REVIEW OF FERAL HORSES

A REVIEW OF FERAL HORSE RESEARCH PERTINENT TO THE REINTRODUCTION OF PRZEWALSKI'S HORSES

Richard Miller

HABITAT USE

In the United States, Canada and Australia feral horses are most often considered to be important as competitors with the livestock and wildlife. Hence, much of the research is directed toward understanding and reducing this competition. There is a volume of feral horse food habits studies (Hubbard and Hansen 1976; Varva and Sneva 1978; Salter 1978; Salter and Hudson 1979; Hanley and Hanley 1982; Krysl et al. 1984a, 1984b). These studies have found feral horses are primarily dependent on grasses but do use some forbs and shrubs depending on the habitat where they are feeding.

Habitat use by feral horses has also been well studied in the attempt to understand competition between horses and other animals. The availability of water is important in determining the distribution of feral horses (Pellegrini 1971; Welsh 1975; Hansen 1976; Keipner 1976; Green and Green 1977; Miller 1983a). Where the water supply is limited, intensive competition for water may occur. In the Red Desert, when water was scarce, most horses (60 to 100% of all locations) were within 4.8 kilometres of water (Miller 1983a). The same study found temperatures are the most important factor influencing use of free water.

Shelter, particularly wind cover, is important to feral horses during severe winter weather (Welsh 1975; Salter and Hudson 1978; Miller 1983a). The areas used for wind cover are often the lea side of ridges, which are also used for feeding if the wind keeps them snow free. Eighty to ninety percent of horse locations in the Red Desert during winter were within 1.6 kilometres of ridges. Duncan (1983) found wind blown sites were used to avoid biting insects in summer. Selection of protected micro climates in winter may be an energy conservation mechanism (Malechek and Smith 1976).

Feral horses select different vegetation types by season (Welsh 1975; Salter and Hudson 1978; Duncan 1983; Miller 1983a). This selection may be related to diet, availability of water and cover, or as Duncan (1983) found, concentrations of high quality foods.

HOME RANGE AND MOVEMENTS

Feral horse home range size and movement patterns appear to be determined by the distribution of resources needed by the horses (Miller 1983b). Home range sizes have been reported (ranging from 90 ha (Welsh 1975) to 30 300 ha (Miller 1983b). The size of the home range extent of movements both daily and seasonal reflect the needs for food, cover and water previously discussed.

Ideas from several theoretical ecologists can be used to predict the home range size and movement patterns to be expected in a reserve for Przewalski horses: Clutton-Brook and Harvey (1977 and 1978) have suggested large home ranges result where resources are strongly clumped, widely dispersed or unpredictable in abundance. Altmann (1974) and Crook et al. (1976) predicted extensive home range overlap where essential resources have very restricted distributions. Denham (1971) argued clumping of populations should be high where predictability of resources is low.

A variety of movement patterns and home range sizes have been shown or suggested for feral horses (Pellegrini 1971; Welsh 1975; Berger 1977; Green an Green 1977; Miller 1983b). The patterns observed seem to fit those predicted by theory. The movements and the size of the use area for reintroduced Przewalski's horses may be determined by the distribution and predictability of resources in the release area.

SOCIAL ORGANIZATION

Like the home range data, social organization and behaviour of feral horses have also varied a great deal from site.

Feral horses social organization are generally variations of the family groups described by Klingel (1974) for Plains Zebra. The basic structure is bands made up of one mature stallion and several mares with their offspring. Groups of bachelor males are also found in the vicinity of the breeding bands. This basic type of social organization is widely found in feral horses (Pellegrini 1971; Hall and Kirkpatrick 1975; Feist and McCullough 1976; Berger 1977; Green and Green 1977; Salter 1978; Miller 1980). However, variations from this basic pattern are also common. Breeding bands containing more than one mature male are found in many areas (Hall 1972; Welsh 1975; Keipner 1976; Salter 1978; Miller 1980). Pellegrini (1971) and Berger (1977) reported bands of horses rarely associated with each other. Pellegrini (1971) suggested that horse bands in the Wassuk Mountains of Nevada might be territorial. All other studies have found that bands frequently associate with one another and are non territorial. Dominance hierarchies are reported within bands for males (Hall and Kirkpatrick 1975; Welsh 1975; Green and Green 1977; Salter 1979; Miller 1981) and for females and young (Boyd 1980). Interband dominance hierarchies are found in feral horses in the Red Desert which determined access to a limited water source (Berger 1977; Miller and Denniston 1979).

The adult membership of feral horse bands is generally quite stable averaging 0.75 adult changes per year in the Red Desert (Miller 1980) or 11.6 adult changes/100 horses/year in Nevada (Feist and McCullough 1976).

Breeding bands vary widely in size from two horses to more than twenty horses.

Breeding bands containing multiple adult males are more stable and larger than single male bands (Miller 1980).

Rates of exchange between bands, average band size and proportion of multiple male bands can change from year to year (Miller 1980).

A social unit made up of breeding bands and bachelor male groups has been suggested by Miller and Denniston (1979) and Miller (1983b). It was defined as follows: “a herd is a structured social unit made up of bands following similar movement patterns with a common home range”. An examination of data presented by Green and Green (1977) and Feist and McCullough (1976) also suggests the existence of herds.

Like home range and movement patterns social organization may be a result of the action of resource distribution on the inherent behaviour patterns. The oretical ecologists have explained both large band size (Altmann 1974; Clutton-Brock and Harvey 1977) and multiple male bands Bourliere and Hadley 1970) in other animals as a reponse to clumped and/or separate resource distribution.

BREEDING BEHAVIOUR

In single male bands the band stallion does the majority if not all breeding (Feist and McCullough 1976; Hall and Kirkpatrick 1975; Salter 1978; Miller 1981). Band stallions from single male bands also breed females in other bands (Miller 1981) sometimes with great frequency (9 of 19 copulations, Nelson 1978). In multiple male bands dominant males may be doing all the breeding (Welsh 1975) or the majority of the breeding (Salter 1978; Nelson 1978; Miller 1981). Within multiple male bands at least three breeding systems are found: (i) several stallions breeding with one mare, all from the band; (ii) consort pair formation; and (iii) the dominant stallion doing all or most of the breeding (Miller 1981). Additionally as in single male bands some breeding is done by males from outside the band. In the Red Desert 13 of 22 mares observed being bred more than once in a season were bred by two or more stallions (Miller 1981).

DEMOGRAPHICS

There has been considerable disagreement among researchers regarding rates of increase in feral horse populations. Field researchers have proposed rates of increase approaching 20% per year while population modelers have generally suggested much lower rates of increase. Conley (1979) reported the conditions necessary for rates of increase approached 20%: (i) the proportion breeding must be 0.8 or greater across all age classes; (ii) age at first breed must be 3 years or younger; and (iii) breeding must extend to age 8 or 10. Additionally adult mortality rates must be very low. Boyd (1980) found 53 to 78% of mares 3 years old and older foaled, and mares were foaling first at 3 years. Seal and Plotka (1983) found pregnancy rates in feral horses in Idaho to range from 35% in 2 year olds to 85% in 6 to 15 year old mares. Lactation rates in that study were 25% for three year olds and 50 to 100% in mares 5 years old and older. Eberhardt et al. (1982) documented populations in which apparent rates of increase were 18% and 20%.

The best available information seems to indicate that in populations with adult survival rates around 80%/year and less than 80% of mares three years and older foaling each year, rates of increase may be between 5% and 10% per year. However, if adult survival approaches 95% and 80% or more of 3 year old and older mares foal rates of increase may reach 20% per year. If young horses are used in the Przewalski horse reintroduction, and if habitat or weather conditions do not limit reproduction, rates of increase may approach 20% until adult mortality rates increase with old age.

CONCLUSIONS

All aspects of feral horse behaviour and ecology which have been studied have varied from site to site. It is reasonable to expect the characteristics of the release rate such as the availability of forage, cover and water to influence the movements, behaviour and perhaps the social organization of reintroduced horses. Habitat and weather conditions were not severely limiting. High rates of increase may occur often on initial adjustment period.

REFERENCES

Altmann S.A. 1974 Baboons, space, time and energy. Amer. Zool. 14(1):221– 248.

Berger J. 1977 Organizational systems and dominance in feral horses in the Grand Canyon. Behav. Ecol. Sociobiol. 2:131–146.

Bourliere F. 1970 and Hadley M. The ecology of tropical savannas. Ann Rev. Ecol. Syst. 1:125–152.

Boyd L.E. 1980 The natality, foal survivorship and mare-foal behavior of feral horses in Wyoming's Red Desert. MS Thesis. Univ. Wyoming, Laramie. 137 p.

Conley W. 1979 The potential for increase in horse and ass populations: A theoretical analysis. Proc. Symp. on Ecol. and Behav. of Feral Equcids. Univ. Wyoming, Laramie. pp. 221–234.

Clutton-Brook T.H. and Harvey P.H. 1977 Primate ecology and social organization. J. Zool. 183:1–39.

Clutton-Brook T.H. and Harvey P.H. 1978 Mammals, resources and reproductive strategies. Nature 273:191–195.

Crook J.H., Ellis J.E. and Gross-Custard J.D. 1976 Mammalian social systems: Structure and function. Anim. Behav. 24:261–274.

Denham W.W. 1971 Energy relations and some basic properties of primate social organization. Amer. Anthropologist 73:77–94.

Duncan P. 1983 Determinants of the use of habitat by horses in a Mediterranean wetland. J. Anim. Ecol. 52:93–109.

Eberhardt L.L., Majorowicz A.K. and Wilcox J.A. 1982 Apparent rates of increase for two feral horse herds. J. Wildl. Manage. 46(2): 367–374.

Feist J.D. and McCullough D.R. 1976 Behavior patterns and communication in feral horses. Z. Tierpsychol 41:337–371.

Green N.F. and Green H.D. 1977 Wild horse population of Stone Cabin valley, Nevada: A preliminary report. Proc. National Wild Horse Forum. Coop. Exten. Serv., Univ. Nevada, Reno. pp. 59–65.

Hall R. 1972 Wild horse: biology and alternatives for management. US Dept. Inter., Bureau of Land Management, Billings District. 117 p.

Hall R. and Kirkpatrick J.F. 1975 Biology of the Pryor Mountain wild horse. US Dept. Inter., Bureau of Land Management, Salt Lake City, Utah. Unpublished paper, 21 pp.

Hanley T.A. and Hanley K.A. 1982 Food resource partitioning by sympototric ungulates on Great Basin Range Land. J. Range Manage. 35(2): 152–158.

Hansen R.M. 1976 Foods of free-roaming horses in southern New Mexico. J. Range Manage. 29:347.

Hansen R.M., Clark R.C. and Lawhorn W. 1977 Foods of wild horses, deer and cattle in the Douglas Mountain area, Colorado. J. Range Manage. 30:116–118.

Hubbard R.E. and Hansen R.M. 1976 Diets of wild horses, cattle and muledeer in the Piceance Basin. Colorado, J. Range Manage. 29:389–392.

Keipner R.R. 1976 Social organization of feral ponies. Proc. Penn. Acad. Sci. 50:69–70.

Klingel H. 1974 A comparison of the social behavior of the Equidae. In: The Behavior of Ungulates and its Relation to Management. V. Geist and F. Walther (es). IUCN New Series No. 24, IUCN, Morges. pp. 124–132.

Krysl L.F., 1984a Hubbert M.E., Sowell B.F., Plumb G.E., Jewell T.K., Smith M.A. and Waggoner J.W. Horse and cattle grazing in the Wyoming Red Desert. I. Food habits and dietary overlap. J. Range Manage. 37(1):72–76.

Krysl L.F.,1984b Hubbert M.E., Sowell B.F., Plumb G.E., Jewell T.K., Smith M.A. and Waggoner J.W. Horse and cattle grazing in the Wyoming Red Desert. II. Dietary quality. J. Range Manage. 37(3):252.

Malechek J.C. and Smith B.M. 1976 Behavior of range cows in response to winter weather. J. Range Manage. 29:9–12.

Miller R. and Denniston II R.H. 1979 Interband dominance in feral horses. Z. Tierpsychol. 51:41–47.

Miller R. 1980 Band organization and stability in Red Desert feral horses. Proc. Conf. Ecology and Behavior of Feral Equids. R.H. Denniston (ed). Univ. Wyoming, Laramie. p. 113–123.

Miller R. 1981 Male aggression, dominance and breeding behavior in Red Desert feral horses. Z. Tierpsychol. 57:340–351.

Miller R. 1983a Habitat use of feral horses and cattle in Wyoming's Red Desert. J. Range Manage. 36(2):195–199.

Miller R. 1983b Habitat use of feral horses and cattle in Wyoming's Red Desert. J. Range Manage. 34:195–199.

Nelson K.J. 1978 On the question of male limited population growth in feral horses (Equus caballus). MS Thesis, New Mexico State Univ., Las Cruces.

Olsen F.W. and Hansen R.M. 1977 Food relations of wild free-roaming horses to livestock and big game, Red Desert, Wyoming. J. Range Manage. 30:17–20.

Pellegrini S.W. 1971 Home range, territoriality and movement patterns of wild horses in the Wassuck Range of western Nevada. MS Thesis, Univ. Nevada, Reno.

Salter R.E. 1978 Ecology of feral horses in western Alberta. MS Thesis, Univ. Alberta, Edmonton.

Salter R.E. and Hudson R.J. 1978 Habitat utilization by feral horses in western Alberta. Le Naturaliste Canadien 105:309–321.

Salter R.E. and Hudson R.J. 1979 Feeding ecology of feral horses in western Alberta. J. Range Manage. 32:221–225.

Seal and Plotka. 1983 Age-specific pregnancy rates in feral horses. J. Wildlife Manage. 47(2):422–429.

Vavra M. and Sneva F. 1978 Seasonal diets of five ungulates grazing the cold desert biome. Proc. Internat. Rangeland Conf. 1:435–437.

Welsh D.A. 1975 Population, behavioral and grazing ecology of the horses of Sable Island, Nova Scotia. PhD Thesis, Dalhousie Univ.

MUSTANGS OF THE MANYCH LAKE

V. Klimov and N. Paklina

Feral horses, mustangs … These words make anyone imagine blue boundless prairies with galloping free wild horses, riding cowboys and Indians. Mustangs are a part of the history of North American prairies and pioneering of the Wild West. They are newcomers to America for the second time. In the pleistocene era, it was from America that the ancestors of horses, asses and zebras crossed the Bering Straits in several streams and populated Europe, Asia and Africa. Then they became extinct in their native land and returned only in the latter centuries, accompanying the whites in developing the vast territories of North America and the natives. Having played a great role in the pioneering of America, the horses gradually lost their former significance and fell into the background, failing to compete with motor vehicles. They were set free and allowed to be on their own. That is how mustangs appeared in America.

Feral horses not only live in the prairies of America but also in the USSR. A few small populations are scattered over the areas, difficult to access. Such a population has existed in one of the islands of the Manych-Gudilo or Great Manych lake (region of the city of Rostov-on-Don) for about 30 years.

The island Yuzhny or Vodny, the topic of our talk, is the largest on the lake. It is 12–13 km long and 2–4 km wide, with the total area being 2658 ha. The island stretches from west to east, its southern shore is steep, while the northern one is sloping. From west to east the island is cut out up to the centre with a bay; the southern shore, being a water-meadow, consists of sand-bars, small islands, reedy bays and serves as a refuge for numerous birds herons, stints, swans, ibises, etc. A great number of grass associations are represented in the island. Moderately dry sod-wheat grass steppe covered with sheep's fescue and feather grass prevails. There is a great variety of meadows on the island. Such species rare for the Rostov Region as splendid meadow saffron (Colchicum), tulips (Tulipa bibersteinniana and Tulipa shrenkiana), licorice and feather-grass (Stipa zalesskiana) are preserved there.

But the main beauty and boast of the island are feral horses. They were experimentally released into the island in the middle of the fifties. The horses survived and reproduced. In some years they numbered 80 individuals. Last summer, when these observations were undertaken, there were about 40 horses: 12 stallions, 14 mares and 14 foals. Regarding their exterior aspect, these are sturdy red and brown horses of the Budyonny breed. The horses were named at the beginning of the observations.

The island herd consists of three bands, i.e. harems, two bachelor groups and four solitary bachelors. The largest band is led by Bury. It includes 5–7 mature mares with young stock. Bury is a sturdy dark-brown stallion of 7–10 years of age. He is an initiator and winner of all ritual contacts in the island. These contacts do not usually turn into real fights since his authority is rather high. For a long period of time Bury had tolerated the claims to superiority in the band by a two-year stallion Adyutant whom he banished later on; he also tolerated a small band of Ryzhy consisting of two mares and a yearling who constantly ranged on Bury's territory. The bands met on grazing land and at watering sites. But in every case the satellite showed respect for the band stallion, and Bury was patient with the presence of the satellite. It is worth speaking in detail of Bury's group since it is the largest band on the island. The first assistant of Bury in guiding the band was Gryaznulya, a massive light-brown mare. She led the band in all the cases when it was necessary to change rangeland or go to the watering site. In doing so, she asked for permission from Bury who himself decided the usefulness of the band's moves and actions. Varvara is a mature red mare and the stallion's favourite lady who enjoyed special privileges. Stesha is a friend of Varvara and due to this fact she had certain privileges in the band. Schuka is an old hairy mare and a regular initiator of quarrels with other horses. She was inclined to make friends with Gryaznulya and Sivka-Burka who had two foals: a newborn and a yearling. This fact explains the attraction that Sivka-Burka drew from other mares who spent a lot of time petting the foals. A mare Chernoburka played a specific role in the band. She made no friends with the others and devoted all her time to her foal. Why was Adyutant banished from the band? In moving in the rangeland he tried to lead the band, guide the juniors, smell the excrements. Since such behaviour characterizes a sexually-mature stallion, this proved his claims for the territory and leadership in the band. In the middle of May Bury banished Adyutant from the group. Analysing the relationships in this group, it can be said that all the threads of superiority, leadership and guidance of the band are within the power of the leader that is a band stallion. As mentioned above, among mares an indisputable leader was Gryaznulya, and the band stallion did not interfere with her attempted leadership. A certain agreement was observed in the actions of these two animals: the mare led the band while the stallion was urging the others, she tried to lead the band but he did not want to move, and the mare returned. When one of them came to a stop, the other also stopped. So, she is a business lady but not a lady of the heart.

The other group is a band led by Pogorelets.

This mature red stallion was named after a small island located near the northwestern shore of the island that was called Gorely (burnt) after a steppe fire.

His band included three mature mares, four two-year foals and two sucklings. Beloglazka, a mature red mare with a star, had a one-year daughter. Polnoch, a mature light brown mare, had a small foal, and Taina, a dark brown female, took care of a newborn foal. In this group Beloglazka was a leader in all the movements while Pogorelets drove the group from the rear, marked the territory, protected the band. He was on friendly terms with one-year Zolushka, Polnoch's daughter, and Kometa, Beloglazka's daughter. These three animals constantly kept together and followed each other. Pogorelets like Bury had his satellites. This was a group of Gomon. It occupied the same territory and accompanied Pogorelets' band as his subordinates. Gomon, a mature dark red stallion, had two mature mares in the group: Bolshaya Medveditsa and Malaya Medveditsa.

Gorely island was also occupied by a bachelor group consisting of Ilya, Dobrynya and Alyosha. A dark brown stallion Ilya is a senior in this group. He marked the territory, fought with stallions from other groups. The other two were 2–3 years old. These three stallions were in Pogorelets' territory though they kept a distance necessary for peace and did not claim for his mares but, when water in the Manych lake reached critical salinity, they followed Pogorelets' band to the rangeland where a fresh water source was available.

In addition to group bachelors, the island housed solitary stallions. Dik, Kavaler and Stas held aloof from other horses.

It is interesting to analyse the way the territory was divided between the horses' groups. As would be expected, the strongest bands occupied the best portions of rangeland on the island. Bury's band settled on the most remote, western portion where sheep did not graze. This is a grassy water-meadow with fresh water puddles and previous year's haycocks. Pogorelets' band also occupied a grassy rangeland far from people. Other groups and solitary bachelors found themselves under worse conditions. They had to range in the areas where sheep were grazing.

The largest area was occupied by Bury's band, stretching from the western shore inside the island by 2 km. In the daytime the band used to stay on the water-meadow, resting and drinking. When it was getting dark, the leader took the horses to the higher southern shore. The view was better here (horses see very well at night), south winds in the spring and summer blew midges off. In spring the horses ranged in dusk hours and in summer they foraged throughout the night, resting in the daytime only. One of the sand-bars was their resting and watering place. In the hot season they stayed with their bellies in the water for a long period of time to protect themselves from heat and midges. When fresh-water puddles near the resting site were dried up, the leader took his band to the Sheep Water Source. All the family territory is spotted with numerous paths that connect sites of resting, ranging and watering. The paths go along the sand-bars and reeds, cross bivouac and ranging sites. In some places heaps of excrement can be seen on the paths that are regularly replenished. These are so-called horses' toilets that also serve as markers to show that the territory belongs to a particular group. In addition to the common toilets, the stallion usually has his own markers and patrol paths that go along the borders of the territory and are marked with excrements. Bury uses an earth road laid out by mowers. Only a mature stallion included in the population's hierarchial structure has the right to leave his patrol marks. Young stallions and mares leave their excrements in the common toilets or in the grass near the road but never on the road that is a patrol path of the stallion. The observations helped detect other bands' ranging and watering sites, directions of moves, resting places in the daytime and at night.

Comparing the use of territory by horses' groups with that by solitary stallions, one could indicate that family groups (bands) get acclimatized to their territories more properly. They have fixed resting, watering, toilet sites, etc. They move along permanent paths, mark and patrol their territory. Naturally, the bachelor groups and solitary stallions use rangelands which are much worse and they patrol their territory from time to time.

THE EDITOR'S COMMENTS: The article covers rather valuable evidence on behaviour of feral horses living in one of the Manych-Gudilo lake islands. The observations undertaken are of scientific and practical value for productive distant-pasture horse rearing involving all-the-year round ranging of horses. They prove once more that the horses can be preserved as a zoological species without man's intervention. Besides, it is possible to make use of remote rangelands with the help of horses for man's benefit - the production of the cheapest meat.

This article was published in ISSN 0023–3285 Horse Breeding and Equestrian Sport, Soviet monthly magazine No. 4. 1985.

TECHNIQUES FOR REINTRODUCTION

TECHNIQUES FOR RE-INTRODUCTION OF EQUUS PRZEWALSKI

J.M. Knowles

There are few precedents for the re-introduction of wild animals into their former habitats. Those that have taken place to date with large ungulates have been in warm climates in desert or semi desert. The re-introduction of the Arabian Oryx (Oryx leucoryx) into Oman, and the introduction of this species and Scimitar horned oryx (Oryx dammah) and Addax (Addax nasomaculatus) into the Hait Bar reserves of Israel offer limited guidance, as may the so-called semi reserves established to date in Europe for Equus przewalski. The experience of the management of the large herds at Askania Nova, in extensive conditions, will be invaluable to the project.

Because of this paucity of historical experience, it is most fortunate that we have the feral horse examples from North America and elsewhere from which information may be gathered for comparison with records of Przewalski Horses in those zoos where the animals live in social groups as would be expected in the wild, even though movements in and out of the herds are constrained by barriers.

There will, therefore, be some guidelines of value to our planning, but all concerned with this most exciting project should be aware that we are at a frontier of knowledge.

What follows therefore cannot be a scientifically developed plan, but rather a series of suggestions based on assumptions. As sites have yet to be chosen and a budget agreed and allocated, the field of assumption has to include the following:

1. Re-introduction sites will have similar climatic extremes and vegetation resources as those of the last known habitat of Przewalski horses - the Takhin Shara-nur mountains.

2. Finance on an adequate scale will be available for the potentially high costs of the necessary fencing and buildings for:

   1. Horses (in the initial stages)
   2. Staff accommodation
   3. Laboratory
   4. Feed and equipment storage

   Similarly that funding will be available for technical and non-technical personnel, transport to the various site stages and the means of monitoring the various project stages. It is not a function of this paper to discuss the question of finance, though it will be fundamental to success. A commitment of funds for at least 25 years will be necessary to ensure success of the project and the detailed monitoring needed to validate scientifically its problems and successes.

Aspects of re-introduction are discussed under appropriate headings below.

VETERINARY

As the constituent animals of release groups will come from diverse breeding centres it will be necessary to have a very full understanding of their health status, particularly in terms of immunology. Insofar as veterinary science can achieve this, it will be necessary to ensure that none of the constituent animals represent a disease threat to others, or is itself threatened by others. Similar knowledge of the health status of any domesticated or feral equines which may have direct or indirect contact with animals in the various stages of this programme will be vital. More easily accomplished but equally vital will be thorough physical examinations of all participant horses before they leave their original home and on arrival at the various “staging posts” in the programme. The stages of reintroduction for zoo-bred animals will be individually rigorous and will call for the same degree of veterinary care as that given by medical practitioners to athletes.

CHOICE OF PARTICIPATING ANIMALS

Genetic planning will play an important part in these choices, and this will be discussed in other papers. There will be some important additional considerations to those and to the veterinary aspects already referred to.

Broadly, these are temperament, which must not be either timid or excessively aggressive, and social experience (having lived in a stable mixed age and sex herd will be an advantage). History of ancestors will be a further consideration with physical, physiological, psychological and behavioural abnormalities being carefully scrutinized for possible genetic origins. One to two years of age would seem to be best for animals entering the programme.

RE-INTRODUCTIN BY STAGES

All this is of course subject to the limitations referred to at the beginning of this paper.

Stage 1

Where possible, bearing in mind international health regulations and geography, animals for the programme should be grouped “en route” and given time to develop social bonds before onward transit. This will facilitate transport, which may be lengthy, in twos or threes. Time should also be given to small constituent groups to be accustomed to their travel containers and be prepared for any significant change of diet.

Stage 2

Arrival at the chosen sites. At least 2 of these should be constructed at strategic points on the edge of the chosen range and should at minimum be out of sight and hearing distance of each other. Each site should have basically zoo conditions for the acclimatizing of the animals. These should consist of a partially open fronted shed capable of protecting the animals from the worst climatic extremes and be large enough to avoid overcrowding of temporarily incompatible individuals and consequent damage to each other. This house would have direct access to isolation stalls for the variety of purposes for which animals may need to be separated from the main herd. These should be at a distance from the main group as the herd instinct in Przewalski horses is stronger than with most other wild equines and isolation is consequently more difficult. The house would open on to a “yard” or “hard standing” which is well drained and has an abrasive surface to avoid hoof troubles during confinement in the early stages of re-introduction programmes. Continuous water supply and food distribution points sufficiently separated to avoid damaging agression at feeding times will be needed. The above described complex should open to an enclosure containing the kind of habitat that the animals will eventually find themselves living in. This enclosure will serve two important functions in addition to habituating the animals to new food plants. The first of these will be a physical toughening process for more stringent exercise than will have been experienced during the times of assembling, transport and settling in and (in the case of some individuals) than they have experienced in their place of origin. The second will be a period of further social adjustment.

Suggested sites for the enclosure would be approximately 100 to 200 hectares (depending on group size). This will be large enough to meet the requirements listed above and yet small enough to enable observation. In the context of observation clear freeze branding of all individuals is recommended so that intensive observation may be carried out from an observation tower with the aid of binoculars. Observation will be assisted by establishing a routine of returning to house and yard daily for supplementary feeding.

Stage 3

The stage (2) complex should lead to a large fenced area where the horses can be released after the criteria suggested below have been met. Here the animals will live as nearly naturally as possible but still be protected from contact with other equines (and consequently disease) and man, and can still be observed at least daily. Size of enclosure should be large enough to accommodate a self-sustaining herd, with opportunity for young males born in the group to form their own bachelor herds and stay in the enclosure. Another possibility is for 2 “bands” to be released into the same enclosure allowing for eventual social re-structuring in a natural way. Of additional importance to size of enclosure will be the nature of the terrain. Valleys and hills allowing visual escape of antagonistic individuals will be as important as space alone. A minimum of 20 000 hectares will be needed.

Stage 4

When the first three stages have been completed satisfactorily and the required knowledge accumulated, it should be possible to release entirely animals of the second generation into nature which will be free in every sense except that the animals will need to be guarded from human interference and that of domestic animals.

COMPOSITION OF GROUPS

If, as suggested above, two sets of facilities can be made available, it would be desirable to import an equal number of males and females and establish these in single sex groups, with ten animals (plus or minus 2) in each group. The females when they reach Stage (3) should be joined by a selected male for breeding to commence providing all has gone well with the preceding Stages and the mares have reached sexual maturity at this time. For this reason and one other it would be desirable for the male to be one year older than the females and arrive one year sooner. The second benefit of this arrangement is that valuable lessons may be learned from having the more expendable males testing the facilities and systems suggested. As near as possible all males should be of genetically desirable composition for mating with the females, so that eventual selection for the first “herd male” can be based on successful adaptation to the environment of the restoration site.

TIMING AND AGE OF ANIMALS

The least stress during transport and arrival would be in Spring when the animals would experience the least climate change. Animals should be approaching their second birthday at this time and therefore reaching the age of natural social adjustment.

LONG TERM

When the first animals reach Stage 3, a review of progress to date should be made and consideration given to the importation of further animals, which in their turn would follow the already suggested Stages. Changes in the methods used and the general plan will then be instituted in the light of previous experience.

It is impossible to anticipate how long after the beginning of the programme this review will take place, as this will depend on the successes and problems experienced.

DISCUSSION

The above suggestions represent a very preliminary view of how this exciting challenge may be approached. These are offered without the necessary detailed knowledge of the are and facilities available for the programme and without adequate knowledge of the climatic variations and their annual dates and duration.

Most importantly this paper has been attempted without knowing the ultimate stocking density of Equus przewalskii which the chosen site or sites may be able to sustain. It will be vital to establish at an early stage what our numerical goals will be, and in what time frame these should be set. The captive breeding community has successfully brought the Equus przewalskii from the point of near extinction to a population size which allows a restoration programme to be begun. A well-conceived and energetically pursued plan will receive the full support of those institutions which breed these animals, that of the wider community of conservationists and from the still larger part of the human population that are known as “horse lovers”.

TECHNIQUE FOR RESTORATION OF THE PRZEWALSKI HORSE IN THE WILD

C. Pohle

The Zoological Garden of Berlin welcomes and supports the project to restore the Przewalski Horse to its original native environment in the People's Democratic Republic of Mongolia, using animals from captive environments.

This restoration to the original environment is needed to conserve the performance of these wild horses, which is otherwise endangered and will alter after several decades of keeping these animals in zoo environments. As a result of good breeding results in the zoos, it is now possible to provide animals out of these descendants for the project. The Zoological Garden of Berlin is willing to take part in the project.

Younger animals up to an age of 5 years seem to be most suitable for adaptation in the wild. The transport to Mongolia could be done by having the Przewalski Horses packed in crates, and accompanied by a horseman who will be responsible for adequate feeding and watering of the animals on the long trip.

The Zoological Garden of Berlin has no experience in the restoration of horses. However, experiences with cattle, coming from an area without vegetation, into an area with natural vegetation, indicate that the animals born in captivity had problems in grazing at first. Although offered a rich source of natural feed, the animals got poorer. Therefore additional feeding for the restored animals must be provided for a lengthy period.

Depending on the number of animals available for the project, one or two female herds with one stallion each and a separate herd of young stallions should be established. Each group should be kept in large enclosed areas. The size of the area will depend on the location but should not be too small. A restoration into the wild should only be done when it is clear that the animals are able to thrive on naturally grown vegetation and when the herd structure has been established and strengthened. A most favourable time would be when the first foals are born in the enclosure where adaptation is taking place.

To keep the animals in the adaptation enclosure (or semi-wild enclosure) offers opportunity for regular care and necessary control which includes feeding and watering.

After liberation from the enclosure of adaptation, observation of the freed groups is important in order to obtain information about nutrition, movements and herd behaviour. It is necessary to make sure that the restored Przewalski Horses are not endangered by large wolf populations and that there is no possibility of contact between the liberated wild horses and domesticated horses.

TECHNIQUES USED TO REINTRODUCE UNGULATES IN ARIZONA WITH POSSIBLE APPLICATION FOR PRZEWALSKI HORSES

Richard Miller

The State of Arizona has been involved in ungulate reintroductions since 1913 to supplement or replace populations decimated by livestock disease and habitat destruction. Some techniques have been developed which may have application to the proposed Przewalski horse reintroduction.

A set of criteria which are used to evaluate potential release sites have proved valuable in avoiding costly mistakes. The criteria we use are:

1. Historic occurrence - Has the animal been found here in the past?

2. Land Status - Who controls use of the land?

3. Topography - Is the topography suitable to the animal?

4. Cover - What is the vegetative cover?

5. Range condition - What is the condition and trend for forage plants?

6. Presence of domestic animals - What domestic animals are present and are they competitors or carrying diseases to which the animal in vulnerable?

7. Presence of wildlife - What wildlife species are present and are they competitors or disease sources?

8. Human disturbance - What level of human use does the area receive?

9. Available water - What quantity and quality of water is available and at how many sites?

10. Dispersion and size of habitat area - What is the size of the available area? How are the resources such as food, cover and water dispersed? How much of the area will be unavailable due to topography, etc.?

11. Potential for expansion - Is there a potential to expand the area if the population does well?

12. Fencing - Are there potential problems with existing or planned fences?

13. Seasonal habitat - Are the seasonal needs of the animal such as wind cover, available?

The majority of our recent experience in Arizona has been in transplanting desert bighorn sheep (Ovis canadensis). Since 1979 we have reintroduced 384 bighorn to 28 separate sites. Along the way we have learned some lessons which may be useful in planning the Przewalski horse reintroductions. When we began reintroducing bighorn sheep we were concerned they might scatter widely or completely leave the area. First we built very large enclosures and left the sheep in them for up to fourteen years before releasing them. Mortality rates have been high and reproductive rates low while animals were held in enclosures. For example the Aravaipa herd increased from 16–22 sheep while held in an enclosure from 1958 to 1972. This herd was released in 1973 and by 1983 numbered about 100 sheep (Dodd 1983). Dodd (1983) also found these sheep failed to utilize all the area available to them and concentrated lambing sites near the old enclosures. He suggested that transplants be made with small groups of animals (6–10) at several locations, and that lambing sites be established by transplanting ewes late in their pregnancy. This technique has been used to supplement a declining sheep population with dramatic success. Sheep transplanted in late pregnancy are using an area approximately one quarter the size of sheep transplanted in winter and have scattered much less (Remington and deVos, in press). In almost all releases some bighorn rams have left the transplant site some travelling up to 75 kilometres (Dodd 1983; Miller and deVos, unpublished data).

The Arizona Game and Fish Department has also been actively transplanting javelina (Tayassa tajacu) moving over 400 animals (Day 1985). In order to promote herd formation we have been placing javelina into artificial herds and holding them together for a minimum of six months in enclosures before release. Although the effort is considered important because of low reproductive rates in small javelina herds, all released javelina herds have fragmented badly following release. No technique we have yet tried has solved the problem.

REFERENCES

Arizona Game and Fish Department. 1984 Draft Pronghorn Management Guidelines. Unpublished memograph.

Arizona Game and Fish Department. 1984 Draft Bighorn Management Guidelines. Unpublished memograph.

Day G.I. Javelina research and management in Arizona. 1985 Arizona Game and Fish Department, Phoenix, Arizona. 128 p.

Dodd N.L. 1983 Ideas and recommendations for maximizing desert bighorn transplant efforts. Desert Bighorn Council, 1983 Transactions. pp. 12–16.

Remington R. and deVos Jr. Preliminary request on Arizona's first desert bighorn sheep transplant into a natural population. Desert Bighorn Council, 1985 Transactions.

PRACTICAL EXPERIENCES IN IMMOBILIZING AND TRANSPORT OF PRZEWALSKI HORSES

H. Wiesner and G. von Hegel

1. IMMOBILIZATION

Since the introduction of the synthetic morphine derivate, Etorphine hydrochloride in zoo medicine the immobilization of specimens of the equine family has lost its horror. This very highly potent drug made obsolete all previous methods using the high risk drug Succinylcholin chlorid and the oral application of 30 g Chloralhydrat per animal. However whilst these old methods have only been used in emergency cases, they have afflicted a high risk on the live animals and the health of the personnel involved. Thus we have with the blowpipe and M99 a new method for a safe and effective immobilization of wild horses.

However, in using Etorphine alone like M99 crystal powder or even the combination of Etorphine with Acepromazin, registered as “Large Animal Immobilon^R” C-vet, England (1 ml contains 2.25 mg Etorphine per ml combined with 10 mg Acepromacin used as tranquillizer), we must await severe side-effects. About 6–10 minutes after the injection severe excitation, violent muscle-tremors, collapses, sweat spasms, cramps, blind forward moving and an increase of the pulse rate up to 200/minute were observed. These side-effects can be reduced with the combination of Xylazin or, in a short operation, by a direct intravenous injection of 1/10 of Diprenorphin of the Etorphine dosage (Wiesner, Rietschel and Gatesmann 1982).

With this combination we have had good results in the last years in immobilizing wild horses. From 1975 to 1984 we immobilized 188 times Przewalski horses without any mortality (Wiesner and von Hegel 1985). The recommended dosage for an adult Przewalski horse would be 2.5 ml of Immobilon^R combined with 50 mg of Xylazine in toto on average. That corresponds to a dosage of 0.019 mg/ka of Etorphine (Wiesner, Rietschel and Gatesmann 1982; Wiesner and von Hegel 1985). In some animals a higher dosage up to 3 ml of Immobilon^R may be necessary, depending not on the body weight but on the nervosity of the animal. Therefore, the dosage can vary individually. We have practised this immobilization not only in emergency gynaecological examinations of our Przewalski horses but also in other cases (Wiesner and Bostedt 1979). However, the immobilization stage of this combination is not sufficient for painful operations. When we had to perform a tail amputation in our mare “Sitka” (born 8.7.80, Studbook No. 932/Hell. 84) in the year 1984 after an injury from the stallion, we had to give some Immobilon^R (0.5 ml) by slow direct means intravenously to get surgical tolerance. No doubt, this tolerance can also be reached by an adequate i.v. injection of Ketamine.

The big advantage of this immobilization is the reversibility by i.v. injection of the antidote diprenorphine, registered as “Revivion” (1 ml contains 3 mg diprenorphine hydrochloride per ml).

Especially in the family of equinae, the so-called “hepatocyclic effect” may occur. In this case it means no metabolized Etorphine passes the liver via the bile and is reabsorbed in the duodenum. Therefore 6–10 hours after the injection of Etorphine heavy excitations may occur, that may lead to a self-injury risk of the animal. To avoid this side-effect, half the i.v. applied dosage is additionally given subcutaneously.

In the year 1984, a thesis was written about the change of the blood parameters during the combined immobilization with Immobilon^R and Rompun^R. During the experiment three blood samples were taken of 20 immobilized Przewalski wild horses every 10 minutes within a period of half an hour, starting 10 minutes after the Immobilon^R-Rompun^R injection (Kuttner 1985).

The samples were analysed for red and white blood cells, hematocrit hemoglobin, chloride, sodium, potassium, calcium and phosphorus, GOT, GPT, AP, LDH, CK, GLDH, G-GT and cholinesterase, bilirubin, creatinin, total protein, urea and glucose.

The results were statistically evaluated.

Changes in blood levels within 30 minutes after the narcotic injection were analysed. Basic data were the first samples, which were taken 10 minutes after injection.

Table 1 shows mean values and standard deviations of Przewalski horses 10 minutes after injection.

Thirty minutes after injection hemoglobin, erythrocytes, hematocrit, glucose and potassium changed significantly.

Glucose increased heavily, the other parameters were diminished.

Leucocytes, calcium, phosphorus, TP, GOT, GPT, AP, LDH and GLDH decreased not significantly but notably.

Bilirubin, creatinin, chloride, sodium, CK, G-GT, cholinesterase, MCV, MCH and MCHC remained nearly constant.

Urea was individually different, so a narcotic-caused change could not be observed.

It may be concluded that blood values taken under narcosis, which differe from reference values, did not change because of the narcotic drugs but on account of pathologic reasons as soon as they leave the physiologic variances. Excluded are the values of red blood cells, hemoglobin, hematocrit and glucose, which have changed significantly during the 30 minute experiment.

We have to point out the fact that Xylazin alone is not an adequate drug to get a sufficient immobilisation stage in the Przewalski horses, even if this drug is only administered to handle the animals for the transport. Two fatality cases with Xylazin first and a following injection of Etorphine are reported. The animals died by shock syndrome in the transport crates (Müller et al. 1980).

2. TRANSPORT

With a dosage of ca. 12 g Vetranquil Granulat^R (Fa. Albrecht) (i.e. ca. 220 mg Acepromacin per adult animal) Przewalski horses under zoo conditions can become sufficiently sedated to handle them easier for the transport.

After 10–20 minutes the horses show uncoordinated movements and are in most cases able to be driven into the transport car or crate.

Nevertheless, like all oral drug intake, the reaction of the animals can vary considerably and the Acepromacin should not be used in very shy and timid animals. In these animals, the darting with Immobilon^R/Rompun^R-combination shows greater advantage and preserves the animals from self-injuries. For this matter, the animals become handable, before the complete immobilization of the Etorphine takes effect. The stiff-legged animals can be quite easily led into a crate, where the antidote injection is given (Heck 1980). The crate should be narrow enough that the animal cannot turn during the transport and broad enough to be comfortable for the animal. A sufficient air conditions especially at the ground of the crate is absolutely necessary. If the animals can be transported by car and not by aircraft the transport without crates in a convenient lorry for horse transport is preferable. According to our experience the animals will remain much calmer in these special transport vehicles than in the crates. For longer distances in the crates, water and food supply should be arranged regularly. In all types of transport only animals in a standing position should be moved.

The animals should starve the day before transportation, even if they are not fully immobilized but only slightly sedated. Chasing wild horses with the object of capturing or immobilizing or netting can easily lead to myoglobinurie of the captive animal with fatal results. In any case, a safe immobilization has the preference to all other handling methods.

For that reason, the staff of the re-establishment programme of Przewalski horses should be specially trained for the immobilization of these animals and a blow pipe and blow gun equipment seems to us to be as necessary as the use of Etorphine and Xylazin, at least during the first period in the arger enclosures before the horses can be successfully released in the wild.

REFERENCES

Heck H. 1980 Use of M-99 on wild equines. Equus, Band 2, Heft 1, 132–133.

Kuttner C. 1985 Veränderungen von Blutwerten von Wildequiden während der Immobilon ^R Narkose. Inaugural Dissertation, München.

Müller R., Rüedi D., Schatzmann U. and Sägesser H. 1980 Transportzwischenfälle bei Przewalski-Pferden. Equus, Band 2, Heft 1, 79–81.

Wiesner H. and Bostedt H. 1979 Zur Sterilitätsbehandlung beim Przewalski-Pferd. Zschr. Kölner Zoo, Heft 2, 22 Jahrg. 55–58.

Wiesner H. and von Hegel G. 1985 Praktische Hinweise zur Immobilisation von Wild- une Zootieren. Tierärztl. Prax 13:113–127.

Wiesner H., Rietschel W. and Gatesmann T. 1982 Erfahrungen mit der Kombination von Immobilon^R und Rompun^R beim Zootier. Zschr. Kölner Zoo, Heft 2, 25 Jahrg. 47–55.

ECOLOGICAL CHARACTERISTICS OF THE PRZEWALSKI HORSE AND THE PROSPECTS FOR ITS ADAPTATION IN THE WILD

V.V. Klimov

SUMMARY

The social and spatial phenotype of the band of Przewalski horses in Askania Nova is a functional biological system with a hierarchic type of structure, having relatively great diversity in its composition and displaying the strategy of social formation in a partially protected territory. The combination of such strategies, with their cooperating and coordinating links, allows the system to respond elastically to the fluctuations of the outer environment and to adapt to it successfully ensuring its survival. At the same time, some elements of this system, when existing separately, are not able to provide the adequate response to the environment and to withstand the external factors.

Therefore it is necessary to form full-value social groups when introducing the horses into the wild: such groups will ensure the regulation of the intra-band relations, hereditary formation of agonistic and reproductive behaviour patterns, the establishment of a specific structure and organization assisting in responding adequately to the environment and to the further propagation of the species. Such groups shall be formed by themselves, on the basis of personal sympathies and abilities of stallions in the territories able to carry several functional units, i.e. harems, bachelor groups, etc. In these conditions, under natural selection, the natural mechanisms of improving the consequences of inbreeding will be developed, such as reproductive success of more experienced and strong stallions, rotation of males and females, formation of new harems.

All activities on the creation of horse groups should be carried out beyond the active reproduction period, i.e. in late summer or in the fall. Sexual activity is reduced at this time; the sired mares reject stallions and the yound stock that were born in the spring grow older. When forming the bands and shipping the horses to other regions, the foals should be kept with their mothers. Moreover, the horses should be transported by harem groups. When the horses have to be transported in cages, the presence of familiar animals that are sensed acoustically, or by smell, will allow them to tolerate better the extreme conditions and avoid stress.

It should be kept in mind that when Przewalski horses are introduced into the natural biotopes, the presence of the domestic Mongolian horse in these regions will lead to the formation of mixed harems and to cross-breeding. In order to avoid this it is necessary to remove the domestic horses from the introduction areas, preferably in the spring, during the active reproduction period. In wintertime, the domestic horses may be of great help, teaching “the newcomers” to get food from under the snow cover. In this situation they will serve as ethological “mentors”.

Thus, there is every reason to assume that the reintroduction of wild horses into nature will be a success if they successfully pass the period of primary adaptation to the climatic conditions of the steppe area in Askania Nova or in some other steppe reserve in the territory of the Soviet Union.

The study has been made of Przewalski horse behaviour in the steppe reserves of Askania Nova from the aspect of its follow-up reintroduction into the wild. Taking into account the major problem, namely whether the horses will manage to withstand the environmental pressure in the severe conditions of Central Asia, the study was focused on individual and social behaviour of the free-ranging populations of horses. The adequate response of the wild horses to environmental fluctuations can assure that their reintroduction into the wild will be a success. The study was carried out in 1980–1984. The band of Przewalski horses is maintained year-round in large enclosures of Stipe and Festuca steppe with the area ranging from 45 to 1550 hectares. In these conditions the horses are under the effect of abiotic factors of the environment (climate, relief), biotic (other animal species, steppe associations of plants) and of man-induced factors (automobiles, aircraft, people).

1. ECOLOGICAL PREMISES

Askania Nova is situated in Europe, near the southern border of the Sivash steppe plain, between the West Sivash and the flood-plain of the Dneiper river. Not long ago this plain was characterized as a virgin Stipa and Festuca steppe with vast depressions, or “bottoms” that were characterized by specific associations of wheat grass, sedge, and motley grass. The solontshaks with their parts on the surface are frequent in the Sivash steppe, near the shores of Sivash, in particular. At present all virgin steppes here are developed. Askania Nova is the only place where the intact plots of steppe have been preserved, their total area is about 11 000 hectares, and they are characterized by specific vegetation.

The geophysical situation of Askania Nova is at latitude 46°28'N. The low position above sea level and abundant solar radiation result in a mild climate. The total number of sunny hours per year is 2255, and 1000 out of this number are registered in June-August (Babich 1960). The absolute maximum air temperature in the summer is 38°C. The summer season lasts for about 140 days, from mid-May till the end of September The summer is dry and hot. The average winter temperture is 1.2–3.3°C; occasionally it can be as low as minus 30°C. Such temperature fluctuations with a range of 41–49°C point to instability of the weather in winter, that is characterized by sharp falls in temperature and by unexpected thaws. Low temperature is accompanied by the growth of air humidity up to 80% that results in formation of winter fogs and glazes, as well as of continuous winds, blowing mostly from the north or northeast with an average velocity of 5.3 m/s. During occasional storms the wind velocity can reach 25 m/s.

The snow cover in Askania Nova is formed in December and melts in early March. Nevertheless, this cover in the steppe areas is not more than 10 cm thick and it can disappear in mid-winter as the result of thaws. Only in most severe winters the snow cover may exist for 60– 70 days in a row. In such cases its thickness may reach 40 cm and more.

One of the most unfavourable meteorological factors that is observed from early spring is cold dry winds of the northern quarter that bring back the cold weather, impede the vegetation growth, and reduce soil humidity. Occasionally, spring winds turn into storms, with a velocity of 25 m/s.

Overall the climate in Askania Nova may be defined as dry, with a long hot summer and a short unstable winter, with the prevailing dry winds; the annual precipitation is 376 mm; the solar radiation is high.

The horses, ranging on a year-round basis, can successfully tolerate the Askanian climate as a whole. Their skin is 5933±199 microns thick, i.e. 37% more than that of the domestic horse in Europe. The length of hair in the summer is 15.0 mm; it increases up to 20 mm and more in wintertime. The hair cover comprises pop-hair of the first degree (82.6%), pop-hair of the second degree (13.2%) and underhair (4.2%). In wintertime the mane grows from 126 to 140 mm (Katsy and Klimov 1983). It should be stressed that the horses imported from the USA in 1982 that lived in the conditions of city zoos managed to overcome successfully two winters, the last one being extremely severe. Moreover, they produced off spring that points to their successful acclimatization. The horses in the wild used to get food from under the snow cover, pawing and digging with the muzzle. The horses in Askania Nova do not have that skill since they have not been taught by their parents as happens in nature. Nevertheless, the horses demonstrate scratching movements that are characteristic of caballus in the attempt to dig snow and get the plants that are hidden under it. This behaviour pattern is not yet formed completely, but there is a hope that availability of “mentors”, i.e. the animals that possess this skill, will help the others to master it. The wild horses are unpretentious as regards the botanical composition of forage, they eat both wheat grass (Festuca, Stipa, Bromus) and various sagebrushes, pod-bearing plants, etc.

Automobiles, tractors and people moving by the road behind the network fence make the horses withdraw only by 20–30 m. The aircraft flying over the enclosure do not produce any defensive response.

2. ETHOLOGICAL PREMISES

2.1 Formation of Behaviour Patterns

The newborn foals strive to maintain permanent visual, olfactory and tactile contacts with their mothers. Their mothers ensure their protection within and beyond the band, and they control the observance of individual distances. The infant foals (till the age of 6–8 months) follow their mothers, seeking protection. They start periodic grazing at the age of 8–9 days, and at the age of 10–14 days they begin to establish personal, competitive contacts with other foals of their age group. When they shift from suckling to grazing, their alliances with the mothers become weaker, and they try to establish contacts with older horses. An element of competition among contemporaries and a certain degree of aggression on behalf of the mature horses are observed at this period. By the end of this period the social ranking is clearly seen. Older band members define individual distances between the infants.

Juvenile horses (till the age of 1.5–2) separate from their mothers but maintain links with them. Personal contacts are strengthened as a result of playing and competitions. The future reproductive behaviour pattern begins to be displayed by individual elements of their play, the division of their roles according to their sex being observed. The juvenile horses try to keep individual distances.

The females at the age of 2–3 years, along with their mothers, automatically become the members of a general or of a specific harem. Cases of ousting of the young mares from the band were observed. They are fully subordinate to the stallion and mate with “their” stallions, when reaching sexual maturity. The alliances within a band are personalized, competitive or polygenic when the family is formed. The mature mares are able to keep individual distances when ranging, recreating, when the band is moving, or in dangerous situations, but they do it selectively and not always. They do not mark their territory, and do not protect it. The passive defensive response to the effect of outer factors prevails. They do not involve other mares into their band and perform passive control over the band. Without a leading stallion the mares are not capable of coordinating the band activity and providing adequate response to outer factors. The mothers with foals increase individual distances and their degree of aggression. In the course of time the mares may change their social ranking and join the leading group.

The males at the age of 2–3 years develop the personalized contacts in their group with the use of grooming, playing, fights and their ranking is more pronounced as compared with the previous age group. They try to court the females, to involve them in their band, demonstrating reproductory behaviour. They get into the focus of the leading stallion's attention and the latter starts to pursue them and eventually drives them out. The ousted stallions form a bachelor group with its own ranking, but the relations within such a group are not competitive till the reproductive stimuli appear and then some stallions try to take hold of a mare The bachelors neither mark, nor protect their territory. The young stallion actions are characterized by incomplete performance of aggressive and reproductive behaviour due to underdevelopment of some parts of these patterns. As a result, they fail to reach the top rank, and consequently cannot take hold of mares. When the leader is absent in a band the most dominant stallions take hold of several mares or the entire band, ousting other mature males, and try to secure their social and reproductive status. During the period of social structure establishment the relations within a band are mostly aggressive, since the possessive efforts of a stallion are repulsed by the dominating mares. Upon stabilization of the social homeostasis in the group such stallions take secondary roles after the dominating mares. They neither mark, nor protect their territory; they perform no goal-oriented control of the band that has its adverse effect. A stallion may occupy the leading position in a group only after it has gained the experience of leading the harem, approximately by the age of 10 years. Such a stallion becomes unquestionably dominant in his band: he patrols, marks and protects his territory. The size of such territory, number of mares and the quality of efforts needed to hold and protect the mares are directly dependent on the masculinity and social experience of a leader. The leading stallion with socially adequate actions displays the concern of the band members, he coordinates all its activities and counteracts the environmental pressure that eventually leads to the prosperity of the population and further propagation of this species.

2.2 Reproductive Behaviour

The growth of reproductive activity, as regards Przewalski horses in Askania Nova is observed in early spring with its warmer weather, long days and sunlight. The young animals at the age of 1–2 years begin to involve each other in active play that results in expanded and strengthened links between the horses. The sexual character of their play, that increases by the summer, is clearly seen. In May the mares start to produce offspring and that leads to the change of their position in the group. They display no interest toward males and other horses, become aggressive and cautious. Mothers aggressively defend their foals, attacking everybody in their vicinity, striking them with their hind legs. The stallion stops his efforts to approach such a mare but resumes his efforts more persistently on the 6–8th day. Copulations are observed 11–15 days after confinement.

A certain peculiarity is observed when the stallions of various ages are mated to mares. Young stallions display inadequate behaviour and their helplessness is characterized by a series of chaotic, unfinished movements. The older stallions are able to perform successful copulation; they court a mare displaying the entire range of sexual behaviour. Nevertheless, the mares probably feel uncomfortable and often reject the stallions. Mature experienced stallions (e.g. Pegasus) do not waste time on preliminary courting but mount the mare without any preparations. In such cases the mares are submissive and obedient. There were no registered cases of rejecting Pegasus. The frequency of mating depends on the age of a stallion. Young stallions, placed in the herd after a long period of rest can copulate once per hour for 4–5 hours in a row. Their activity increases if the mares are receptive. Moreover in some cases when a stallion had a surplus sexual motivation he managed to mount the mare that was not in sexual heat and displayed vigorous resistance. At the expense of his physical strength he managed to neutralize her actions, pressing her croup to the ground and managed to cover her (Klimov and Paklina 1983).

Within the framework of sexual behaviour a number of anomalies and inadequate actions can be outlined:

Group 1: Anomalies caused by incomplete formation of young horse sexual behaviour, their response to animals of the same sex, the disrupted chain of actions in a sexual ritual, absence of certain links in this chain, by the absence of social skills to behave with partners and ignoring them when physical strength prevails.

Group 2: Preference of interspecific sexual relations, i.e. mares in heat prefer domestic stallions of the Reserve workers, or formation of a harem that consists of other species representatives (Shetland pony, domestic mare) by a stallion.

It is our view that such phenomena are caused by the following factors: incomplete formation of social behaviour patterns of stallions due to their frequent isolation from mares and grouping with other stallions; mares have no contacts with stallions which results in the absence of social knowledge; maintenance with representatives of related species (domestic horse) that leads to a breach of adequate behaviour patterns, of social skill formation.

2.3 Social Structure and Organization

One of the major factors of uniting the horses in the structural system, of their distribution in a given area and functioning in a strange environment, is the social, or ethological band structure. It comprises several structural units, i.e. age and hierarchic groups, permanent or temporary formations and their modifications. The hierarchic steps that are passed consecutively by each animal in the course of life are defined, in major part, by the morpho-physiological and psychological status of the horses. The position, ranking and social role in a band are formed due to the combination of these factors.

Age factor is of great significance for social ranking. Newborn foals stick to their mothers and occupy the same position in the ranking system. While they are growing and begin to graze the distance between mother and foal increases and by two-months of age they may be included into the lowest ranking group of juvenile horses. This group is followed by yearlings, two year olds, semi-mature and grown-ups. This formal age ranking is the basis for the band hierarchy. The animals of the junior age groups may be easily distinguished due to their considerable differences in physical features; however the mature animals have almost no visual differences between them having approximately the same size, etc. Nevertheless observations proved that each animal occupies its own position in the general hierarchy of a band.

A band's stallion, that is also called a “leader”, occupies the top position in the social structure of his band. For a number of years the Askanian band of wild horses was headed by Pegasus, the oldest stallion in the group (22 years old). The activity of this stallion is characterized by the full range of behaviour patterns characteristic of the males of the Equus species, that have its functional and evolutionary meanings. All his actions may be divided into the following: (i) ways and methods to establish his dominant position; (ii) ways and methods to control and defend his band; (iii) performance of a reproductive function that is both the result and the tool of his dominance. The first group of activity stipulates, in major part, the aggressive forms of influence, and their ritual meaning. When performing control over the band the stallion searches for the optimal location of his group, defines the routes of their movement, controls the activities within the band, sets the “schedule” of visiting the watering sites. The leader drives out other stallions, stops fighting, maintains order in his group, takes care of the mares and the young stock. The external factors, such as appearance of other stallions, of other species, of people and their objects (e.g. domestic horses, a cart, automobiles) become the area of its special concern.

A leading group, or “the nucles” can be distinguished within the herd; such a group comprises predominant mares (3–5) that is the basic structural and functional unit. The mares of the “nucleus” define direction at grazing, when on the march, or escaping from danger. In extreme conditions the lead is often taken by the oldest and most experienced mare, Volga. Besides individual qualities, the family ties are one of the factors that have an effect on a band hierarchy. A mother and foal are the primary family unit. When a mare has a yearling, or two year-old foal, which is less frequent, it is possible to regard it as a family cell inside the band.

Along with the basic structural units within the band, other, more or less permanent and temporary formations are observed that introduce changes into the band structure. The presence of such formations depends on the season, this or that enclosure, ecological situation, demographic composition of the herd, and on the physiological state of individual animals. The permanent, or stationary units are as follows:

1. Harem groupings. When stallion Pegasus is present in the band, all mores are members of his harem. When this stallion is absent and other mature stallions are released into the herd that were in isolation prior to this, one of the newcomers takes hold over the mares driving out the others; when two stallions are similar, as regards their physical strength, they can divide the herd into two parts.

2. Bachelor groups and solitary horses. Such groups are observed when the band has a leader that ousts other semi-mature and mature stallions that are grazing at some distance from the herd. They can occasionally occupy the herd's territory but never mix with it and stick to their individual group.

3. Groups of horses with personal bonds. Such bonds may exist for a number of years. Such bonds can form between the animals, representing different species.

   Temporary formations are as follows:

1. Groups that are formed during a mare's cycle. Two options are possible; first, when a mare has a “train” of courting stallions striving to sire her, and, second, when a receptive female is striving to copulate with a stallion.

2. When foals are born, the character, behaviour and rank of their mothers changes radically. They become the most cautious and aggressive animals in the band and since they are motivated by the same drive, that is to preserve their offspring, they undertake joint actions when in an emergency, or a strange stallion penetrates their territory, etc. Their actions are characterized by extreme cruelty and completeness of reactions toward the enemy, that allows them to occupy the second position after the leader in the band's hierarchy, the latter often preferring not to have any conflicts with the mothers.

The age status and the rank of each horse that makes up a population are relatively balanced but the time comes when some horse upsets this balance and tries to change its rank. Such actions are accompanied by conflicts with the animals of the same rank and with those of higher ranking. Thus, the rank of each horse is defined and preserved by the individual qualities of each animal, by the horses of the adjacent ranks, by high-ranking horses, and by the leader. The change of psychological and physical status of individual animals is caused by their own development, reproductive processes, and by their physiological state, e.g. injuries, diseases. Along with external factors, such as climate, season, new horses in a group, or removal of some old ones, the internal reasons change the ranking of many horses eventually upsetting the entire band structure. Such changes are very painful, they are accompanied by increased excitement, stress, etc. The number of aggressive actions sharply grows, up to 40–50 per 5 minutes. The number of activity shifts grows from 27–30 to 68–70 during the light hours. The time spent on grazing and rest sharply decreases (46%, 34% correspondingly), the time of marching increases up to 20% of the total, while the normal indicator is 8–12%. Depending on the factors that have caused the change in social structure, the balance is reached in 1–3 days, each animal occupying its own cell within the general structure that remains stationary till a new “coup”.

The relations within the group are regulated by the demonstrative and agonistic forms of behaviour. At the initial stage the contact between two mature stallions is based on visual stimuli and ritual movements, characteristic of their rank, motivation, and claims. If the stallions' interests remain unrealized at this stage their contact continues with increasing excitement. The stallions draw closer to each other, rear, start to bite each other and to hit with the front and hind feet. such fights may lead to serious injuries. It is not characteristic of mares to use ritual actions, they just start aggressive actions.

2.4 Spatial Organization

The mode of behaviour and the spatial distribution of a band depend on the enclosure where it is placed: its size, topography, locality, configuration of the territory position and availability of water reservoirs, shelters and saline plots. According to these factors, the group forms its spatial structure, i.e. the “central” and “border” zones, the area to be patrolled, marking places, areas for rest, activities, “toilets”, etc. The principal territory where the horses are maintained during the reproduction period (May-June) is enclosure No. 3.

The “central” zone is situated in the western part of the enclosure and represents a dusty plot of land, 18 m in diameter. This plot is a so-called “tyrlo” used for rest and recreation where the horses spend most time of the day. Along its boundaries this plot is surrounded with auxiliary “services”, e.g. “dust baths” with an area of 12–15 m² and “public toilets” with heaps of horse faeces, that are regularly supplied with new portions. The “toilets” are an important part in the system of communicative and hierarchic links between the horses, they are also significant for establishment and maintenance of the territorial principles in this group. Additional “toilets” and “recreational facilities” are situated in the most frequently visited parts of the enclosure. There is another “tyrlo”, an outlying one, with a diameter of 16 m that is visited by horses less frequently; it is situated at a distance of 50 m from the “central” one. Both plots are overgrazed and lack vegetation. The plot for “after-watering rest” is situated near the watering site and is used by horses only in the evening or in the daytime when the weather is cloudy. The area of this plot is 140 m²; it has several “dust baths” and 3± “toilets” along the boundary. The horse paths start directly from the watering site. They cross the entire enclosure and end in the “central zone” or by the grazing areas. According to observations, such paths are renewed every spring and used during the entire season. The paths are divided into “arterial” ones that link the watering site with the “central” zone by a straight line, and they are 3–4 cm lower than the soil surface, and the outlying paths that cross the enclosure in every direction and can hardly be noticed in the thick grass cover. In any case when a horse is not grazing and is moving alone or with a group it prefers to move by path. There are several “toilets” along the paths, especially the arterial ones, that are situated with a distance between them of 15– 20 m.

The patrolled zone is situated along the boundary of the enclosure. It is especially well pronounced along the northern and eastern fences behind which other species are managed. This zone consists of a similar path along the fence, with all its curves that is 50 cm from the net. The patrol path skirts the water reservoir and at the same time crosses it. The patrolling stallion usually goes round the reservoir; when he is in an excited state, following an external irritant (other stallion, riders) that are behind the fence, he can cross the reservoir directly. There are marking places near each gate, in the northeast part of the enclosure, around the geodetical column and in some sections of the patrol path. Such places represent small plots with an area of 1–1.5 m² that are filled with horse faeces, periodically renewed by the leading stallion. It should be stressed that he does so only in the case of a threat coming from the adjacent areas (wild animals, domestic animals, patrol men). In a calm situation when the external irritants are absent he may only study such a plot and leave it since there are no other stallions that could leave markings and the mares are not allowed in these patrol areas.

When calm, the group of horses managed in enclosure No. 3 covers a distance of 4–7 km per day, in the large enclosure the horses move by 8–10 km; on hot days this distance is sharply decreased. During the driving of the band to another place in the fall, the resistance of the band is sharply decreased after 10–15 km of driving. It indicates that the horses suffer from lack of movement, as compared with the horses in the wild when the riders could follow a band for 2– 3 days, tiring out several horses and managing to obtain only foals (Klements 1903).

2.5 The Band as a Functional System

The peculiar feature in behaviour of the wild horse band is its permanent readiness to respond to external irritants, with the use of group forms of adaptation that are characteristic of the social animals that lived in the severe conditions of Central Asia; these forms are clearly seen when studying the animals born and maintained in captivity. Thus, when ranging the horses display social constructions, such as “circle”, “horseshoe”, “wedge”, “arc”, “rank”, “band”, etc. In hot weather they protect themselves against midges, using the coupled modifications of the “tail-head” stand, or collective ones standing by a double rank, horses waving the faces of their neighbours. In some cases the band is concentrated in an amorphic or dense group. When on the march the horses move in a chain, when each horse strictly follows the front one. There may be several such “chains” when the band is moving.

During any function, rest, ranging, marching, 2–3 horses study the environs, assisted by different spatial orientation of horses when resting, or grazing. The leader is always at some distance from the group and spends most time of the day observing the surroundings. Occasionally he patrols the territory.

When a strange object appears it is marked by a loud “inhalation-exhalation” and alert of the horse that was the first to notice it. This signal is passed over to all band members in a radial direction, after which all horses cautiously watch the object. The leader is the first to see danger, as a rule, and he meets the object at the boundary of the enclosure, near the gate or the net. The response of the band depends on the nature of the irritant. When the object is familiar enough, or totally unfamiliar, it can cause curiousity and not aggressive actions. Thus the horses may approach an observer and stop at a distance of 20–30 m, surround him and watch for 2–3 minutes. The leader usually prevents such actions, and actively grouping the mares drives them to a remote area. When the object causes alarm, the band consolidates into a solid group, upon the first warning signals, and is ready to follow the leaders. The mares with foals are placed in the rear of the group, the mature animals are responsible for choosing the direction and the young stock that are “in the way” and have no experience in extreme situations remain in the front of such a group. The leader usually stands halfway between the band and the object, demonstrating the aggressive ritual behaviour and marking the territory. Both he and the mares in the front thoroughly study the object (visually) and are ready to respond to its possible actions. When this object is a rider, or a man in a horsecart, the leading stallion attacks the domestic horse, trying to kill it. When the leader desires to drive the mares away, he makes a signal, taking a “goose-like” stand, lowering his head to the ground and waving the head and the mane sideways. The signal is usually received by one of the mares in the “nucleus” that takes the lead. There is a need to stress the joint coordinated actions of the horses avoiding the pursuers. When pursuing the band during the inter-season drives it is clearly seen how the leader, or the leading group perform a “shuttle” manoeuvre, in an attempt to change the route, imposed by the pursuers. The animals make false movements and at the right moment make a spurt and get beyond control. Such a breakthrough may be performed not by the members of the leading group alone but also by other, most resolute, animals. After a single horse breaks out it is impossible to hold the band anymore. Like an avalanche the horses follow the leader, passing the riders. At this moment the animals do not display the former fright of man and do not keep the necessary distance (30–50 m). Only after the horses are grouped again may their driving be resumed, starting from the very beginning.

Thus we observe the functioning of the intra-group mechanisms that coordinate and organize the band's actions; we can use them to control the horses' behaviour and their territorial distribution. It should be stressed that all activities of a horse group described above, both on the range and in response to external objects, are characteristic only of the band with a full-value social structure, formed in the course of several subsequent generations. The absence or replacement of an experienced leading stallion with one which is young and inexperienced will turn a well-organized functional band system into a chaotic mass. Such a group changes the activity patterns spontaneously, does not develop its territory, does not display stable rhythm in its daily activities, and when exposed to the strange, threatening irritants finds itself at a loss and gets into an impasse. It is possible to explain the rapid extinction of the wild horse in Mongolia and China by this very factor. The Arats (local residents) shot the stallions in the first place, because they caused damage and prevented the successful hunting of the wild horses. Without a leader the mares failed to withstand the pressure of the environment.

As part of the stallion exchange programme, stallion Sigor was imported from the USA. He used to approach a man, allowed him to pat, etc. When he was given an opportunity to form his own band in a separate enclosure he failed to do so, even in the absence of competition with other stallions. Having no skills of aggressive or reproductive bahaviour he failed to become dominant and to take hold over the mares. When he made an attempt to copulate with them they rejected him. Later, during the second reproductive season, he managed to form a harem and tried to protect it, attacking people in a horse-cart in 1984 when he also started to protect his territory. Despite imcompleteness of these behaviour patterns, we may hope that in the course of time this stallion will gain all the social skills that are characteristic of stallions in the wild.

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FAO TECHNICAL PAPERS

FAO ANIMAL PRODUCTION AND HEALTH PAPERS:

1. Animal breeding: selected articles from World Animal Review, 1977 (C* E* F* S*)

2. Eradication of hog cholera and African swine fever, 1976 (E* F* S*)

3. Insecticides and application equipment for tsetse control, 1977 (E* F*)

4. New feed resources, 1977 (E/F/S*)

5. Bibliography of the criollo cattle of the Americas, 1977 (E/S*)

6. Mediterranean cattle and sheep in crossbreeding, 1977 (E* F*)

7. Environmental impact of tsetse chemical control, 1977 (E* F*)

8. Rev. Environmental impact of tsetse chemical control, 1980 (E* F*)

9. Declining breeds of Mediterranean sheep, 1978 (E* F*)

10. Slaughterhouse and slaughterslab design and construction, 1978 (E* F* S*)

11. Treating straw for animal feeding, 1978 (C* E* F* S*)

12. Packaging, storage and distribution of processed milk, 1978 (E*)

13. Ruminant nutrition: selected articles from World Animal Review, 1978 (C* E* F* S*)

14. Buffalo reproduction and artificial insemination, 1979 (E**)

15. The African trypanosomiases, 1979 (E* F*)

16. Establishment of dairy training centres, 1979 (E*)

17. Open yard housing for young cattle, 1981 (E* F* S*)

18. Prolific tropical sheep, 1980 (E*)

19. Feed from animal wastes: state of knowledge, 1980 (E*)

20. East Coast fever and related tick-borne diseases, 1980 (E*)

20/1. Trypanotolerant livestock in West and Central Africa, 1980; Vol. 1 - General study (E* F*)

20/2. Trypanotolerant livestock in West and Central Africa, 1980; Vol. 2 - Country studies (E* F*)

21. Guideline for dairy accounting, 1980 (E*)

22. Recursos genéticos animales en América Latina, 1981 (S*)

23. Disease control in semen and embryos (E* F* S*)

24. Animal genetic resources - conservation and management, 1981 (E*)

25. Reproductive efficiency in cattle, 1982 (E*)

26. Camels and camel milk, 1982 (E*)

27. Deer farming, 1982 (E*)

28. Feed from animal wastes: feeding manual, 1982 (E*)

29. Echinococcosis/hydatidosis surveillance, prevention and control: FAO/UNEP/WHO guidelines, 1982 (E*)

30. Sheep and goat breeds of India, 1982 (E*)

31. Hormones in animal production, 1982 (E*)

32. Crop residues and agro-industrial by-products in animal feeding, 1982 (E/F*)

33. Haemorrhagic septicaemia, 1982 (E* F*)

34. Breeding plans for ruminant livestock in the tropics, 1982 (E* S*)

35. Off-tastes in raw and reconstituted milk, 1983 (E* F* S*)

36. Ticks and tick-borne diseases: selected articles from World Animal Review, 1983 (E* F* S*)

37. African animal trypanosomiasis: selected articles from World Animal Review, 1983 (E* F* S*)

38. Diagnosis and vaccination for the control of brucellosis in the Near East, 1983 (E*)

39. Solar energy in small-scale milk collection and processing, 1983 (E*)

40. Intensive sheep production in the Near East, 1983 (E*)

41. Integrating crops and livestock in West Africa, 1983 (E*)

42. Animal energy in agriculture in Africa and Asia, 1984 (E/F*)

43. Olive by-products for animal feed, 1985 (Ar* E* F* S*)

44/1. Animal genetic resources conservation by management, data banks and training, 1984 (E*)

44/2. Animal genetic resources cryogenic storage of germplasm and molecular engineering, 1984 (E*)

45. Maintenance systems for the dairy plant, 1984 (E*)

46. Livestock breeds of China, 1985 (E*)

47. Réfrigération du lait à la ferme et organisation des transports, 1985 (F*)

48. La fromagerie et les variétés de fromages du bassin méditerranéen, 1985 (F*)

49. Manual for slaughter of small ruminants in developing countries, 1985 (E*)

50. Better utilization of crop residues and by-products in animal feeding: research guidelines - 1. State of knowledge, 1985 (E*)

50/2. Better utilization of crop residues and by-products in animal feeding: research guidelines - 2. A practical manual for research workers, 1986 (E*)

51. Dried salted meats: charque and carne-de-sol, 1985 (E*)

52. Small-scale sausage production, 1985 (E*)

53. Slaughterhouse, cleaning and sanitation, 1985 (E*)

54. Small ruminants in the Near East: Vol. I, (E***) Selected papers presented at Tunis Expert Consultation

55. Small ruminants in the Near East: Vol. II, (E***) Selected papers from World Animal Review

56. Sheep and goats in Pakistan, 1985 (E*)

57. Awassi sheep, 1985 (E*)

58. Small ruminant production in the developing countries, 1986 (E*)

59/1. Animal genetic resources data banks, 1986 (E*) 1 - Computer systems study for regional data banks

59/2. Animal genetic resources data banks, 1986 (E*) 2 - Descriptor lists for cattle, buffalo, pigs, sheep and goats

59/3. Animal genetic resources data banks, 1986 (E*) 3 - Descriptor lists for poultry

60. Sheep and goats in Turkey, 1986 (E*)

61. The Przewalski horse and restoration to its natural habitat in Mongolia, 1986 (E*)

Availability: September 1986

Ar - Arabic
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* Available
** Out of print
*** In preparation

The FAO Technical Papers are available through the authorized FAO Sales Agents or directly from Distribution and Sales Section, FAO, Via delle Terme di Caracalla, 00100 Rome, Italy