLER-MCCANCE2

Despite their large size equids can be surprisingly cryptic on the landscape and individuals can be difficult to tell apart without good photographs and/or in-depth knowledge of the population. Understanding space use of individuals can inform their management and conservation, but this information can be difficult to obtain through conventional surveys. We tested whether distribution of individuals could be determined using genetic information in feces. We studied a feral horse population in Colorado, USA using non-invasively collected fecal samples, to explore whether multilocus genotypes can be used to determine space use and potential sub-population delineation. We collected >1,800 fecal samples from across the146 km2 Little Book Cliffs Herd Management Area (HMA) in May, August, and October 2014. Of these, 976 samples amplified across all eight loci, resulting in 128 unique multilocus genotypes (each representing a unique individual). Eighty-one individuals were found from 2-18 fecal piles (mean = 6.1 piles) within a single sampling area across sampling sessions, suggesting residence in this area. Individuals were more related within an area than between areas, although relatedness was not higher in some areas than others. Where individuals were found in more than one sampling area, 73% of samples were found in only one of the areas, and the areas tended to be close to each other, sharing an average

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of 1.1 borders. Our data indicated that most individuals have a very specific use of available space. Ultimately this could lead to sub-population formation. Discriminant function analyses indicated there were five genetically similar groups in the population, but these did not correspond to resident areas where genotypes were found so sub-populations were not yet evident. We suggest that fecal DNA can be a useful tool to show space use of individuals from their multilocus genotypes, and potentially identify sub-populations. This method could be useful for understanding the distribution of individuals in endangered populations, and to enable more targeted conservation efforts.

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POSTER 22

Population dynamics of Persian wild ass (Equus hemionus onager) in Qatrouyeh National Park, Iran

Haniyeh NOWZARI1

1 Department of Environment, Abadeh Branch, Islamic Azad University, Abadeh, Iran

As an endangered species, populations of Persian onager have been declining within the country over the last few decades. Unfortunately, the basic ecology of onagers in Iran is poorly understood and there is a need to understand the demography of the remained populations. According to my study on the largest remaining population of onager occurring in Qatrouyeh National Park (QNP) the abundance of the population size was estimated by distance sampling method over two consecutive years (2009-2010) at 302 with 95% confidence interval of 213– 429 onagers. So, my population model projected an increasing population over the next 30 years incorporating a maximum survival rate for adult age class. The Department of Environment’s rangers count the population each year by direct observation method. A ten-year-population size monitoring (2008-2018) shows population increasing therefore it seems that the model foresighted correctly. The result of population dynamics estimation shows population growth rate of 0.09 and 0.14 during 1998-2008 and 2008-2018, respectively. The intrinsic population growth rate indicate a 917% growth over twenty-year-period (1998-2018). Comparing these estimates with those reported previously, suggested an expanding population for QNP. As a result, doubling time will be just five years for onagers to reach nearly 2000 individulas in QNP. The model prospected that if for any reason the survival rate of matures drops then sustainability of the population will

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be challenged. Hereon, we will have to increase the survival rate of foals using management methods to prevent population extinction. The obtained results have implication for conservation and management of the species e.g. through reintroduction programs.

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POSTER 23

North American horse spatial ecology: GPS tracking insights from desert to boreal

John Derek SCASTA1, Jacob D. HENNIG1, Jeffrey L. BECK1, Sonja E. R. LEVERKUS2,3, Samuel D. FUHLENDORF3, Marten GEERTSEMA4, Brady W. ALLRED5, Mark GREGORY3, Alexandre R. BEVINGTON4, David M. ENGLE3

1 University of Wyoming, Laramie, Wyoming, USA 2 Shifting Mosaics Consulting, Fort Nelson, British Columbia, Canada 3 Oklahoma State University, Stillwater, Oklahoma, USA 4 British Columbia Ministry of Forests, Lands, Natural Resource Operations and Rural Development, Prince George, British Columbia, Canada 5 University of Montana, Missoula, Montana, USA

Our understanding of the spatial ecology of free-ranging horses in North America has been constrained by the limited use of global positioning system (GPS) collars. The use of such technology can generate robust location data to understand how horses use the landscape relative to disturbance, plant communities, topography, jurisdictional boundaries, etc. I present 2 recently published North American studies from GPS collars on 9 mares in Wyoming, USA (i.e., cold desert) during summer of 2017 and 13 male horses in British Columbia, Canada (i.e., boreal forest) from 2010-2012. Horses in the cold desert inhabited a 3,413 km2 federally managed area with annual precipitation that ranges from 146-207 mm and elevation ranging from 1,883 to 2,506 m. These horses are under the purview of the US federal government. Horses in the boreal inhabited 2 separate areas that range in size from 12,950-19,655 km2 in boreal (white and black spruce), spruce- willow-birch, and alpine tundra with annual precipitation of 446 mm and elevation

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ranging from <800 m to >2000 m. These horses are owned by private guide outfitters. In both studies, Lotek GPS collars were used with 2-hour and 1-hour fix rates respectively. The study about the spatial ecology of cold desert horses was an initial assessment of movement data during the summer 2017 specifically to quantify home ranges, movement across jurisdictional boundaries, and identify potential issues with horse distribution. The cold desert horses had mean daily movement distances of 9.0 km (SE = 0.3), and mean summer total home range sizes of 40.4 km2 (SE = 6.7). The cold desert horses used both areas outside of the designated horse management area (HMA) as 44.9% GPS fixes were outside of HMA. In addition, 10.8% of the GPS locations for these federally owned and managed were on private land. These results provide a first GPS-based quantification of home ranges for horses in the cold desert and demonstrate the difficulty of managing free-roaming horses on landscapes with heterogeneous sociopolitical patterns and designated management areas with permeable boundaries. The study about the spatial ecology of boreal horses was an assessment of horse resource selection functions relative to disturbances such as fire, vegetation cover type, and topography. The boreal horses preferentially selected recently burned areas and areas that burned more frequently when they were available. These boreal horses also avoided steep slopes and forest cover types but were attracted to grass cover types and in some cases aspen cover types. As such, fire and the ecological process of pyric herbivory (i.e., fire driven grazing), are important considerations when managing boreal rangelands and large free roaming ungulates like horses in the boreal. Because historical fire regimes and land ownership patterns of the boreal region of Canada differ from the cold desert of the United States, the role of disturbance and jurisdictional boundaries vary in their role in horse management. These studies in tandem demonstrate the robust spatial data that can be developed to answer specific questions in the diversity of habitats that horses inhabit in North America.

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POSTER 24

Individual behavioral features of harem stallions in a population of feral horses

Natalya SPASSKAYA1

1 Zoological Museum of Lomonosov Moscow State University, Russian Federation

A population of feral horses has been being existed since the 1950s on the island Vodny, Lake Manych-Gudilo (the territory of the Rostov Biosphere Reserve, Rostov region, Russia). Human interference in the life of the population involves organization of the watering and regulation of the number of animals at a level corresponding to the forage resources of the territory (some animals used to be removed from the island each year since 2012). Horses maintain a natural social structure consisting of harem bands, bachelor and mixed groups (multi-male, youth groups, etc.). The continuous monitoring of the population has been carried out since 2006. We began to conduct detailed studies using the biographical method after 2010, when the entire social structure had been completely reformed due to significant population decline caused by some natural disasters. It is known that individuals use to move constantly between social groups in the populations of wild and feral horses. This includes first of all dispersion of young animals, namely females into harem and mixed groups, and males into bachelor groups. Adult animals move between groups as a result of mare captures by stallions, usually with disintegration of existing and formation of new harem or mixed groups. Such movements maintain integrity of the population. The conducted long-term observations showed a significant variation in the behavior of stallions that lead social groups. We analyzed the cases of the movements of the adult mares and the dispersion of the youngsters between harems or harem and mixed groups persisting

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as long as 3 to 8 years (all 19 groups). In general, the population is characterized by a relatively stable social structure: we recorded an average of 6.1 transitions of adult mares per year, mainly due to the disintegration of groups resulted from regulatory measures or due to aging of their leaders. Several types of groups were revealed in the population studied: a) in which the process of leaving of adults and young mares significantly prevailed over the arrival; b) in which the arrival of adults and young mares prevailed over leaving; c) groups with active mutual exchange of young individuals; d) groups that accumulated dispersing young mares. The process of dispersion of young mares in the population was slowed down: in 30% of the population groups, only 30-50% of animals in this category left the family groups by the age of 3 years. The uncovered behavioral patterns did not depend on the individual age of the harem stallions and the time of persistence of social groups. They are apparently depend on individual characters of the animals.

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Genetics & Morphology

POSTER 25 – Student

Characteristics of the enamel ultrastructure of Przewalski's horse (Equus przewalskii) in the Chernobyl Exclusion Zone (Ukraine)

Vitalii DEMESHKANT1, Kateryna SLIVINSKA2

1 Faculty of Biology and Animal Breeding, Wrocław University of Environmental and Life Sciences, Wrocław, Poland 2 Schmalhausen Institute of Zoology NAS of Ukraine, Kyiv, Ukraine

The morphology of the teeth and the structure of their enamel reflect the morpho- functional characteristics of the process of consumption and chewing of plant foods and is closely related to the function and conditions of the habitat of the species. The structural and functional features of this process are species-specific, evolutionarily parallel in both onto- and phylogenesis and characterize the correlation dependences in the dynamics of adaptatiogenesis. Predominantly transverse (and longitudinal for incisors) enamel sections of various morphological structures (conids) of Equus przewalskii molars were studied using an electron microscope. An analytical comparison of the data obtained with morphologically close species (Polish Konik, tarpan, domestic horse) was carried out. It is confirmed that the ultrastructure of enamel on the transverse sections (thin sections) of the teeth and their constituent structures - conids is of the same type and consists of a radial layer. Tangential and lamellar layers of enamel are absent. There were no differences in the structure of the enamel of the molar and pre molars, and the enamel of the incisors is characterized by the presence of HSB structures. In the post embryogenesis of permanent teeth, the enamel is also not

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significantly different. That is, an important characteristic of the enamel structure is characteristic of the Przewalskii horse - its relative stability and insignificant individual variability, which may also apply to other types of horse. The radial layer of enamel - R includes a prismatic matrix - IPM, linear or cellular structure and crystalline prisms - PE, laid in a different way, but almost always inclined towards the OES boundary. This layer is differentiated and can be structurally divided into three types - I, II, III (Kilic et al. 1977). A trend was observed in the quantitative ratios of IPM and PE, aimed at reducing the proportion of IPM and strengthening the structure by increasing the beams of prisms. In rare cases, molars have a very specific type of enamel - HSB (Hunter-Schreger-Band) uniserial variations, which is characteristic of the enamel of Equidae incisors, including the hipparions. The enamel structure of the incisors of the species consists of linearly laid and stripes of HSB prisms. Studies of many authors have shown that the ecological-adaptive characteristics of the species — the Przewalskii horse in the Chernobyl zone is very positive, which indicates the optimal conditions for its dwelling under the conditions of Polesie. The most functional organs - the teeth and the ultrastructure of their enamel also indicate the optimality of these characteristics, which is confirmed by our comparative morphological studies. As a result, this allowed us to characterize the structure of the enamel Equus przewalskii in the Chernobyl zone in close relationship with the function of the teeth and justify its importance as a morphological feature.

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POSTER 26

Group size, group composition and level of intestinal parasites of Przewalski`s horses in the Chernobyl Exclusion Zone

Nataliya YASYNETSKA1, Daniel KLICH2, Kateryna SLIVINSKA3, Vitalii DEMESHKANT4

1 Biosphere Reserve "Askania Nova", Askania-Nova, Ukraine 2 Warsaw University of Life Sciences, Warsaw, Poland 3 I. I. Schmalhausen Institute of Zoology, NAS of Ukraine, Kyiv, Ukraine 4 Wroclaw University of Environmental and Life Science, Wroclaw, Poland

In 1998 and in 1999, the Reserve Askania Nova transferred and release to the wild 2 reproductive (12 and 7 individuals) groups and 6 males of Przewalski`s horses to Chernobyl Exclusion Zone (CEZ). Results of last four-years observation of the group size, group composition and level of intestinal parasites of the Przewalski`s horses are analysed. Five field trips per 4–9 days to CEZ using car were conducted during vegetation seasons in 2015-2018. When horses were sighted, they were observed with the aid of binoculars or a telescope. The number of individuals were recorded in each group. Clinical signs of parasitosis such as body conditions and delay in the molting were noted. The McMaster method was used to document the level of intestinal parasites. Totally, 98 individual fecal samples of horses from different groups were collected, examined and calculate was the level of infections as the number of main groups of parasites` eggs per 1 gram of horses` feaces (EPG). In total, 187 Przewalski`s horses were counted in CEZ during 2015-2018. At the beginning of 2015, more than 100 individuals in 10 groups (10.71.4 individuals per group, range 7–22) were recorded. Six reproductive (8.71.7; 4- 15), one bachelor group consist of 7 males and two single bachelors were recorded

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in summer 2015. Four reproductive (6.81.8; 4-12), one bachelor group consist of 2 males and two single bachelors were recorded in 2016. Three reproductive (8.33.2; 3-14), two bachelor (2 and 5 males, respectively) groups and two single bachelors were recorded in 2017. Five reproductive (9.02.3; 3-13), three bachelor groups (2, 6, 2 males, respectively) and one single bachelors were recorded in 2018. During 2015-2018 average number of animals in the reproductive groups was 7.70.9; in bachelor groups – 3.90.9. An increase in population size led to the formation of a larger number of groups and a change in the average number of animals in groups. The level of infection of horses by the main groups of intestinal parasites has been determined. The eggs of intestinal nematodes (Strongylidae; Parascaris equorum; Habronematidae) and tapeworms (Anoplocephalidae) were found. The prevalence of Strongylidae was 100%, the mean eggs per gram range from 50 to 888,9±565,7. Parascaris equorum eggs were recorded in the various groups of horses (33-100%; 25-200 EPG). In additional, a few eggs of habronems and anoplocephalides were noted in some horses (16,7%; 25 EPG and 9,09-66,7%; 25 EPG, respectively). The visible outer changes in animals with high parasites infestation were not observed. We consider necessary to continue regular study of Przewalski's horses in the Chernobyl exclusion zone.

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Equid reproductive biology

POSTER 27

Reproductive management of the feral horse population of Letea Sandbank – Danube Delta

Ovidiu ROȘU1,2

1 University of Agronomic Sciences and Veterinary Medicine Bucharest 2 Four Paws International

The following represents the summary of the PhD thesis assessing the feasibility of PZP vaccination following remote capture on the free roaming horses from Letea Sandbank from 2011 to 2017. Five annual flights were conducted in order to provide a count of the horse population, estimated to X̅ = 535. Throughout the research, several horse capturing methods were tested, which, depending on field results, were modified and/or abandoned. In 2011–2013, funnel enclosures capture was tested on 55 feral horses, which had to be blood sampled and load for transport several times. On this group, 102 cases of remote chemical immobilization were done. In performing these procedures, a standard combination of 25 mg detomidine, 62.5 mg tiletamine, 62.5 mg zolazepam and 10 mg butorphanol delieverd by 3.5-ml darts with the blowpipe. In most cases (78.4%) the horses had a complications-free induction and recovery. The combination of anesthetics proved to be appropriate and safe, capable of inducing and maintaining a good anesthetic plan for an average duration of 35 minutes. Furthermore, chemical capture method using ketamine, medetomidine and hyaluronidase was tested on a number of 35 free-range horses. The average induction time recorded for 28 ketamine-immobilized horses (X̅ = 2.32 mg / kg)

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and medetomidine (0.09 mg / kg) was 8.85minutes, while mean induction time for the group of 7 horses immobilized using ketamine (X̅ = 1.74 mg / kg), medetomidine (X̅ = 0.119 mg / kg) and hyaluronidase (X̅ = 3.64 IU/kg) was 9 minutes. The mean duration of anesthesia was 69.56 min, respectively 73.8 minutes. Peripheral oxygenation (SpO2) in most of the cases was below the normal limit. The mixture proved to provide adequate muscle relaxation and a sufficient anesthesia, with gradual wakening, requiring no antagonists. Following the testing of capturing methods, between 2013 and 2017a total of 246 females were tranquilized remotely, for the purpose of PZP vaccination. PZP effect was initially studied in 150 chemically immobilized mares, with subsequent remote administration of a booster, using the dart gun. Ear tagging was used, due to hard individual identification. Fecal estrogens and progesterone metabolites (20a-OH- pregnane) were analysed. Results showed that the gestation rate in the group of PZP-treated mares was 14.6%, as compared to 81.5% for the control group, despite that 44% of the vaccinated females had not receive the booster. Horses showed a cumulative behavioural aversion to being targeted (from 20m to 50m). Following PZP vaccination, the average contraception rate for the entire free-ranging horse population of Letea Sandbank, was 46.25%. A contraceptive program based on the PZP vaccine, using individual chemical immobilization followed by remote administration of boosters, is a viable alternative to the classical capturing of free- range feral horses. However, this type of approach consumes time and resources, evidencing the need for developing a product that can work effectively for several years, supplanting annual administration of boosters.

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Feral equid issues and rewilding

POSTER 28

Meta-analysis of diet composition and potential conflict of wild horses with livestock and wild ungulates on Western Rangelands of North America

Jeffrey L. BECK1, J. Derek SCASTA1, Catherine J. ANGWIN1

1 Department of Ecosystem Science and Management, University of Wyoming, Laramie, Wyoming, USA

Wild horse (Equus ferus caballus) management in western North America is an escalating concern for ecological integrity on these landscapes. Identifying potential diet overlap among horses, livestock, and wildlife will inform management decisions to optimize multiple interests. To understand dietary relationships, we conducted a quantitative synthesis of microhistological fecal studies for wild horse, beef cattle (Bos spp.), domestic sheep (Ovis aries), elk (Cervus elaphus), pronghorn (Antilocapra americana), and mule deer (Odocoileus hemionus) diet composition on western rangelands of North America. Our search yielded 60 studies from 14 states, 1 Canadian province, and 2 Mexican states with 392 unique species-season samples. We summarized plant species into graminoid, forb, and browse functional groups. For wild horses, seasonal diet composition means for graminoids (77−89%), forbs (4−15%), and browse (3−10%) did not vary seasonally for any plant group (P ≤ 0.05). Univariate analyses and the calculation of effect sizes corroborated our finding that graminoid composition explained the potential overlap of wild horses with cattle regardless of season, with sheep and elk in the spring, with sheep in the summer, and with elk in the fall and winter. Although data indicate wild horse diets are primarily composed of graminoids,

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several studies reported unusual, regionally specific shifts in response to winter snow that limited graminoid accessibility, leading to higher browse composition. Season, plant composition, and ungulate assemblage may all influence dietary competition between wild horses and other large ungulates sharing western North American rangelands; however, the low and nonsignificant heterogeneity values at alpha 0.01 for cattle:horse effect size comparisons suggest that cattle and horses respond to regional and seasonal variation similarly—a result not observed for other ungulate:horse comparisons. Our meta-analysis provides a robust data set for evaluations of diet composition for wild horses, livestock, and wildlife, whereas no empirical studies have assessed all species together.

Note: this abstract originates from our published paper of the same title (Scasta, J. D., J. L. Beck, and C. J. Angwin. 2016. Meta-analysis of diet composition and potential conflict of wild horses with livestock and wild ungulates on western rangelands of North America. Rangeland Ecology and Management 69:310–318).

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POSTER 29 – Student

Parasites in the context of rewilding

Julia HATZL1, Amos BOUSKILA2, Juán José NEGRO3, Dennis HANSEN1

1 University of Zürich, Switzerland 2 Ben-Gurion University of the Negev, Israel 3 Estación Biológica de Doñana, CSIC, Spain

Human development has led to population declines and extinctions for many large herbivore species. This results in the loss of ecological functions, and cascades of secondary extinctions for species which depend on grazing pressure. Rewilding aims to restore lost ecological functions in ecosystems by reintroducing large herbivores to landscapes. Before reintroduced animals can implement a function in an ecosystem, a healthy, self-sustaining population has to be established. Parasites influence the health of host species and have an impact on the establishment and viability of populations through increased mortality. We quantified the prevalence of gastrointestinal endoparasites in Retuerta feral horses (Equus ferus caballus) in the Biological Reserve of Doñana National Park, Spain, and assessed the possible impacts of parasite burden on individuals’ health. Results of faecal egg counts showed that i) Retuerts show a moderate infection rate compared to other feral horse populations, and ii) only the infection with Parascaris equorum had a negative effect on body condition. These results suggest that a high infection rate with P. equorum, which is only prevalent in young horses, could have a negative effect on the establishment and recruitment of feral horse populations. However, infection with other parasite species appears to be of little concern for this feral horse breed.

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POSTER 30

Response of sagebrush habitat characteristics to feral horse grazing

Jacob D. HENNIG1, J. Derek SCASTA1, Jeffrey L. BECK1

1 Department of Ecosystem Science and Management, University of Wyoming, Laramie, Wyoming USA

Following the extinction of native equid species in North America during the Pleistocene, shrubland communities existed in absence of large mammalian grazers for roughly 10 millennia. Shrublands evolved with less grazing pressure compared to grassland and savannah systems and therefore may be more susceptible to damage from introduced large grazers such as feral horses (Equus ferus caballus) and cattle (Bos spp.) Compared to cattle, feral horses are relatively unmanaged in the United States and little is known about how these animals impact fragile sagebrush (Artemisia spp.) systems. Sagebrush habitat is crucial to native wildlife species such as pronghorn (Antilocapra americana), mule deer (Odocoileus hemionus), and greater sage-grouse (Centrocercus urophasianus; hereafter ‘sage- grouse’) to fulfill the food and cover requirements of these iconic species in many areas of western North America. High utilization by horses is thought to decrease sagebrush habitat quality through two mechanisms: 1) grazing that may in turn decrease native vegetation cover and height, and 2) trampling that may increase shrub fragmentation and soil penetration resistance. For sage-grouse, these impacts can translate to decreased nesting habitat, escape cover, and forage availability. Empirical examination of how feral horses affect sage-grouse habitat is lacking; however, these evaluations are crucial as the sage-grouse is an imperiled species that has been petitioned for endangered species listing under the U.S. Endangered Species Act of 1971. We set out to address this knowledge gap by measuring a

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suite of sagebrush habitat characteristics along a gradient of horse utilization within the Adobe Town Herd Management Area in southern Wyoming, USA in summers 2017-2019. We used spatial regression models to examine the effects of horse utilization on canopy cover of plant functional groups, grass height, and visual obstruction. Increased horse utilization was significantly correlated with increased bare ground cover along with decreased shrub cover and grass height; however, horse use was not significantly correlated with visual obstruction or native herbaceous cover. Our results show that horse utilization may decrease several characteristics of habitat quality for greater-sage grouse. To assess if sage- grouse avoid areas of high predicted horse use, we also will present the degree of resource selection overlap between the two species in summer 2019.

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POSTER 31

Human dimension of Galician wild pony traditional management in Serra do Xistral

Laura LAGOS1, Carlos MUÑOZ-BARCIA1, Jaime FAGÚNDEZ1

1 Biocost Group, Faculty of Biology and Advanced Scientific Research Center (CICA). University of A Coruña, Spain

The Natura 2000 site “Serra do Xistral”, located in the North of Galicia, NW Spain, holds a large population of over 1500 ponies living with very limited human intervention. The presence of these wild ponies contribute in a decisive way to the conservation of heathlands, blanket bogs and raised bogs, a valuable mosaic of European priority habitats included in the Habitats directive (92/43/CEE). The project Life in Common Land (LIFE16 NAT/ES/000707) aims to preserve this system by aiding pony owners to maintain or increase the number of ponies. A preliminary study was performed to address the main constraints for the pony owners. Between January and November 2018 we conducted 20 semi-structured interviews with pony owners of the Xistral mountains. About 160 pony owners maintain the activity in the area because they want to follow their family tradition, but also due to the indirect benefit they obtain from ponies: the ponies control the shrub vegetation, and specifically the gorse (Ulex spp.), improving pasture for cattle. The trend of this activity is regressive. In the last 50 years there has been a reduction of more than 50% of the pony population. At the same time, management has changed in some of the Commons in Serra do Xistral mainly due to fencing and to the creation of artificial pastures. In general, it has become more intensive. The main problems detected for the continuity of the wild pony traditional management are the abandonment of the rural area and the advanced

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age of the owners, wolf damage and the difficult verification of wolf attacks which is required to receive compensation, the application of CAP aids, low price of meat, and administrative problems. Wild ponies may be used as a management tool for habitat conservation and the maintenance of the high natural value of the Xistral mountains, but this will need efforts on promoting better regulations for pony owners.

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POSTER 32 – Student

Media, misinformation, myth and mystery – Brumbies against the flow?

Colleen O’BRIEN1,2

1 Victorian Brumby Association, Glenlogie, Australia 2 Australian Brumby Alliance, Melbourne, Australia

We read much about the damage that Brumbies do to the environment in Australia. Whilst headlines about Brumbies decimating the populations of endangered native Stocky Galaxia fish read well, how much truth is in the story…

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POSTER 33

Use of GPS collars and geospatial technology to evaluate horse habitat use in Western United States

Steven L. PETERSEN1,2,3

1 Professor of Forest and Rangeland Landscape Ecology and GIS 2 Department of Plant and Wildlife Sciences 3 Brigham Young University, Provo, Utah USA

In western North America, over 80,000 free-roaming horses (mustangs) and burros range across 11 million hectares of open public land in the United States. Horses utilize these rangeland ecosystems as a source of forage, water, and protection. With unrestricted access to these resources, the potential impact that horses have on rangeland condition and ecological processes is of tremendous importance to rangeland and wild horse managers. While the patterns of movement and animal distribution are fundamental in assessing horse impacts, this information remains poorly understood and lacking in current literature. We utilized GPS collars and geospatial technology to quantify horse movement and land use in an effort to evaluate horse impacts across heterogeneous rangeland landscapes. We collected a coordinate location every 30 minutes from 12 collared lead mares representing 9 horse bands in northeastern Nevada, USA. Each mare carried a collar between 6-24 months. We used Brownian Bridge and Resource Selection Functions to model horse movement, home range, and optimal habitat suitability. Horse habitat use was influenced by distance to water, forage suitability, resource availability and topography. With suitable grass and water availability, distances travelled were minimized, however, this varied across seasons with higher distances travelled away from water during late summer and fall months. Managers can use this

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information to predict horse habitat use and help minimize impacts to sensitive or vulnerable rangeland resources.

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POSTER 34

Effect of radio collars and tail transmitters on feral equids in the USA

Kathryn A. SCHOENECKER*1, 2, Sarah R. B. KING2, Jacob D. HENNIG3, Jeffrey L. BECK3, and J. Derek SKASTA3

1 U.S. Geological Survey, Fort Collins Science Center, Fort Collins, CO 80526, USA 2 Natural Resource Ecology Laboratory, Colorado State University, Fort Collins, CO 80523, USA 3 Department of Ecosystem Science and Management, University of Wyoming, Laramie, WY 82071, USA

Radio collars and other radio tracking devices have been invaluable tools for wildlife managers for over 40 years, providing a greater understanding of wildlife population movement rates, spatial ecology, habitat selection, seasonal range, and demographic parameters such as immigration and emigration, that facilitate development of models for conservation and management. This essential tool has been used on virtually all North American ungulates, but rarely on feral horses or burros. To determine if radio collars could be applied to feral equids we conducted an observational study to investigate fit and wear of radio collars affixed on feral horses and burros in 5 free-roaming populations (2 burro populations, 3 horse populations). Radio collaring equids is challenging because their neck can be much thicker when the head is elevated/alert compared to head-down grazing position. We measured a circumference difference of 15 cm in these two different positions in feral horses. This makes fitting a radio collar more difficult and increases probability of complications. We conducted a study from 2016 to 2019 to assess collar impact on individual behavior, body condition, and evaluated neck surface

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for wounds, signs of chafing, or other effects. We also tested longevity/duration and effects of GPS transmitters that we braided into the tails of feral horses. We found no significant impact of collars on the necks of mares or burro jennies, nor any changes in body condition. However, 10% of collars flipped forward over the ears onto the foreheads of equids, which posed a risk of mild to moderate discomfort or obstruction of movement. In several cases incorrect collar position in front of the ears caused wounding and collars had to be removed. In several cases radio collars flipped back into the correct position on their own. We found no behavioral impact from tail transmitters, and most lasted 6-14 months after which tail hair grew and the tag was expelled. We had better results in terms of tag longevity when trail transmitters were lighter (~40 grams as opposed to 70 grams). We conclude that collars can be used on free-roaming feral equids, but collars need to be placed correctly just behind ears, affixed tightly, and care should be taken to get visual observations regularly to assure collars are remaining in correct position on the neck. We recommend making sure collars can be removed if needed with iridium signal or some type of remote controlled drop-off mechanism.

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POSTER 35 – Student

Strangers in their own lands. How to manage wild horses and burros in a modern American society

Elke TUKKER1

1 University of Amsterdam, Amsterdam, History student; University of Washington, Seattle, Environmental Sciences and Resource Management Exchange student; The Cloud Foundation, Colorado Springs, intern

Modern equids (genus Equus) evolved for most of their evolutionary history in North America. Being one of the larger grazers, they performed a crucial ecological role in keeping the landscape open, providing habitats for birds, rodents, and other small mammals, and supporting the growth of a large variety of plants. After spreading across the world over the Beringia land bridge, North American equids likely went extinct only 6,000 to 10,000 years ago. After this short interval, horses and burros were reintroduced with the arrival of the Spanish conquistadors about 600 years ago, returning to their ecological home. For many generations after this reintroduction, millions of wild horses roamed what is now the western United States, once again filling their ecological niche. Not only did wild horses fill an ecological niche, they also transformed native-American and American identity. Native-Americans made extensive use of gone-wild horses in their battle against settlers. In addition, horses were used for hunting purposes and became an important aspect of many spiritual rituals. Through time, the elusive wild horse herds became an integral part of the American landscape. The essentialism of wild horses for both native and non-native Americans has caused these animals to attain a legendary, even mythical place in American history. However, modernization, domestication and industrialization have changed the fate of American wild horses.

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In 1970, hunting had brought wild horse populations down to a mere 25,000 and they were considered “fast disappearing”. Public outcry to protect wild horses caused the American Congress to unanimously pass an Act designed to protect them. Continuous mismanagement of the users of public lands, including livestock, wild horses and other wildlife has caused habitat degradation in some of America’s most fragile landscapes. As of today, many interest groups such as ranchers, ecologists, advocacy groups, politicians, and native Americans are in conflict over wild horse and burro issues. While stakeholders are polarized on issues regarding wild horses and burros, such as range degradation and ethical concerns, pressure on America’s soil, water, wild horses and burros, and other wildlife increases. Arguably, it is one of the most critical wildlife issues in the western United States. Although there is a historic niche for equids in America’s ecology, wild horses and burros have become strangers in their own lands. How do we manage wild horses and burros in a modern American society?

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POSTER 36 – Student

The use of GPS monitoring for analyzing moving behavior in horses during the presence of wildlife

Linda VOGEL1, Sarah WIEZOREK1, Peter SCHUETTE2, Ernst-Hermann SOLMSEN3, Hanne MARTIN3, Enno HEMPEL3, Theo GRUENTJENS3, Markus BATHEN3, Peter HEROLD4, Sophia VOIGTLAENDER-SCHNABEL1, Konstanze KRUEGER1

1 Faculty Agriculture, Economy and Management, Nuertingen-Geislingen University, Germany 2 Nature And Biodiversity Conservation Union, Lower Saxony, Germany 3 Working Group Horse Wolf, Lower Saxony, Germany 4 Society for the Protection and Conservation of Wolves, Germany

With the return of wolves to Germany, horse keepers and owners fear injuries and predation in horses caused by wolves, as well as panic-like flight responses in horses, which may cause escapes from pastures and accidents on the roads. To obtain valid information about the reactions of horses towards wolves and other wildlife under natural conditions, we tested the usefulness of monitoring in groups of horses via GPS combined with camera trapping of wildlife. For the data collection, two horse groups kept on separate pastures were equipped with GPS devices (GPS Lap Timer BT-Q1000eX). In total the movements of seven horses were recorded over a period of six weeks (20th August – 30th September 2018) every night between 6pm in the evening until 8am in the morning. For each horse we recorded the longitude, latitude and the moving speed once per second. Additionally, five camera traps were positioned around each pasture. The GPS data were analysed with a newly developed R script and the computer software R. The

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horses’ moving distance, speed, position and distances within the group were compared with occurrences of wildlife recordings at the camera traps. In total, 228 wildlife recordings were documented, of which 26 could be connected to flight responses in the horse groups.

The analysis of speed showed that movements responding to wildlife occurrences were significantly slower (GLM N = 64, t = 2.574, p = 0.013) than the average speed of horses responding to other stimuli (≥12km/h). This leads to the assumption that the presence of wildlife does not frighten horses per se. Only in 11% of the 228 wildlife occurrences, the horses showed faster movements. This supports the argument, that horses do not generally show flight responses in wildlife presence. The analysis of the distance between the horses demonstrated that, in general, horse pairs were significantly closer to each other in slow movements (≤12km/h) than in faster movements (GLM N = 153, t = 5.755, p < 0.001). However, the distance between the horses declined during a wildlife occurrence with movement reactions (GLM N = 84, t = 5.919, p < 0.001). The results indicate that horses reduce their individual distance even in mild frightening situations and move closer together.

The GPS monitoring allowed the analysis of the group structures. The distance between horse pairs varied greatly (GLM N = 153, BD t = -4.682, p < 0.001, BE t = -3.965, p < 0.001, DE t = -3.589, p < 0.001, FG t = -4.614, p < 0.001, FH t = - 5.123, p < 0.001, GH t = -4.314, p < 0.001), especially when movements were close to zero (≤0.126km/h). We suggest that horses kept lower distances with socially bonded animals in calm situations. This assumption needs closer analysis with the help of network analysis. The monitoring of horse groups with GPS devices is a promising tool for field research. To validate the results, further research in horse groups with various sizes and compositions, and the application of network analysis is needed.

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POSTER 37

Hide nor hair –the illicit trade in donkey hides is a threat to wild asses

Zoe RAW1

1 The Donkey Sanctuary, Slade House Farm, Sidmouth, Devon EX10 0NU, United Kingdom

The global donkey population is estimated at 44 million and is largely associated with economically developing nations where donkeys are used as working animals. Donkeys play a central and critical role in supporting the livelihoods of millions of people accross the world, providing support for farming, enabling access to resources, and in food production. Global demand for diverse products of donkey origin has escalated rapidly, with a particular interest in the premium products resulting from donkey skins.

Ejiao is a traditional Chinese medicine product which is based upon extracts of donkey gelatin from donkey skins, mixed with herbs and other ingredients to form a gelatinous bar, which is marketed as a miracle cure for multiple health probems. Since 2010, consumer demand for ejiao has increased rapidly, and subsequently, so has the demand for donkey skins. The Donkey Sanctuary estimate that a minimum of 1.8 million donkey skins are being traded per year, but this may be a gross underestimate. The increasing wealth and diaspora of the Chinese middle classes, alongside the apparent credibility of ejiao products, appears to have created such a high level of demand for donkey skins that global supply is struggling to keep up, leading to high prices and widespread claims of fraud. Such high levels of demand by the Chinese market are fuelling global reports of donkey theft and a sudden increase in the purchase price of donkeys. Products of donkey origin are so highly sought after that ejiao can sell for up to USD $500/kg.

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The alarmingly high demand for donkey skins, and high prices that a donkey skin can fetch, positions donkey skin in a similar position as ivory or rhino horn. There is a complete lack of regulation over the utilisation of donkeys for the skin trade to fuel ejiao production, and new slaughter houses are opening at rapid rates to keep up with demand. Consequently, there has been a sharp rise in donkey thefts.

As well as being unsustainable and harmful to rural livelihoods, this illicit trade could have devastating effects on populations of wild asses. Donkey skin is highly valuable, yet increasingly scarce, making it feasible that traders will start targeting wild asses. The Afrcan Wild Ass Equus africanus is Critically Endangered, with fewer than 200 mature indivduals remaining in the wild; unfortunately, the species occupies an area of Africa where the trade in donkey skins is high, exposing it to risk of being targeted. There is also some indication that populations of Asiatic Wild Ass Equus hemionus could also come under threat, given their proximity to China and surrounding socioeconomic climate.

The international tade in donkey skins has emerged rapidly and fiercely, and is grossly unsustainable. The trade has the significant potential to eradicate populations of donkeys accross Africa, and poses a serious threat to the conservation and survival of African Wild Ass populations. In this presentation I raise and highlight these issues, and open up discussion for how this new threat may be mitigated for the conservation and management of wild equids.

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POSTER 38 – Student

Ongoing research on social dynamics of reintroduced Przewalski´s Horses in the Great Gobi B Strictly Protected Area (Mongolia)

Anna BERNÁTKOVÁ1, Ganbaatar OYUNSAIKHAN2, Jaroslav ŠIMEK3, Martina KOMÁRKOVÁ1, Francisco CEACERO1

1 Faculty of Tropical AgriSciences, Czech University of Life Sciences Prague, Czech Republic 2 Great Gobi B SPA, Takhin Tal, Mongolia 3 Zoo Praha, U Trojského zámku 120/3, 171 00, Czech Republic

Przewalski's horse was listed as Extinct in the Wild (EW) from the 1996, with the last individual seen in 1969. Thanks to the ongoing reintroduction efforts, the species is listed as Endangered from 2011. At the Great Gobi B SPA, the first reintroduced horses were released in 1997 and currently, there is a mixture of recently reintroduced, long-term reintroduced and wild-born animals (wild-born individuals are in majority). Behavioural research may support the conservation efforts since social factors like the stability of the harem, its social structure and the role of the group members in decision making are critical in the response of populations to the environmental factors. Our research is focused on the social dynamics of the species at the individual, harem and population level. During this research, the social network of certain selected harems will be studied and monitored along time. The individual behaviour of group members with different origin and their role within the harem will be studied. Differences in behavioural preferences among recently re-introduced, long-term re-introduced and wild-born individuals will be investigated, and the behavioural patterns explaining an adequate adaptation to the environment will be identified. The data collection is

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being conducted in three different seasons (late spring, summer, autumn). The harems are located by the local rangers during the daily monitoring routines. Once located, the selected harems are approached and filmed from a close distance (from 150m to 500m) by standard Panasonic video camera. Kestrel 4500 Pocket Weather Tracker is being used for the collection of weather conditions. Location and cohesion of the harems is assessed with a rangefinder and digital compass. Behavioural Observation Research Interactive Software (BORIS) is being used for event logging, video coding and observation of the captured videos. Understanding the influence of environmental factors on the activity budget, the impact of the animal origin (recently re-introduced, long-term re-introduced and wild-born) on the structural group properties, and the links between activity and patterns of habitat use, are the main outputs expected from this research.

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POSTER 39

Effects of ex-situ captivity and lactation on immune functions in zebra mares

Peter A. SEEBER1,2, Thomas MORRISON3, Alix ORTEGA4, Alex GREENWOOD1,5, Gábor CZIRJÁK1

1 Limnological Institute, University of Konstanz, Germany 2 Leibniz Institute for Zoo and Wildlife Research, Germany 3 Institute of Biodiversity, Animal Health and Comparative Medicine, University of Glasgow, United Kingdom 4 Réserve Africaine de Sigean, France 5 Department of Veterinary Medicine, Freie Universität Berlin, Germany

Immunity is affected by genetic, physiological, and environmental factors. Ex-situ captivity differs from natural environments in many respects, e.g. diet, climate, social factors, movement space, and anthropogenic interference. Abundance and diversity of pathogens (pathogen pressure) can be assumed to be higher in the wild than in captivity for animals that live in tropic and subtropic environments. Lactation can also affect the immune system as the associated energetic demands may impose energetic constraints on immune functions whereas passive immunization of nursing offspring demands up-regulation of certain immune factors. We investigated immunity in zebra mares in ex-situ captivity in European zoos versus zebras in African national parks, and the effect of lactation on functional aspects of the constitutive and induced innate and of the adaptive immune system in plains and mountain zebras (Equus quagga and E. zebra). As a proxy for general immune function, we screened for lytic equine herpesvirus (EHV) infection, a common and mostly latent pathogen which is reactivated in response to stress and immune challenge. We observed significantly lower levels

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of constitutive innate immune factors in captive than in free-ranging zebras. In contrast, factors of the energetically costly induced immunity were higher in captive animals. Lactation was associated with higher titers of natural antibodies and lysozyme. Lytic EHV infection was not significantly correlated with any of the measured immune markers. Our results indicate that captivity and lactation may be important immunomodulators in zebra mares.

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POSTER 40

PZP-induced mare infidelity increases male aggression and stress in the feral horse

Cassandra M.V. NUÑEZ1,2 and Maggie M. JONES1

1 Iowa State University, United States of America 2 University of Memphis, United States of America

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POSTER 41

Social network dynamics and home-range overlap among horses in extremely seasonal habitats

Amos BOUSKILA1,2, Dafna BRICMAN1, Roni BELENKI1, Juan José NEGRO2 and Eloy REVILLA2

1 Department of Life Sciences, Ben-Gurion University of the Negev, Beer-Sheva, Israel 2 Estación Biológica de Doñana (CSIC), Spain

Feral Retuerta horses in Doñana Biological Reserve experience dry summer and fall and in winter marshes may flood large areas of the reserve. These dramatic environmental changes affect availability of forage and water and may thus affect the social structure of the horses. We tested the hypotheses that during the dry season 1. Groups show a greater overlap in home-range, due to scarcity of patches of food and water and 2. Network modularity will be lower, due to increased opportunities for mixture of groups when using the few available patches. We documented horses' group affiliation between July 2014-Feb 2015, and divided these months to a dry and a wet season, based on monthly rainfall and NDVI. We analysed the weighted social networks separately for the two seasons and compared them.

Ten harems were detected in both seasons and a few of them remained mostly unchanged. However, the composition of some of the groups was very different: a few groups split or merged, and a few horses died towards the end of summer. Group home-ranges overlapped more during the dry season, however, network modularity was higher in the dry season, i.e., groups were less inclined to mix. We

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hypothesize that in addition to the distribution of resources, the presence of flies may have affected the movement and the cohesion between individuals in summer.

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List of Participants

Afonso Luís Filipe Faculdade de Ciências da Universidade Portugal [email protected] de Lisboa Alonso Garcia Enrique Council of State / Franklin Institute-UAH Spain [email protected] Bar-David Shirli Ben-Gurion University of the Negev Israel [email protected] Bayarbaatar Buuveibaatar Wildlife Conservation Society's Mongolia Mongolia [email protected] Program Beck Jeffrey University of Wyoming USA [email protected] Beja-Pereira Albano Research Center in Biodiversity and Portugal [email protected] Genetic Resources (CIBIO-InBIO), Faculty of Sciences, University of Porto Berman David University of Southern Queensland Australia [email protected] Bernátková Anna Czech University of Life Sciences Prague Czechia [email protected] Bews Willa accompanying Hannah Davie, from UK [email protected] Association Takh and Nottingham Trent University Bhatnagar Yash Veer Nature Conservation Foundation & Snow India [email protected] Leopard Trust Bobek Miroslav Zoo Praha Czechia [email protected] Bouskila Amos Ben-Gurion Univ. of the Negev Israel [email protected] Boyce Paul University of Saskatchewan Canada [email protected] Boyd Lee Washburn University USA

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[email protected] Brandlová Karolína Czech University of Life Sciences Czechia [email protected] Britnell Jake University of Manchester England [email protected] Burnik Šturm Martina Research Institute of Wildlife Ecology, Austria [email protected] Vetmeduni Vienna Byamba Munkhtuya Khomyn talyn takhi, NGO Mongolia [email protected] Ceacero Herrador Francisco Czech University of Life Sciences Prague Czechia [email protected] Clancy Cara The Donkey Sanctuary UK [email protected] Cooke Fiona The Donkey Sanctuary UK [email protected] Čulík Luděk ZOO Dvůr Králové a.s. Czechia [email protected] da Silva Mendonça Renata Kyoto University Institute for Advance Japan Study [email protected] Davie Hannah Association pour le cheval de Przewalski: USA [email protected] TAKH; Nottingham Trent University De Munnik Norbert Hustai National Park Trust, Mongolia Netherlands [email protected] Demeshkant Vitalii Wrocław University of Environmental and Poland [email protected] Life Sciences Dobiášová Barbora Zoo Praha Czechia [email protected] Drouard Florian Association pour le cheval de przewalski : France [email protected] TAKH Elliott Alice Port Lympne Hotel & Reserve UK [email protected] Faulkner Charlotte Friends of the Dartmoor Hill Pony UK [email protected] Feh Claudia Tour du Valat research station France [email protected] Fekete Eva Hortobagy National Park Hungary [email protected] Funnell Sheila Grevy's Zebra Trust Kenya

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[email protected] Gloneková Markéta Derbianus Conservation Czechia [email protected] Gogan Dana USA [email protected] Gogan Peter USGS Fort Collins Science Center USA [email protected] Goodrich Erin Cornell University USA [email protected] Gosling Morris Newcastle University UK [email protected] Grams Kayla The Science and Conservation Center USA [email protected] Griffin Paul USA Bureau of Land Management USA [email protected] Gutiérrez-Cepeda Luna Universidad Complutense de Madrid - Spain [email protected] Horse1 Spillers Hall Carol National Equine Welfare Council UK UK [email protected] Hatzl Julia ETH Zürich, Switzerland Switzerland [email protected] Heckel Jens-Ove ZGAP / Zoo Landau Germany [email protected] Hennig Jacob University of Wyoming USA [email protected] Hinchcliffe Danielle University of Manchester UK [email protected] Ibler Benjamin AG Zoologischer Garten Köln Germany [email protected] Inoue Sota Wildlife Research Center, Kyoto Japan [email protected] university, Japan Jagdag Enkhbayar Hustai National Park, Mongolia Mongolia [email protected] Jocelyn Poissant University of Calgary Canada [email protected] Kaaria Timothy Lewa Wildlife Conservancy Kenya [email protected] Kerekes Viola Hortobagy National Park Directorate Hungary [email protected]

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King Sarah Colorado State University USA [email protected] Kloska Alina Śląski Ogród Zoologiczny Poland [email protected] Komarková Martina Czech University of Life Sciences in Czechia [email protected] Prague Krug Martin Zoologická zahrada Hodonín p.o. Czechia [email protected] Lagos Laura University of A Coruña Spain [email protected] Langenhorst Tanya Marwell Wildlife UK [email protected] Linklater Wayne California State University, Sacramento USA [email protected] Linnell John Norwegian Institute for Nature Research Norway [email protected] Lovász Lilla University of Basel Switzerland [email protected] Maeda Tamao Wildlife Research Center, Kyoto Japan [email protected] University McLean Amy University of California Davis USA [email protected] McLoughlin Philip University of Saskatchewan Canada [email protected] Mekarska Anna Wroclaw Zoo Poland [email protected] Melichar Luboš Zoologická zahrada Liberec, příspěvková Czechia [email protected] organizace Messmer Terry Jack H. Berryman Institute, Utah State USA [email protected] University Moehlman Patricia IUCN/SSC Equid Specialist Group Tanzania [email protected] Mohammed Redwan Samara University, PhD candidate at Ethiopia [email protected] Addis Ababa University Nasanbat Battogtokh Institute of General and experimental Mongolia [email protected] biology - Mongolian Academy of Sciences Neumann-Denzau Gertrud independent researcher Germany [email protected]

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Nowzari Haniyeh Department of Environment, Abadeh Iran [email protected] Branch, Islamic Azad University Nuñez Cassandra University of Memphis USA [email protected] O'Brien Colleen Victorian Brumby Association Australia [email protected] Ochi Sakiho Wildlife Research Center, Kyoto Japan [email protected] University Oyler-McCance Sara US Geological Survey USA [email protected] Ozogány Katalin Department of Evolutionary Zoology and Hungary [email protected] Human Biology, University of Debrecen Parrini Francesca University of the Witwatersrand South Africa [email protected] Perrin Jean-Louis Association pour le cheval de Przewalski France [email protected] Petersen Steven Brigham Young University USA [email protected] Pinto Pandora Wildlife Research Center, Kyoto Japan [email protected] University Pluháček Jan Ostrava Zoological Garden and Botanical Czechia [email protected] Park Prevot Louise Netherlands [email protected] Pukazhenthi Budhan Smithsonian Conservation Biology USA [email protected] Institute Radcliffe Rolfe Cornell University College of Veterinary USA [email protected] Medicine Rademacher Ulrike Wilhelma Zoo Stuttgart Germany [email protected] Raw Zoe The Donkey Sanctuary UK [email protected] Reeder Sabrina Vectronic Aerospace Germany [email protected] Rees Lucy Proyecto Pottokas en Piornal Spain [email protected] Ringhofer Monamie Kyoto University Institute for Advanced Japan Study [email protected]

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Robovský Jan Department of Zoology, Faculty of Czechia [email protected] Science, University of South Bohemia Roșu Ovidiu ARCA - FOUR PAWS INTERNATIONAL Romania [email protected] Rubenstein Dan Princeton University USA [email protected] Samuelsson Indra Followit Sweden AB Sweden [email protected] Scasta John Derek University of Wyoming USA [email protected] Seeber Peter Univ. of Konstanz, Limnological Institute, Germany [email protected] Environmental Genomics Shah Nita Bombay Natural History Society India [email protected] Shultz Susanne University of Manchester UK [email protected] Schoenecker Kate United States Geological Survey and USA [email protected] Colorado State University Snook Victoria the Aspinall foundation UK [email protected] Souris Anne-Camille Association GOVIIN KHULAN Mongolia [email protected] Spasskaya Natalia Zoological Museum of Lomonosov Russia [email protected] Moscow State University Starbuck Gareth Nottingham Trent University UK [email protected] Steck Beatrice Zoo Basel Switzerland [email protected] Szabados Timea Hortobagy National Park Directorate Hungary [email protected] Šandlová Kateřina University of South Bohemia in České Czechia [email protected] Budějovice Šimek Jaroslav Zoo Praha Czechia [email protected] Tatin Laurent Association TAKH France [email protected] Tesfai Redae Teclai University of the Witwatersrand, School South Africa [email protected] of Animal, Plant & Environmental Sciences

200

Thynell Cecilia Followit Sweden AB Sweden [email protected] Tserendeleg Dashpurev Hustai National Park, Mongolia Mongolia [email protected] Tserendorj Munkhbat "Khomyn Talyn Takh" NGO Mongolia [email protected] Tukker Elke The Cloud Foundation USA [email protected] Valášková Irena Zoo Praha Czechia [email protected] van Dierendonck Machteld Equine Clinics Utrecht (NL), Gent & Netherlands [email protected] Antwerp (BE) Vodička Roman Zoo Praha Czechia [email protected] Vogel Carine Sav'âne Switzerland [email protected] Voigtlaender-Schnabel Sophia Nuertingen-Geisslingen University Germany [email protected] Wieczorek Ellen Smithsonian Conservation Biology USA [email protected] Institute Yarnell Kelly Nottingham Trent University UK [email protected]

201

202

Author index

Briana ABRAHMS 93 Shannon E. ALBEKE 93 Brady W. ALLRED 160 Enrique ALONSO GARCIA 114 Ariel ALTMAN 58 Valentin AMRHEIN 140 Catherine J. ANGWIN 170 Rachael ANTWIS 42, 87 Russell AVRAMENKO 48

Lkhagvasuren BADAMJAV 93 Shirli BAR-DAVID 36, 58 Zoltán BARTA 85, 130 Jitka BARTOŠOVÁ 62 Markus BATHEN 184 Nasanbat BATTOGTOKH 34 Jeffrey L. BECK 93, 99, 101, 160, 170, 173, 180 Albano BEJA-PEREIRA 29 Afework BEKELE 140 Roni BELENKI 193 Jennifer BELLAW 48 Tamar BEN-NUN 58 David BERMAN 112 Anna BERNÁTKOVÁ 188 Alexandre R. BEVINGTON 160 Yash Veer BHATNAGAR 71 Miroslav BOBEK 75 Coby BOLGER WILSON 52, 114 Randall B. BOONE 93 Amos BOUSKILA 36, 58, 172, 193 Adéla BOUŠKOVÁ 122 Paul BOYCE 144 Samantha BREMNER-HARRISON 64 Dafna BRICMAN 193

203

Jake BRITNELL 87 Lauren BRIZGYS 50 Martina BURNIK ŠTURM 32 Bayarbaatar BUUVEIBAATAR 92, 93

Francisco CEACERO 34, 122, 136, 188 Simon CHAMAILLE-JAMMES 93 Cara CLANCY 146 Justine CORDINGLEY 77 Paul C. CROSS 93 Luděk ČULÍK 118 Gábor CZIRJÁK 190

Hannah DAVIE 64 Eric DAVIS 50 Vitalii DEMESHKANT 164, 165 Helmut DENZAU 126 Barbora DOBIÁŠOVÁ 75, 118 Usukhjargal DORJ 38, 93, 150 Florian DROUARD 152

Lars S. EKERNAS 97, 108 David M. ENGLE 160 Saeideh ESMAEILI 67, 93, 138 Paul H. EVANGELISTA 140

Jaime FAGÚNDEZ 40, 175 Claudia FEH 64, 152 Jennifer FIKE 108, 156 Ilya R. FISCHHOFF 93 Adam T. FORD 93 Samuel D. FUHLENDORF 160 Sheila FUNNELL 77

Mariana GAMIERO 146 Stefan GAVRILIUC 48 Marten GEERTSEMA 160 Jean-François GERARD 36 Abhishek GHOSHAL 71 Jennifer GIANDHARI 87 John GILLEARD 48

204

Nina GIOTTO 36 Clark Kendrick C. GO 134 Peter J. P. GOGAN 81 Jacob R. GOHEEN 93, 136 Erin GOODRICH 54 Morris GOSLING 26 Kayla GRAMS 106 Gili GREENBAUM 58 Alex GREENWOOD 190 Mark GREGORY 160 Theo GRUENTJENS 184 Cassandra GUARINO 54 Luna GUTIÉRREZ-CEPEDA 52

Carol HALL 64 Dennis HANSEN 172 Julia HATZL 172 Mahmoud-Reza HEMAMI 93, 136 Enno HEMPEL 184 Jacob D. HENNIG 93, 99, 101, 160, 173, 180 Peter HEROLD 184 Danielle HINCHCLIFFE 42, 153 Satoshi HIRATA 124, 128, 132, 148, 155

Benjamin IBLER 140 Kazushi IKEDA 134 Sota INOUE 134, 148, 155

Enkhbayar JAGDAG 38, 150 Emily JENKINS 48 Kate JENKS 38, 93 Sara JERELE 50 Brett R. JESMER 93, 136 Tomáš JIRÁSEK 118 Maggie M. JONES 105, 192

Timothy KAARIA 79 Petra KACZENSKY 32, 73, 93, 138 Shirin KARRYEVA 73 Fanuel KEBEDE 140 Viola KEREKES 85, 130

205

Graham KERLEY 87 David KIMITI 79 Sarah R. B. KING 56, 62, 67, 93, 99, 108, 156, 180 Daniel KLICH 165 Martina KOMÁRKOVÁ 122, 136, 188 Konstanze KRUEGER 184 Takatomi KUBO 134 Ralph KUEHN 73

Joshua LABARAKWE 77 Laura LAGOS 40, 175 Jess LEA 42 Corey LEE 87 Peter LEIMGRUBER 38 Sonja E. R. LEVERKUS 160 Wayne LINKLATER 27 John D. C. LINNELL 73 Badamjav LKHAGVASUREN 92 Cornelius J. LOUW 46 Lilla LOVÁSZ 140 Belinda LOW 77

Tamao MAEDA 124 Samantha MAPES 50 Jan MAREK 75 Elena MARIOTTI 46 Jason P. MARSHAL 46 Hanne MARTIN 184 John McEVOY 38, 93 Amy McLEAN 50, 54 Philip D. McLOUGHLIN 48, 83, 144 Aránzazu MEANA 52 Lubomír MELICHAR 118 Jorg MELZHEIMER 93 Renata S. MENDONÇA 134, 148, 155 Jerod A. MERKLE 93 Terry A. MESSMER 110 Patricia D. MOEHLMAN 44, 69, 140 Redwan MOHAMMED 140 Rita MONTEIRO 29 Thomas MORRISON 190

206

Thomas MUELLER 93 Byamba MUNKHTUYA 152 Carlos MUÑOZ-BARCIA 40, 175

Máté NAGY 130 Dejid NANDINTSETSEG 38, 93 Francisco Javier NAVAS GONZALEZ 50 Juán José NEGRO 172, 193 Gertrud NEUMANN-DENZAU 126 Martin NIELSEN 48 Haniyeh NOWZARI 158 Cassandra M.V. NUÑEZ 105, 192

Colleen O’BRIEN 177 Sakiho OCHI 124, 128 Tulio de OLIVERIA 87 Kirk OLSON 93 Alix ORTEGA 190 Norman OWEN-SMITH 69 Sara OYLER-McCANCE 108, 156 Ganbaatar OYUNSAIKHAN 32, 75, 188 Katalin OZOGÁNY 85, 130

Francesca PARRINI 46, 69 Steven L. PETERSEN 178 Sureshnee PILLAY 87 Pandora PINTO 132, 148 Jan PLUHÁČEK 62, 118 Jocelyn POISSANT 48, 83 Budhan S. PUKAZHENTHI 103 Nicola PUSTERLA 50

Zoe RAW 186 Libby REDMAN 48 Lucy REES 60, 114 Charlotte REGAN 83 Sharon RENAN 58 Eloy REVILLA 193 Monamie RINGHOFER 124, 134, 148, 155 Jan ROBOVSKÝ 118 Ovidiu ROȘU 168

207

Daniel I. RUBENSTEIN 66, 93, 133 Eldar RUSTAMOV 73 Allen T. RUTBERG 106

Mikhail V. SABLIN 32 Albert SALEMGAREYEV 73 Ravchig SAMIYA 32 Kateřina ŠANDLOVÁ 136 Istvan SANDOR 85 Radka ŠÁROVÁ 62 Hall SAWYER 93 John D. SCASTA 89, 93, 99, 101, 160, 170, 173, 180 Kathryn A. SCHOENECKER 56, 67, 93, 97, 99, 108, 156, 180 Peter SCHUETTE 184 Peter A. SEEBER 190 Susanne SHULTZ 42, 87, 153 Jaroslav ŠIMEK 75, 188 Ronald O. SKOOG 81 Kateryna SLIVINSKA 164, 165 Steve SMITH 29 Ernst-Hermann SOLMSEN 184 Melissa SONGER 38, 93 Natalia N. SPASSKAYA 32, 162 Jared A. STABACH 38, 93 Christina STANLEY 42 Seth STAPLETON 93 Theresa STRATMANN 93 Sajid SULTAN 71 Siva R. SUNDARESAN 93

Ito Y. TAKEHIKO 93 Alan TEMPELTON 58 Redae T. TESFAI 44, 69 Dashpurev TSERENDELEG 150 Vladimíra TUČKOVÁ 62 Elke TUKKER 182 John W. TURNER Jr. 106

Sophia VOIGTLAENDER-SCHNABEL 184 Roman VODIČKA 75 Christian C. VOIGT 32

208

Linda VOGEL 184

Chris WALZER 138 Annsarah WANGUI 77 Sarah WIEZOREK 184 Matthew WORKENTINE 48

Shinya YAMAMOTO 124, 134, 155 Nataliya YASYNETSKA 165 Nicholas E. YOUNG 140

Alon ZIV 36 Steffen ZUTHER 73

209

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