The distribution of fallow deer: a worldwide review

Mammal Review Volume 10, Numbers 2 &i 3 June/September 1980

The distribution of fallow deer: a worldwide review NORMA G. CHAPMAN and D. I. CHAPMAN ‘Larkmead ’, Barton Mills, Bury St. Edmunds, Su ffolk IF28 6AA

CONTENTS

1. Introduction . 2. Distribution of European Fallow deer .

2.1. Africa . 2.1.1. Democratic Republic of Madagascar . 2.1.2. South Africa .

2.1.2.1. Cape Province . 2.1.2.2. Orange Free State . 2.1.2.3. Transvaal .

2.2. Australasia . . 2.2.1. Australia .

2.2.1.1. New South Wales . 2.2.1.2. Northern Territory . 2.2.1.3. Queensland . 2.2.1.4. South Australia . 2.2.1.5. Tasmania . 2.2.1.6. Victoria . 2.2.1.7. Western Australia .

2.2.2. Fiji 2.2.3. New Zealand .

2.3.1. Japan . 2.3.2. Turkey .

2.4.1. Austria . 2.4.2. Belgium . 2.4.3. Bulgaria . 2.4.4. Czechoslovakia . 2.4.5. Denmark . 2.4.6. Finland . 2.4.7. France . 2.4.8. German Democratic Republic (East Germany) . 2.4.9. German Federal Republic (West Germany) . 2.4.10. Greece . 2.4.11. Hungary . 2.4.12. Italy . 2.4.13. Netherlands . 2.4.14. Norway . 2.4.15. Poland . 2.4.16. Portugal . 2.4.17. Republic of Ireland . 2.4.18. Romania .

2.3. Asia .

2.4. Europe .

0305--1538/80/0900-0061 $02.00 0 1980 Blackwell Scientific Publications

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62 N. G. Chapman and D. I. Chapman

2.4.19. Spain . 2.4.20. Sweden . 2.4.21. United Kingdom .

2.4.21.1. England . 2.4.21.2. Northern Ireland . 2.4.21.3. Scotland . 2.4.21.4. Wales .

2.4.22. U.S.S.R. . 2.4.23. Yugoslavia .

2.5. North America . 2.5.1. Canada . 2.5.2. United States of America.

2.5.2.1. Alabama . 2.5.2.2. California . 2.5.2.3. Colorado . 2.5.2.4. Georgia . 2.5.2.5. Kentucky . 2.5.2.6. Maryland . 2.5.2.7. Massachusetts . 2.5.2.8. Nebraska . 2.5.2.9. New Mexico . 2.5.2.10. Oklahoma . 2.5.2.1 1. Tennessee . 2.5.2.12. Texas .

2.6. South and Central America . 2.6.1. Argentina . 2.6.2. Chile . 2.6.3. Peru. 2.6.4. Uruguay . 2.6.5. Leeward Islands .

3. Distribution of Persian Fallow deer . 4. Discussion . 5. Acknowledgments . 6. References .

99 100 101 102 106 106 107 108 109 110 110 111 112 112 113 114 114 114 114 115 115 115 115 115 116 116 118 119 119 119 120 123 130 132

1. INTRODUCTION The genus Duma comprises two subspecies, the European Fallow deer (Duma duma duma) and the Persian or Mesopotamian Fallow deer (Duma duma mesopofamica). Formerly, the same generic name was used also for the American White-tailed or Virginian deer, named Duma vulgaris, which is sufficiently different from Fallow to be in a separate subfamily. This led to much confusion, especially in North America where White-tailed deer are indigenous whereas Fallow are introduced. The matter was finally settled in 1960, following a petition by many eminent zoologists to retain the widely-used name Duma for Fallow deer but, for the White-tailed deer, to revert to the name Odocoileus virginianus, first given in 1832 (Anon., 1960). Some authorities still place Fallow deer in the genus Cervus, in which Linnaeus placed it when he first described the animal.

The Persian Fallow (Plate 1) is larger and typically its antlers have palms near the base whereas they are at the distal end in the European Fallow (Plate 2). The tail colouring of the two forms differs slightly. These are the most obvious differences on which the two subspecies are separated. The two forms have inter-bred in captivity and produced fertile young (Gray, 1972): such hybrid- ization points to a very close relationship. The two subspecies must have evolved from the same stock, so at some time they must have lived in the same area or at least have had adjacent ranges. Fossil and historical evidence does indeed suggest that in former times the distribution of the two types either overlapped or was contiguous (Chapman & Chapman, 1975). Both forms of Fallow deer have been identified from archaeological sites in Iraq and that country could well have seen the divergence of the two forms, but stronger evidence of the coexistence of the two subspecies is required. The Persian Fallow became the typical Fallow of the Middle East whereas the European Fallow was the form that became established westward, in Asia Minor (Turkey) and Europe. As we shall show, the present distribution of the two subspecies is totally different.

Distribution of fallow deer 63

It has long been known that European Fallow deer have been introduced widely to various parts of the world (Millais, 1906). Recent reviews on the distribution of Fallow deer have been very brief and have dealt with only part of the Fallow’s range (Niethammer, 1963; Uloth, 1971). The review by Siefke (1977) is largely a compilation from the literature, apart from the section on the German Democratic Repub1ic.Whilstreviewing the literature for our book (Chapman &Chapman, 1975) it became apparent that the present-day distribution of this animal wasincompletelyknown That information which had been published was sometimes conflicting and had suffered also from plagiarism. It has been stated that inTurkey European Fallow are now restricted to thenorthern slopes of the Taurus Mountains, yet the accompanying map shows their distribution as being over much of northern and western Turkey (Whitehead, 1972). In fact, the deer are restricted to threesmallareas near the south coast, on the southern foothills of the Taurus Mountains (Heidemann, 1976). We de- cided, therefore, before writing our book to review afresh the distribution of Fallow deer. This could not be done by personal visits so we wrote to potentially knowledgeable people in every country where we knew or suspected that Fallow deer may exist. In some cases these first correspondents were able to supply the required information. In many other instances a chain of correspondence eventually led to a person with the relevant knowledge. To try to avoid repetition of possible inaccuracies, information given in earlier reviews has been discounted. Published accounts of the presence of Fallow deer in an area have not been accepted unless they were the result of a personal survey by the author of the paper, or unless we have been able to have the information verified by someone in the country concerned. Even so, for some areas it has been extremely difficult to obtain information and errors and omissions have doubtless occurred: we should be grateful to have these brought to our attention.

A great deal of information was collected and a brief resume of that on distribution was published (Chapman & Chapman, 1975). This led to other populations being brought to our attention. In some cases there were doubts about the identity of the deer being described. We have satisfied ourselves, from photographs or specimens, that in all these cases Fallow deer certainly are present and that other species are not being misreported as Fallow.

The European Fallow is frequently kept in parks, particularly in Britain, and in other enclosures in many countries. In general, this review considers the distribution of wild or free-ranging animals, not those in captivity. This decision has presented problems. In Hungary, for example, Fallow deer live within a 7300 ha hunting enclosure at Gyulaj. Similarly, in the Netherlands some of the forests are fenced. Where such enclosures are very large or numerous they have been included, but the fact that they are fenced has been mentioned. The antlers of bucks from the Wedellsborg estate on the island of Funen in Denmark were disqualified at the World Exhibition of Hunting held in Budapest in 1971 on the grounds that they were from an enclosed herd (Carne, 1975). The deer are not completely fenced and are free to roam away from the estate. The deer, however, do stay on the estate because of the practice of providing food for them. In this review these deer are regarded as wild animals. Antlers from the enclosed deer at Gyulaj, on the other hand, were allowed to be entered in the Hunting Exhibition.

Figures for the total number of Fallow deer in a country, particularly European ones, are frequently published, sometimes even to the nearest unit (Alex-Hansen, 1970; Oko & WXodek, 1978) but rarely are details given as to how these estimates have been obtained. Even in Great Britain, it has recently been stated that 50 000 wild Fallow deer are present (Gibbs el al., 1975). I t is notoriously hard to count deer accurately and Fallow are no exception. Most censuses probably result in an under-estimate. For example, an attempt was made to exterminate Fallow on the 250 ha wooded Danish island of Aebeb. The population had been estimated at 30 deer but 42 were shot and still more remained (Sporon-Fiedler, 1930). In this case, the population was at least 40% greater than estimated although they were on an island and the probability of emigration and immigration during the culling period can be ruled out. If such a large error can be obtained in a small restricted area, how much greater the possible error must be in a large forest. Figures have been quoted for some countries because they may give a guide to the size of the population: a country claiming to have 20,000 deer obviously has far more than one claiming to have 200. In

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our view, such estimates are no more than inspired guesses and are probably best considered as minimum values. It should also be borne in mind that poorly organized surveys could result in deer being over-estimated.

This review attempts to record the present distribution of free-ranging Fallow deer throughout the world (Fig. 1) together with information on the habitat and climatic conditions of the areas where they are found. To appreciate the reasons for the present distribution, a knowledge of the history of Fallow in a country is important and, where known, this has been mentioned.

N. C. Chapman and D. I. Chapman

90 30 0 30 90

The draft manuscript of the majority of sections has been read by a knowledgeable person in the country concerned. The sources of information, where not published, are given for each country in the acknowledgements.

The greater part of the paper is concerned with the European Fallow deer and all mentions of ‘Fallow deer’ refer to this subspecies. The distribution of the Persian Fallow deer is discussed separately, in section 3.

2. DISTRIBUTION OF EUROPEAN FALLBW DEER 2.1. AFRICA With the exception of the Barbary red deer (Cervus elaphus) which occurs in Tunisia, and possibly Fallow deer in North Africa, deer are not indigenous to the African continent. There are no wild Fallow deer in North Africa to-day but until late last century, there were reports of them from the borders of Algeria and Tunisia and from Tunisia and Libya (Millais, 1906). The origin of these deer is unknown. The deer may have been introduced by the Phoenicians or Romans, as suggested by Joleaud (1935) who also reports their introduction to Egypt in the sixteenth century.

Fallow deer have been re-introduced to Africa more recently and wild or feral animals are present in the Republic of South Africa and possibly in the Democratic Republic of Madagascar.

2.1.1. Democratic Republic of Madagascar In 1932, Fallow deer from Czechoslovakia were released on the high wooded-plateau near Mamdja- katompo, 60 kilometres south of Tananarive (R. Albignac, pers. comm., 1974). Although much of Madagascar is situated in the tropics, the climate of the plateau is moderated by the maritime

Distribution of fallo w deer 65

influences and relatively high altitude. The temperature at Tananarive (1400 m) ranges from 13°C in July to 19°C in February and rainfall averages 1370 mm per annum.

Most of the plateau is now denuded as a result of “tavy”, the practice of itinerant agriculture whereby forest is partly felled, burned and the land cultivated, and the process repeated at intervals of about 10 years. Much of the area has become pseudo-steppe with Aristida rufescens as the dominant grass and some areas have been re-afforested with Eucalyptus and Pinus. By 1962 the deer had apparently moved into the rain forest to the east (Vincent, 1962). This dense forest, lying between 800 and 1300 m above sea level, and with Tambourissa and Weinrnannia as dominant tree species, is also greatly reduced and now remains only as isolated patches and a narrow strip. In 1974, the Director of Waters and Forests and of the Conservation of Soils was unable to provide any information on the current distribution of the deer and in 1976, after a visit to the island, Sir Hugh Elliott informed us the deer introduced in the Mamdjakatompo area had been exterminated some years ago. Brygoo (1972) referred to the only wild ungulate on the island as being a type of Wild boar (Sus scrofa). It would seem that Fallow deer have probably become extinct on the island.

2.1.2. South Africa Fallow deer are present in three of the four provinces of South Africa, namely Cape Province, Orange Free State and Transvaal (Fig. 2). The fourth province, Natal, has only two small popu-

/ Transvoal

250 krn Fig. 2. The distribution of Fallow deer in South Africa

lations of Fallow deer in fenced enclosures. One is in Highland Sourveld in south-west Natal; the other on the boundary between Valley Bushveld and Coastal Hinterland (Natal Parks, Game & Fish Preservation Board, pers. comm., 1977).

The origin of Fallow deer in South Africa is obscure. Cecil Rhodes has been attributed with the first introduction when, in 1897, he released some animals on the Groote Schuur estate on Table Mountain, Cape Town (Siegfried, 1962). Fallow deer were, however, already kept in 1869 in a park at Newlands House in Cape Town. When the government sold Newlands House, the herd of 100 deer was sold for E200 to a farmer from Somerset West (Thompson, 1968). Fallow deer from the Groote Schuur estate were taken to Lourensford farm also in Somerset West. Many animals from this district have been supplied as breeding stock to other places in South Africa, Fallow deer have been distributed widely to many farms and estates in the Republic and this practice is continuing at the present time.

Farms in this context in many parts of South Africa are not small areas of intensively cultivated land, but large areas of ‘native’ land which have been fenced for rearing stock. Some of these farms have turned from conventional farming to game ranching, the change in emphasis from meat and

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wool to hunting (i.e., harvesting) giving a better financial return for fewer animals (Deane & Feely, 1974; Johnstone, 1974). Besides native species, several species of exotic ungulates such as Barbary sheep or Aoudad (Ammotragus lervia), Hog deer (Axis porcinus), llama (Lama glama), nylgai (Boselaphus tragocamelus) and Asiatic buffalo (Bubalus bubalis), in addition to Fallow deer, have been introduced to farms (Bigalke & Bateman, 1962; Kettlitz, 1962; van Ee, 1962). In the areas of medium and low rainfall where extensive pastoral farming is practised, these farms may be very large. For example, in the Transvaal they range from 400 to 85,000 ha ( h e y & Kettlitz, 1964). Consequently, although Fallow deer are frequently reported as occurring on farms, it is probably justified, to consider them as feral even if not wild animals.

2.1.2.1. Cape Province Fallow deer are widely distributed in Cape Province and in 1970 they were present in at least thirty-two of the 11 3 districts, from Cape Town in the south-west to Mafeking in the northeast (Fig. 2, Table 1). This information is based on farmers’ written replies to a questionnaire sent out

N. G. Chapman and D. I. Chapman

Table 1

Presence of Fallow Deer in Cope Province, c. I970*

District Number of reports Number of deer (minimum)

Albany Albert (Burgersdorp) Bed ford Bredasdorp Britstown Caledon Calvinia Cape Town Colesburg Cradock Fort Beaufort Hanover

Hopetown Kim berley Laingsburg Middelburg Mafeking Molten0 Paarl Philipstown Piketberg Queenst o wn Somerset East Stellenbosc h Sterkstroom Stey nsburg Sutherland Tarkastad Vryburg Wodehouse Worcester

Hay

3 1 ? 2 1 3 1 2 1 4 3 1 1 1 2 2 1 1 4 3 2 1 1 4 3 4 2 1 1 2 1 2

4 45

124 91

1 7 1 2

5 5 4 3 1 1

15 2 ?

160 1

24 56 4 1 1

95 2 8 2

150 1 5 1 2

Total 61 869

*R. C. Bigallce, Pers. Comm.


by the Department of Nature Conservation of the University of Stellensbosch (R. C. Bigalke, pers. comm., 1975). In addition to all these mainland localities, Fallow are established on the notorious Robben Island which lies 8 km off the coast in Table Bay, about 10 km north of Cape Town. Three deer from Groote Schuur were taken to the island in 1963. BY 1977 the population had increased to about forty (South African Department of Prisons, pers. comm.).

About half the localities for Fallow deer occur on the Karoo, which is largely semi-desert with a low (125-375 mm per annum) and uncertain rainfall. The vegetation is mainly of open, dwarf shrub communities, particularly xerophytic members of the familv Asteraceae, although patches of forest occur along streams (Werger, 1978). Most of the other localities are situated either in temperate grasslands or in the south-western Cape. The latter has moist, mild winters and hot, dry summers: annual rainfall is in excess of 250mm and mostly between 300 and 2500mm. This Mediterranean type of climate supports a vegetation similar to chaparral and known as fynbos (formerly termed ‘sclerophyllous bush’). The fynbos comprises a rich flora of evergreen, sclerophyllous shrubs in wluch the families Testionaceae, Ericaceae and Proteaceae are particularly well represented (Taylor, 1978). The native mammalian fauna of the region is mostlv small species apart from the blesbok (Damaliscus dorcas) and the grysbok (Raphicerus melanotis) (Bigalke, 1978). Few Fallow deer were recorded in the Kalahari region with its savanna-type vegetation.

Although little precise information on distribution and density are available, Fallow deer obviously exist in a variety of habitats in the Cape Province. In the Middleburg district, for example, nine bucks and three does were introduced to Culmstock farm in 1908: now there is a flourishing population of about 200 which range over the farms Culmstock, Hielston, Manor Holme, Mount Melsetter and Southfields. On these farms the deer keep mainly to an area of about 4000 ha composed of brakveld, which is shrubby vegetation on brackish soil, and vleis, low-lying ground wluch is covered with water in the rainy season. In good seasons, however, they move out to graze over a wider area, including other farms, but when it becomes dry they return to the favoured localities. The deer frequent also the banks of the river which has dense bushy vegetation. Between twenty and thirty deer are shot each year, a practice which has been followed since about 1920 (R. C. Bigalke, pers. comm., 1974). Cecil Modes’ introduction of Fallow deer to Table Mountain was also successful. There in 1974, the deer on the Groote Schuur estate were estimated to number about 350. They occupy an area of about 400 ha on the lower slopes of the Devil’s Peak at altitudes between 65 m and 500 m where the habitat is oak and pine woods alternating with open grassland (Puttick, 1972).

2.1.2.2. Orange Free State Fallow deer occur in at least two of the forty-nine districts of the Orange Free State, namely Harrismith and Heilbron (Fig. 2) but they are probably distributed more widely than this.

The first introduction of Fallow deer to the province appears to have been in the early 1900s when Sir Percy FitzPatrick took them to his farm, Buckland Downs, in the Harrismith district (R. C. Bigalke, pers. comm.). These animals thrived and at least 183 deer were present in the district in 1972 (von Richter, Lynch & Wessels, 1972). Before 1914, Fallow deer had been liberated also by the Vereeniging Estates, on the border with Transvaal, 60 km south of Johannesburg. By 1937, these animals had increased to about fifty (Best, Edmond-Blanc & Witting, 1962) and at least 250 were present in 1978 (R. C. Bigalke, pers. comm.). Deer from here have been distributed to other farms in the Republic, according to the Director of Nature Conservation for the Transvaal Prov- incial Administration. Fallow deer were released in 1910 to Pyrnnsberg Farm, between Clocolan and Gumtree, in the district of Ficksburg on the borders with Lesotho and at least forty deer were present in 1937 (Bigalke, 1937) but their present status in unknown. Fallow deer have also been introduced to the Franklin Game Reserve in Bloemfontein (van Ee, 1962). All the above areas are situated in the Highveld, an area of largely treeless, undulating grassy plains, much of which lies over 1400 m. Summer is the rainy season with 635-760 mm precipitation per year, drought and heavy frosts occurring in the winter.

68 N. C. Chapman and D. I. Chapman

2.1.2.3. Transvaal Fallow deer appear to have been introduced to Transvaal soon after 1900 when some animals from the London Zoo were exchanged for tame springbok (Antidorcas marsupialis) by Mr Fick. Fallow deer still occur at the Fick family’s farm, Palmietfontein, near Volksrust, over 350 being present in 1967. About eighty to 100 are sold as breeding stock each year to other farms in the Republic (Uys, 1967). In 1973, Fallow deer were present also at Bryanston, near Johannesburg, according to the Director of Nature Conservation, Transvaal Provincial Administration but we have been unable to obtain any further information on their status (Fig. 2).

2.2. AUSTRALASIA Deer are not indigenous to either Australia or New Zealand but about fifteen species have been introduced with various degrees of success (Bentley, 1978). Particularly in the latter half of the nineteenth century many introductions of Fallow deer were made by individual settlers and by acclimatization societies whose members wanted deer primarily for sport. Fallow now occur in New South Wales, Queensland, South Australia, Tasmania, Victoria, in both North and South Islands of New Zealand and on the Fijian island of Wakaya.

Fallow are reputed to have been introduced to the Madang area of New Guinea by German settlers in about 1900. The deer fed in the areas of tropical rain forest that had been cleared for plantations and in 1953 the herd was estimated to number about 150 (Bentley, 1967). Doubts have been raised as to the identity of these deer and there appears to be no evidence to prove or dispute the introduction of Fallow (Bentley & Downes, 1969). Dr E. Lingren (pers. comm., 1975) has informed us that the Timor Deer (Cervus rimorensis) is now the only species of deer living in Papua New Guinea, the Fallow and Axis deer (Axis axis), if they ever existed, being extinct,

2.2.1. Australia In An Introduction to the Deer OfAustralia, Bentley (1978) gives a detailed history of the various species of deer. The earliest liberation of Fallow deer to the mainland was in the 1840s to Victoria, where hunting the carted Fallow became popular.

Although adaptable to a wide range of habitats in Australia, Fallow have not penetrated the heavily timbered mountain country but have shown a preference for open forest at lower altitudes. After an initial period of increase following their liberation, the numbers of Fallow deer have declined in most areas. This appears to be due, in part, to the fact that rural settlement and the consequent disturbance to wildlife also has taken place in these areas. Fallow are now present in five of the eight States and Territories but in only one, Tasmania, are they numerous (Fig. 3).

2.2.1.1. New South Wales New South Wales probably has the largest number of Fallow deer on the mainland of Australia (Fig. 3). In 1886, a pair of Fallow were released on the eastern side of Lake George, north-east of Canberra. Others were liberated nearby at Bungendore, at an unknown date. During the following 30 years they increased to an estimated 200 and extended their range southwards to Quanbeyan and eastward to Braidwood and Goulburn (Allison & Coombes, 1969). Subsequently roads and intensive agriculture separated the deer into two discreet populations. Now the Lake George Fallow herd, perhaps numbering over 1000, is probably the largest and most stable in Australia, excepting Tasmania. The deer are scattered over the hills around the Lake within a radius of 80 km from the point of release. The second population is established nearer to Goulburn which is 30 km north of Lake George (C. Allison, pers. comm., 1978).

Another large herd occurs in the northeast of the State, north of Glen lnnes in the New England Range. Animals were liberated in this district at various dates up to 1924 when six were released in Ranger’s Valley. The deer established themselves on both sides of the Severn River in an area bounded by Glen Innes, Emmaville and Deepwater (Allison & Coombes, 1969). In recent years some new stock has been introduced and, following a period of study and protection, culling now takes place annually (C. Allison, pers. comm., 1978).


390 krn Fig. 3. The distribution of Fallow deer in Australia

The habitat in the Lake George and Glen Innes areas is similar although they are over 600 km apart. The tableland, averaging 1000 m above sea level, has an annual rainfall of approximately 880 mm but at Goulburn it averages 735 mm. The average summer temperature is about 20°C. Although winters are not usually harsh, they can be in some parts such as Ranger's Valley. Typically the days are cool with cold nights giving an average temperature of about 8'C. Some freezing niglits are followed by sunny days but blustering wind, sleet and snow also occur, a pattern sirnilar to that of southern England. The plateau supports a park-like woodland (savanna) with well-spaced gum (Eucafyptus) trees and wide expanses of grass and isolated shrubs such as Yellow box (Euca- lyptus meffiodora) and New England peppermint (Eucalyptus nova-anglica). Such a habitat is reminiscent of an English deer park, except for the species of plants present, and may have prompted the early settlers to introduce deer for aesthetic reasons as well as for sport.

Two other introductions of Fallow deer to New South Wales led to the establishment of herds mainly in the neighbouring State of Victoria. Those released near Albury about 1884 crossed the River Murray, whch forms part of the State boundary. Other deer released in the second decade of this century in the Delegate-Bombala area, in the southeast corner of New South Wales, established a population in the Snowy Mountains, extending into Victoria and occasional reports are still received from the extreme south of New South Wales (Bentley, 1978). A small herd also exists near Nowra which lies 130 km south-west of Sydney (C. Allison, pers. comm., 1978). Another small herd occurs in the southern part of the Royal National Park, about 60 km south of Sydney. These animals, of the black variety, originate from deer rrleased in the 1880s (W. P. Crowcroft, pers. comm., 1977).

2.2.1.2. Northern Territory Fifty-two Fallow were introduced to the Cobourg Peninsula in the Northern Territory in 1912 but are thought to have become extinct. The four barasingha (Cervus duvaucefi) also failed to become

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established but, by contrast, the three sambar (Cervus unicolor) introduced at the same time flourished (Calaby & Keith, 1974).


2.2.1.3. Queensland Probably the largest herd of Fallow in Queensland (Fig. 3) is that in the north of the State, just south-west of Cairns (C. Allison, pers. comm., 1978). There, on the seaward side of the Atherton Plateau, the humid conditions support tropical monsoon forest. At Cairns the mean annual rainfall is 2235 mm. The mean temperature in January is 28°C and in July it is only slightly lower, 2 1°C.

In the south-east corner of the State, Fallow are present in two areas on the Darling Downs. One of these localities is west of Stanthorpe where a herd survives in the Jubbinbar Mount - Pikedale Station vicinity, following the release of animals at the latter place in 1890. The second population lies about 90 km to the north-east in the Maryvale State Forest in the rough mountain country of the Upper Freestone Valley. The main tree species in this open forest include Narrow- leaved ironbark (Eucalyptus crebra), Blue gum (E. feretecornis) and Ribbon gum (E. viminalis). The deer also feed on the crops cultivated adjacent to the forest (G. W. Saunders, pers. comm., 1979). Fallow were released at Maryvale in 1870 (Roff, 1960). The land is 600-1000 m above sea level and has an annual rainfall around 700 mm. The temperature in mid-summer is about 25°C and in mid-winter it lies between 10" and 20°C. The Fallow deer introduced in the 1870s at two other places on the Darling Downs, at Westbrook near Toowoomba and near Warwick, were exterminated (Bentley, 1978).

2.2.1.4. South Australia Now Fallow probably occur in four or five areas of South Australia (Fig. 3) although they were introduced to at least eight places, mostly in the Adelaide area. Most of the information for South Australia has been supplied by Mr Jack Conquest (pers. comm., 1974). In the days of their introduction the undulating scrubland in the south-east part of the State provided an ideal environment for them. Various eucalyptus, Silver banksia (Banksia rnurginafa), myrtle (Acacia myrrifolia) and other scrub vegetation, including Yacca tussock grass (Poa australis), provided browse and cover. South Australia is the driest State, with much of the land desert or semi-desert, and so where natural vegetation did occur much has been cleared for farmland. This loss of habitat has contributed to the decline of the Fallow deer in South Australia (Anon., 1971).

Before 1895, Fallow were introduced to Yallum Park, in the southern tip of the State, from where some were taken later to neighbouring Nangwarry and Penola. Escapees from this area gave rise to the population that still exists in the pine plantations and scrub in the Mt. Gambier- Millicent-Naracoorte region. Small numbers are also reported in native forest, in which Brown stringybark (Eucalyptus baxteri) predominates, in the Furner area (B. C. Gepp, pers. comm., 1979).

About 1900, some Fallow were taken to Bungaree Station, about 168 km north of Adelaide, and to Angaston and Pewsey Vale, about 56 km and 46 km respectively northeast of Adelaide. In each place the deer were kept initially in fenced parks but later escaped into the surrounding countryside. The descendants of the Pewsey Vale deer ranged over the Adelaide Hills but the last few were shot during the 1950s. The other two herds are also now extinct.

Two lorry loads of Fallow were taken from Pewsey Vale, about 1912, to a 4 ha enclosure at Buckland Park Station, near Port Gawler, from where some were subsequently released and found cover in the coastal country, well timbered with Gum trees and Polygonurn bushes. The deer had increased to about 1500 by the mid 1940s. Despite increased hunting the population maintained this level but during the 1960s shoothg by commercial interests greatly reduced them. The deer now favour an unusual habitat, the broad estuary of the Gawler River. The estuary fans out into many tidal rivuletsafter forming a lake which is fringed by flat land covered by succulent samphire (Arthrocnemurn spp.) bushes. The rivulets are lined by mangroves (Avicennia marina) and then by a zone of lignum (Melofhria rnicrantha). The deer sometimes wade through the water and pen- etrate into the mangrove wilderness to reach shellgrit banks above the surrounding areas. For


shelter they ‘burrow’ into the very dense lignum bushes. At Buckland Park itself a small herd remains and some deer from here were taken to Ashbourne, about 45 km south of Adelaide, to provide foundation stock for a Fallow farming enterprise.

Fallow still exist in the Mount Crawford-Springton vicinity, about 45 km northeast of Adelaide where refuge is provided by the extensive State-owned pine plantations in which Monterey pine (Pinus radiata) is predominant. This population arose from deer that escaped from Rosebank Station, near Mount Pleasant, to where they were introduced early this century.

In all these regions where Fallow occur, although the mean temperature in mid-summer (January) is about 2OoC, occasionally the temperature may exceed 4OoC. Mean mid-winter (July) temperatures are about 10°C although a few degrees of frost occur on some days. Annual rainfall in the Gawler area is about 450 mm but is higher, reaching 600-700 mm, in the Mount Gambier and Mount Crawford regions.

A late introduction of Fallow took place in 1949 when nine were liberated in privately-owned forest in the Flinders Ranges which lie northeast of Port Augusta, rise to 1000 m and have an annual rainfall of about 350 mm. In 1978 C. Allison (pers. comm.) believed that at least 100 Fallow were present, but the following year the Woods and Forests Department of the state knew of no feral deer in these ranges (B. C. Gepp, pers. comm.).

2,2. I . 5. Tasmania The island of Tasmania has more Fallow than any other Australian State with populations well dispersed through the Midlands.

Most of the introductions were made privately, before the foundation of the Tasmanian Ac- climatisation Society. The first consignment may have arrived in 1836 (Bentley, 1978). Some authors give the date as 1829, the year in which Axis deer were introduced to Tasmania. The earlier date is plausible since a meeting was held in Hobart in 1826 to discuss the ordering of game from England (Rolls, 1969), but Allison & Coombes (1969) give the year as 1834 when a Mr James Cox of Campbell Town imported Fallow deer from an English park. By 1863 several releases had been made and the Fallow population was estimated to be between 600 and 800 (Bentley, 1978).

Acentury later the increase in the number of Fallow, the only free-ranging species of deer on the island, was beginning to cause conflict between farmers and hunters. A conservative estimate of the population in 1972 was 7000-8000 (Wapstra, 1975). The deer occur within an approximately circular area of about 400 000 ha in the central and eastern Midlands (Fig. 4). Their range is divided into two main sections by the Midland Highway which runs approximately north- south. East of t h s road Fallow are found scattered within the area to the east of Launceston,

- 60 hm

Fig. 4. The distribution of Fallow deer in Tasmania (after Wapstra, 1975)

72

Campbell Town and Ross and extending eastwards, bounded approximately by Mathinna, Lake h a k e and Tooms Lake. The deer are not distributed continuously throughout this block of country, which includes Ben Nevis, Ben Lomond and Mount Hobgoblin, there being an area from Conara Junction eastwards along the South Esk Valley, and particularly north and south of Avoca, where Fallow do not seem to occur regularly. On the west side of the Midland Highway the main area inhabited by the deer lies within an area roughly bounded by Oatlands, Bothwell and Poatina and includes some of the great lakes. Another population occurs east of Poatina, between the Macquarie River and the Midland Highway, in the vicinity of Epping (Wapstra, 1975).

The deer live in the dry eucalypt woodland and pastures on the fringe of sclerophyll forests (Plate 3) in whichEucalyptus spp. predominate and bracken, Pteridium uquilinum, is a conspicuous feature of the ground cover. The effect of Fallow on the natural environment is relatively unknown but most of their time is spent in areas already grazed by sheep and cattle (Green, 1974). Most of the habitat lies between 500 m and 1000 m above sea level but a few localities are lower. The average annual precipitation in the region is about 550 mm. The mean minimum and mean maximum temperatures in winter are around 2°C and 10°C respectively, and in summer 9°C and


21°C.

2.2.1.6. Victoria The Fallow populations in Victoria today (Fig. 3) are few and small despite many early liberations by individuals and the Victorian Acclimatisation Society (Bentley , 1978). Cowling (1975) reported wild Fallow ‘scattered throughout’ the State but this would seem to be out-dated: his accompanying map indicated only two localities.

One of the largest herds to be established was at Werribee Park, west of Melbourne, from where the deer moved towards the coast and where, by the 1870s deer battues or drives were held to reduce the numbers. No wild deer remain in the vicinity now. Similarly, the herds which were established on Philip Island and also between the Bass and Powlett Rivers are extinct, the former being shot when damage to crops reached an intolerable level (Bentley, 1978).

Fallow were introduced in the 1850s near Lilydale and Healesville in the Yarra Valley, about 40 km north-east of Melbourne. They increased enormously in the then undisturbed river flats and open-forested foothills of the Warburton Ranges. By 19 19 protection was lifted but, even so, press reports 3 years later referred to thousands of deer in the Ranges. Now small populations survive near Healesville, in the vicinity of Maroondah Dam. The main species in the open forest are Narrow- leaf peppermint (Eucalyptus rudiatu), messmate (E. obliqua), blackwood (Acacia melunoxylon), Silver wattle (E. deulbatu), Hop bitter-pea (Daviesiu lutifolia), Forest wiregrass (Tetruwhena juncea) and bracken. There are also occasional reports from a few other localities in the eucalypt bush, particularly Kinglake where the vegetation includes Narrow-leaf peppermint, Broadleaf peppermint (Euculyptus dives), Brown siringybark, Red stringybark (E. macrorrhynchu), low shrubs and Tussock grass, but ground cover is generally sparse (A. Bentley, pers. comm., 1979). At Healesville the annual rainfall is 1000--1200 mm. The mean temperatures for mid-summer and mid-winter are 16°C and 7°C respectively. Fallow were released in the latter part of the nineteenth century in the rugged Blackwood Ranges, about 24 km north of the town Bacchus Marsh, and small numbers may exist there now. There are occasional reports from the Grampian hills in the west of the State (Bentley, 1978) but in the Wimmera country to the north their survival to the present time is uncertain (Allison & Coombes, 1969).

On the northern border of Victoria, Fallow are occasionally sighted along the Corowa-Albury- Lake Hume stretch of the Murray Valley. They originate from animals released in New South Wales and inhabit open forest along the river where Black box (Eucalyphts largiflorens) and River Redgum (E. camuldulensis) are the main species, with a ground cover of grasses and reeds. The deer also feed on the pastures and cultivated land on either side of the river (A. Bentley, pers. comm., 1979). Just to the east, where Fallow were formerly established, lies the Tallangata Forest where nineteen Fallow were released late in 1978 to the Railway Block in the Koetong Plantation on the Koetong Plateau. For the previous 6 years a few Fallow had been kept in an enclosure


within the plantation to observe their effect on the Monterey pine, the principal species grown. The plantation is part of 5200 ha of forest, whch will eventually be greatly extended, and includes some native hardwood trees and adjoins 6800 ha of marginal farmland and scrub to which the deer also have access. In this undulating country, lying between 600 m and 1000 m, the annual rainfall is between 1270 mm and 1520 mm and snow occurs in winter. The average temperature is about 21°C in mid-summer and 7°C in mid-winter (Bentley, 1978).

The Fallow reported near the Snowy Mountains in the far east of Victoria (Allison & Coombes, 1969) originated from animals released in the adjacent district in New South Wales. On the western boundary of Victoria, in the vicinity of Casterton, occasional reports of Fallow were made, at least until a few years ago. These resulted from a release of deer in South Australia. On Sunday Island, which has an area of 1600 ha and lies in Corner Inlet and north of Wilson’s Promontory, a small number of Fallow live with Hog deer: the Fallow were introduced for a specific breeding programme and can hardly be considered as a wild herd (C. Allison, pers. comm., 1978).

2.2.1.7. Western Australia The first of several releases of Fallow deer was made near Cape Leeuwin in 1899 and others were also liberated near Gingin, north of Perth. In both areas the deer failed to survive (Long, 1972) and there are no recent records of wild Fallow in Western Australia.

2.2.2. Fiji Fallow deer are present on Wakaya, one of the Fijian islands, lying in the Pacific Ocean about 1600 km north of New Zealand (Fig. 1). The deer were introduced about 1880 by a Captain Padd and about 1000 were estimated to be present in 1929 (Spencer, 1929). The deer population was believed to be about 400 in 1977 (F. Marseu, pers. comm.).

There has been confusion as to the identity of the deer on the island. Derrick (1951) reported that ‘During the early years of the present century Red Deer were introduced, probably from New Caledonia, and they have increased to such an extent that they tend to become a nuisance’. Whitehead(l972) believed the animals on Wakaya to be Timor Deer, basing his identification on a description and photograph of a male taken in 1954. He also recorded the presence of Timor deer in New Caledonia, which is about 1200 km west of Fiji. Derrick may have confused Timor deer and Red deer since the two are similar in appearance although the date that he gave is later than that of the Fallow introduction. Photographs of the deer and of antlers from culled animals, taken by Mr Bill Beckon in 1977 and seen by us, confirm that the deer are indeed Fallow. The Beckons saw no evidence of another species of deer when they visited the island to survey the vertebrate fauna but it is just possible that a few Timor or Red deer are present.

Wakaya which lies 90 km east of Viti Levu, the largest of the many Fijian islands, has an area of 770 ha. The highest point is 200 m above sea level and the island eoxperiences a steady prevailing wind from the south-east. The maximum summer temperature is 32 C and the minimum in winter is 16°C. Annual rainfall is around 420 mm but this occurs mainly during the humid summer. Droughts are not uncommon in winter and may last for several months but small pools may still remain in the stream beds to which the deer go to drink.

The deer inhabit light forest and scrub. The most conspicuous trees are candlenut (Aleurites moluccana), raintrees (Albizia lebbek), Dysoxylum quercifolium, D. richii, Ficus spp., mango (Mangifera indica), guava (Psidium guajava) and rose-apple (Syzygium malaccensis). In contrast with forests on other Fijian islands, there is very little understorey. The unusual bareness of the ground presumably results from grazing and browsing by the deer, which have been introduced only to this island, and the feral goats which sometimes feed with the deer. The dominant scrub plants are Turnera ulmifolia, Lantana camara and Dendrocnide harveyi (B. Beckon, pers. comm., 1978).

2.2.3. New Zealand New Zealand holds a rare ecological position in the world because its indigenous flora and fauna

74

have evolved in the total absence of browsing mammals, the only truly native mammals being two species of bat. Nineteenth century colonists were quick to alter the situation and during approximately 50 years over fifty species of vertebrates werd introduced (Wodzicki, 1950). These included predators, game animals and many which were considered only for their aesthetic appeal. Of the animals introduced, some thirty-five species became successfully acclimatized and of this number eight were species of deer. The European Fallow was one of the more adaptable and after Red deer is the most widespread of the deer in New Zealand (Harris, 1970). The situation regarding the distribution and density of the Fallow populations has changed so much in the last few years that most published sources of information (Christie & Andrews, 1966; Harris, 1967; Joll, 1968; Wodzicki, 1950) are out of date.

The introductions of Fallow deer to New Zealand are well documented. Most of them are described by Donne (1924) and Wodzicki (1950) but additional releases are cited by Clarke (1976) who dates the first, but unsuccessful, liberation as 1862 at Flaxbourne, Marlborough province in South Island. The second, better-known liberation, in 1864, was also in the north-east of the South Island, in the Aniseed Valley, near Nelson, where three animals from Richmond Park, England were set free. A larger consignment, from Richmond Park together with some from Carshalton Park, left London in the ‘Thurland Castle’ in November 1876. Of the thirty-three deer taken on board, twenty-nine arrived at Auckland Harbour ninety-six days later (Brett, 1924). The fate of one animal is uncertain, but ten were released near the Wanganui River in the south-west of North Island. The other eighteen were divided into two groups to be liberated in the Waikato district, one group at Matamata and the other in the Maungakawa range. Following a 2 d a y journey by paddle steamer up the Walkato River, the latter group completed the journey by being taken on sledges up themountainside,(L.H. Harris,pers. comm.),amorespectacular arrival than that of the fawns led on ropes for 6 km before their release at Flora Saddle in the Arthur Range some years later, (Clarke, 1976). Further liberations of Fallow from England, Tasmania and later from herds already established in New Zealand, followed until by 1910 the species had been released in at least eight places in the North Island and eleven in the South Island or their off-shore islets.

Most of the liberations were successful. An abundance of food and lack of predators favoured the Fallow and allowed it to exploit the forest below 600 m, scrub and grassland (Williams, 1973). To help all deer species to become established full protection was afforded until 1890 when Fallow could be shot under licence in certain areas. Of the few unsuccessful introductions, one was at Morven and another at Cairn Bush in the South Island (Baker, 1972). A pair of Fallow was released also on Kapiti Island off the south-western shore of the North Island but was soon removed because the island had been designated a sanctuary for native flora and fauna in 1897 (Wodzicki, 1950).

The rate of dispersal of Fallow deer has been relatively slow when compared with that of other ungulates. From Wanganui, North Island, they spread at the rate of 0.8 km/year over a 50 year period (Caughley, 1963). A similar figure, 0.9 km/year, was found over a decade (1920-30) in the Paporoa Range in South Island (Clarke, 1976). In consequence, the Fallow populations have mostly remained discreet and closely associated with the areas in which the liberations took place.

Most herds flourished so well that by the 1920s Fallow had reached such high numbers in some areas that they were declared ‘vermin’ and all hunting restrictions were lifted. Even before this time, about 1912, a herd at Whangamoa in the North Island had been exterminated by farmers because of intolerable damage to crops (Clarke, 1976). Numbers in many areas remained high, sometimes with serious effect on the habitat, until the 1950s. Bucks were hunted for trophy antlers and to-day Fallow remain a sought-after trophy animal by stalkers. Further reductions have occurred in the last few years, following the establishment of a profitable export market for Fallow venison mainly to the German Federal Republic and the Netherlands.

The only small islands on which Fallow now occur are in Hauraki Gulf, North Island. Some deer are present on Rangitoto and on Kaikoura islands and a few may still survive on Kawau too. On the North Island itself Fallow occur in several areas (Fig. 5). On the southern headland on the west side of Kaipara Harbour the deer live mainly in sand dunes and scrub where the tea-tree or



- P 175 krn

Fig. S.The distribution of Fallow deer in New Zealand

manuka (Leptospermum scopurium) predominates (R. Pittaway , pers. comm., 1976). South of there, small herds are present on the South Head of Manakau Harbour, extending to the estuary of the Waikato River. At the base of the Coromandel Peninsula, Fallow were present until recently in the Puriri-Matatoke area but a single female seen late in 1977 was believed to be the last survivor (L. H. Harris, pers. comm.). Due south of there, Fallow also occur at Maungakawa (Te-Tapui), between Matamata and Cambridge, while to the east another herd lives south of Te Puke, half-way between the coast and Lake Rototua.

The largest area of the North Island which has Fallow deer, and which probably now accounts for the greatest population in New Zealand, lies in the south-west, just north of Wanganui and centring on the river of that name. The deer are found within an area up to 60 km long and 40 km wide at its greatest width, here extending almost as far as the Waitotara River in the west and the Whdngaehu River in the east. The Wanganui valley lies at about 200 m above sea level but much of the deer's habitat is higher, some hills extending up to 1000 m. Part of the habitat is on privately owned land used for cultivating crops or grazing sheep but some scenic reserves and State Forests fall within the area. The Fallow lie up in manuka scrub, gorse (Ulex sp.) and kiekie (Freycinetia bunksii) which are dense in the damper situations adjoining the farmland and native forest, which comprises mainly a mixture of conifers and hardwoods. Hunting is encouraged in the State-owned parts and the Fallow are subjected to sustained and often heavy year-round pressure from sportsmen. The species is also being commercially farmed for venison on land within the area which is uneconomic for sheep or cattle (Anon, 1976a). In the lowland the July temperature is between 4" and 12°C and in mid-January it is between 12" and 22'C. Annual rain-fall is around 1000 mm.

Most of the six widely separated areas of the South Island where Fallow occur (Fig. 5) are at higher altitudes than most of those of the North Island and, therefore, coupled with being further south, are cooler and in winter are liable to receive snow which is uncommon in New Zealand except above 1200 m.

Within the North-west Nelson State Forest Park a scattered population is found in the Mount

76

Arthur area. Their range is bordered approximately by the Mount Arthur and Pikirunga Ranges in the east, the Leslie and Peel Streams in the south and west, and the Cobb and Takaka Rivers in the west and north (Anon., 1974a). This population owes its origin to two liberations, in the Arthur Range in 1910 and in the Upper Takaka about 2 years earlier; the two herds merged in the 1920s. The deer mainly inhabit the native forest where Red beech (Nothofugus fuscu) is dominant but in some areas is mixed with Silver beech (N. rnenziesii). Fallow here are often sighted in the snow- grass (Chionochloa sp.) above 1400 m. Red deer also inhabit the region.

South of Nelson, small numbers of Fallow still inhabit the head of the Aniseed Valley, near where some were liberated in 1864. During the 1920s the herd extended its range northeastwards along the Bryant Range, eventually merging with Fallow from the Teal Valley, at the north-west end of the Range, where some animals were released in 1906.

On the west coast, between Brunner and the Buller River, a population of Fallow exists in the Paparoa Range. They frequent the scrub and, higher up, the alpine tussock lands at an altitude of 1200 m where Chionochlou spp. are dominant (Clarke, 1976). Annual rainfall is very high, over 6000 mrn in some places. In the central part of South Island small herds of Fallow are present near Albury in the valley of the Opihi River, about 10 krn south of Fairlie, and extending onto the Two Thumbs Range. This Range extends inland from Fairlie in the north-south direction between Lake Tekapo and the Rangitata Valley and rises from 200 m to 3000 m, but here the deer are seldom seen above 1500 m. Rainfall is about 600 rnm a year.

The Blue Mountains in Otago province were formerly the major stronghold of New Zealand Fallow but heavy culling in recent years to protect newly planted trees has reduced the population substantially. The history of the herd, discussed by Baker (1972), dates back to 1869, not to 1867 as stated by Wodzicki (1950) as the deer imported in that year went to Morven near the east coast, not to the Blue Mountains. The deer increased rapidly: between 1893 and 1903, 2000 bucks are reputed to have been shot (Donne, 1924). Almost 4000 deer were killed in 1926 and a further 2000 the following year. During 1957 and 1958,6000 were taken within 8000 ha and a similar number were killed in 1961-62. At least 1550 Fallow were shot by sportsmen in 1972 according to permit returns but Baker (1973) considered that this figure may be as much as 40% below the actual number because some deer that are shot are not recovered owing to the rough nature of the terrain. Three State Forests - Beaumont, Rankleburn and Tapanui - now comprise the Blue Mountains hunting area which, for administrative purposes, is divided into thirty-five blocks ranging from 400 to 800 ha. Exotic timber crops, mainly Monterey pine but also Corsican pine (Pinus nigu), Douglas fir (Pseudotsuga rnenziesii) and Ponderosa pine (Pinus ponderosa), have been planted in less than a half of the area. New plantations are sometimes grazed by Fallow wlule young unthinned stands provide shelter. The mature pine forest lacks an understorey, which would provide cover and browse, so the deer seldom remain there. The greater part (over 7000 ha) of the State Forest, many of them steep, rugged and intersected by gullies and streams, is native forest in which Silver beech dominates over broadleaf (Griselinia littoralis), putaputaweta (Curpoderus senatus) and fuchsia (Fuchsia excorticatu). The scrub vegetation in the moist gullies is predominantly fern (Blechnurn sp.) or pepperwood (Winteru coloratu) or manuka, according to aspect. At higher altitudes this vegetative cover gives way to Snow tussock (Chionochloa rigidu). The altitude ranges from 30 to 1036 rn. Snow occurs at elevations over 450 m for about 10 days a year: heavy falls can benefit the deer by breaking branches and so presenting an abundance of browse. Pre- cipitation averages 900 mm/year. For mid-summer the mean daily temperature averages about 15OC and the equivalent mid-winter figure is 4OC (Baker, 1973).

About 90 km north-west of the Blue Mountains, the glacialvalleysassociated with Lake Wakatipu have long been important Fallow areas but sustained hunting over many years has resulted in scattered populations. The northeast side of the Lake and the Caples and Greenstone Valleys on the north-west side are ideal habitats for Fallow. Extensive river flats are surrounded by native forest, mainly of Southern beech (Plate 4). When the grassy river flats at about 350 m above sea level are under heavy snow the deer sometimes move higher up the valley sides where the steepness prevents deep drifts forming. In the spring they seek the new growth of Mountain flax (Phormiurn



colensoi) above the tree-line (Harris, 1967). About 20 years ago the excessive number of Fallow in the Caples Valley was reduced, not only by hunting but also by poison bait dropped from aircraft. In recent years the Fallow population in this valley has stabilized at a lower number which is likely to be maintained by recreational hunting. Fallow deer in both the Caples and Greenstone Valleys have also been subjected to spasmodic commercial hunting for venison in recent years (L. H. Harris, pers. comm., 1977).

2.3. ASIA Fallow deer have, as far as we know, never occupied the greater part of Asia: the most easterly records, from Iran, are of the Persian subspecies. Turkey is the only Asiatic country where the European Fallow deer exists in the wild and is probably the only country where this animal has lived continuously since it evolved.

2.3.1. Japan Fallow deer currently live in Japan only in five zoological gardens ( T. Ito, pers. comm., 1974). We have found no evidence of the presence of feral Fallow in this country although de Vos, Manville & Van Gelder (1956) listed the introduction of Fallow to Texas from Japan at an unrecorded date. If t h s information is correct the stock must have been captive animals. An alternative explanation is that confusion may have arisen with Sika deer (Cervus nippon) which is native to Japan and which also has been introduced to Texas.

2.3.2. Turkey Fallow deer are now very rare in Turkey. Only three populations are known, all located in south Turkey-in-Asia between the Mediterranean Sea and the Taurus Range of Mountains (Fig. 6). The total number was estimated to be approximately 100 in 1975 (Heidemann, 1976).

225 krn #+

Fig. 6. The distribution of Fallow deer in Turkey and the island of Rhodes

Fallow were formerly widespread in Turkey, particularly in the southern half of the country. They extended in a southerly arc from Maritza, in the extreme north-west of Turkey, in the European portion adjacent to Greece, to Erzutam and Patros in the east and north of Lake Van (Kumerloeve, 1967). In 1835 Fallow were common in the Kari Be1 and Chamlai mountains in the Taurus range, on the plains of the south coast and the adjacent foothills of the Taurus range and occurred in bushy scrub within a mile of Antalya (Danford & Alston, 1877). The species remained in the most westerly part of Turkey until 1860-70, being reported at several places north-east of Gallipoli on the Gulf of Zeros. To the southeast, on the opposite side of the Sea of Marmara, they survived until at least early this century. About 1905, a buck shot by Whitall 28 km from Karaboga established a place in the record books for the length of its antlers (Barclay, 1955).

Two of the present localities for Fallow deer are near to Antalya. Duzlercami Forest lies 17 km north-west of this town and Manavgat Forest lies 65 km to the east of Antalya. Both areas have similar habitats. Pinus brutiu forest covers over 60% of the land, maquis scrub accounts for over 30% and the rest is grassland (Swift & Holloway, 1967). The annual rainfall is about 1050 mm: the

78

mean temperature in January is about 8°C and in July 27°C. The deer mostly occur at heights between 100 and 900 m above sea level.

Within Duzlercami Forest, 1700ha were declared a protected area for Fallow deer in 1966 when only seven individuals were believed to survive there. Grazing by domestic stock had ceased in the area 4 years previously with the consequent encroachment of maquis species into the grassland and clearings among the pines. Most troublesome of the rapidly invading shrubs was Cistus laurefolius which is unpalatable to Fallow deer but other maquis species such as Quercus coccifera, Phyleriu media, Arbutus unedo and Pistacia spp. and Simylax sp. are browsed by the deer (Swift & Holloway, 1967). By 1975 the Fallow population was estimated to be between 60 and 80 (Heidemann, 1976).

Despite the similarity of habitat in Manavgat Forest, the population has fallen from estimates of 25-45 in 1966 to not more than 15 in 1976 (Heidemann, 1976), thus continuing the decline which has been occurring since they were hunted heavily between 1950 and 1960. Although here too shooting of Fallow is illegal, enforcement of the law is more difficult, there being 28 villages scattered within the 50 000 ha in which the deer occur. In Duzlercami Forest no villages lie within the Fallow deer area. Associated problems not encountered at Duzlercami but serious in Manavgat are the heavy grazing and browsing pressure by thousands of domestic stock in the maquis scrub zone, the Fallow deer’s main habitat (Swift & Holloway, 1967).

The third and most recently discovered population of Fallow deer in Turkey is in the vicinity of Pos Catalan near the Seyhan River and about 80 km north of Adana. In 1970, when it was declared a reserve for Fallow deer, a dozen animals were believed to be present: Heidemann’s more recent estimate is 18. The habitat is similar to the other localities, with Pinus brutia dominant but, as in the Manavgat Forest, there are considerable problems with domestic animals and poaching. The temperature here is very similar to the other two Turkish Fallow localities but rainfall is less, about 600 mm a year (Heidemann, 1976).


2.4. EUROPE Most countries in Europe have some free-ranging Fallow deer. We believe that the only exceptions are Albania, for which it has not been possible to obtain any information, Cyprus, Gibraltar, the Grand Duchy of Luxembourg, Iceland, Liechtenstein, Malta, the Principality of Monaco, the Republics of Andorra and San Marino, and Switzerland.

2.4.1. Austria Wild Fallow deer occur in only three localities in Austria (Fig. 7). Usually any Fallow that escape from parks are shot before they can establish a new population (K. Bauer, pers. comm., 1974). The annual shooting figures for the whole country reflect the comparative rarity of the species. Whereas about 160 Fallow and 170 Sika deer are shot, the number for Red deer is around 46 000 and for Roe deer (Capreolus capreolus) 220 000 (Anon., 1978).

Between the two World Wars, feral herds existed in the Leithagebrige in northern Burgenland and in the Dunkelsteiner Wald in central lower Austria (Amon, 1931). In 1935, over 450 Fallow were shot in Austria, mostly in these two areas (Fuchs, 1959) but by the late 1940s the populations in both these places had been exterminated as had almost all the deer in parks (Kerschagl, 1964).

Since 1932, when deer from Luneberg Heath in West Germany were introduced, Fallow have been present near the River Salzach, at Anthering, just north of Salzburg and close to the German border. Further releases of deer imported from Czechoslovakia were made in 1943, The deer range over 1000 ha of damp woodland in whch the main tree species are willow (Sulix sp.), Common oak (Quercus robur), lime (Tilia sp.), poplar (Populus sp.), alder (Alnus sp.), elm (Ulmus sp.), ash (Fraxinus sp.) and Norway spruce (Picea bbies). Not all the habitat is forested. Meadows and fields, specially created for the deer and planted with rape (Brassica napus) and oats (Avenu orientalis), are interspersed among the woodland, giving a park-like appearance to the land. As a further enticement to keep the deer in this area, and to encourage trophyclass animals for the


50 km Fig. 7. The distribution of Fallow deer in Austria

satisfaction of stalkers who shoot about 45 a year, supplementary feed is given. This provision of hay, oats, turnips (Brassica rapa) and maize (Zea sp.) satisfies the legal requirement in Austria that feed be provided for deer in winter. In this district the average temperature in January is -2°C and in July it is 18°C. The annual precipitation is 1270 mm, of which 200 mm falls as snow (W. Herbst, pers. comm., 1977).

The second locality where Fallow occur is near Horn, about 65 km north-west of Vienna, in northern Austria in country 200-500 m above sealevel. About 60 were present in 1972 (Dobschova, 1972). A similar number also occur in central Austria, near Mautern in Steiermark. They range over woodland, grass meadows and cattle pastures. Although conifers predominate in the woodland, with Norway spruce, European larch (Lark decifua) and pines (Pinus mugo and P.cembra) as the main species, there is some ash (Frauinus sp.) and sycamore ( h e r pseudoplatanus). The altitude varies from 700 to 1860 m. The annual rainfall is 1400 mm and the temperatures are very similar to those for Anthering (Phillips & Mutch, 1974).

2.4.2. Belgium In Belgium the Fallow deer seems never to have been a popular species. The only population of wild Fallow lives in the Ardennes region of south-east Belgium, just west of Rochefort (Fig. 8). Here, within the 1500 ha of the royal forest of Ciergnon, Fallow were introduced about 1850. The population in recent years has been maintained at about 100 or less (S. de Crombrugghe, pers. comm., 1973). The forest lies at an altitude of 200 to 300 m. The annual rainfall is about 800 mm. Mid-summer temperatures may range between 13 and 23°C and January temperatures range from just below to just above freezing point.

Fig. &The distribution of Fallow deer in Belgium


2.4.3. Bulgaria Wild populations of Fallow deer are widely dispersed in Bulgaria but with the greater number in the eastern half of the country (Fig. 9). They are being encouraged to increase, hunting being banned in all but a few areas. Much of the information presented here has been obtained from E. Clay (pers. comm., 1973) and Sartorius (1970).

- 55 hm

Fig. 9. The distribution of Fallow deer in Bulgaria

Probably the first introduction of Fallow to Bulgaria was in 1904 when a buck and two does were imported from Germany. They and subsequently more Fallow, were enclosed in the Kricim hunting reserve near the River Maritsa and to the east of Plovdiv. A herd of about 80 are still enclosed at Kricim: in all the other reserves the Fallow are free-ranging. As the herd increased, animals were taken to other localities where they were released and allowed to establish themselves in the wild. In 1973 the country's total number of Fallow deer was believed to be around 1000.

Probably the largest population of Fallow in Bulgaria is at Voden, near Kubrat, north of Razgrad in the north-east of the country where the Danube Plain broadens to form the Deli Orman plateau. The deer were estimated to number 350 in 1969, within 3000 ha of deciduous woodland. Fallow originating from Voden reserve have colonized areas to the west and east and northwards to the Danube which forms the boundary with Romania. To the west, herds are established in the woodlands extending towards the River Lorn. East of Voden they occur north of Novi Pazar and extend almost as far as the Black Sea, near Shabla.

About 145 km south of Voden, near Yambol, is the Yambolski-orman Forest where, in 1970, over 250 Fallow were estimated to live within 900 ha of woodland lying at 100 to 200 m above sea level. Further east, close to the Black Sea, Fallow occur in the Ropotamo reserve which covers 2700 ha in the Stranja Mountains district, south of Burgas and west of Mitschurin.

In central Bulgaria, on the southern side of the Balkan Mountains, Fallow are found in forest and scrub of the Stara Zagora plain. Further south they live in the hills between Khaskova and Kurdzhali, on the north side of the River Arda and near the Studen Kladenez Dam. West of the Stara Zagora plain, Fallow are found near Pazardzhik on the River Maritsa.

In western Bulgaria Fallow occur in the Vitoscha Nature Park south of Sofia. The reserve covers 10 000 ha, mainly meadows and pine woods, ranging from 500 to 2000 m in altitude.

The areas where Fallow deer occur in Bulgaria generally have January temperatures of -2 to f2"C and July temperatures around 22°C. The annual rainfall is usually about 500 mm in the east, increasing to about 650 mm in the west.


2.4.4. Czechoslovakia The earliest documentary evidence of the keeping of Fallow deer in Czechoslovakia is dated 1465 (NoiiEka, 1966) but apparently the species was not released until last century. Now Fallow are widely distributed in the wild (Fig. 10) and are kept in about twenty-four enclosed reserves (Sartorius, 1970; Velek, 1970, 1976). Recently the total population was estimated as. approximately 7000 (Kotrla & Kotrlg, 1977), an increase on the 1974 estimate when 2200 were thought to be free with a further 2900 in enclosed reserves (V. Mimra, pers. comm.). The enclosed herds are not described here: their history has been discussed by Flasar & Flasarova (1975).

100 km

Fig. 10. The distribution of Fallow deer in Czechoslovakia

Bohemia, in the west of Czechoslovakia, has the greatest number of Fallow, both free-ranging and enclosed. The main localities for wild herds lie south-west of Plzeil, close to the West German border, where three populations are established in an area stretching from west of the River Radbuza to east of the River Uhlava. The middle of these three populations is near Kydne, close to an enclosed herd. Fallow also occur west of PlzeK, near StEbo on the River Mk. South of Prague, in the Dobns area, is a further wild population but most of the Fallow in central Bohemia are enclosed. However, in Northern Bohemia there are four localities inhabited by wild Fallow. These are around M6lnik; to the north, against the East German border near Varnsdorf; eastwards near Jan Nisou close to the Polish Border and southwards, around D d r Kralovi. East of the River Elbe and southeast of Hradec Krilove' in Eastern Bohemia are two further populations but in the Southern sector of the province there is only one Fallow deer area, in the Luznice valley between Tabor and BechynB.

In Moravia, in central Czechoslovakia, there are six areas where Fallow occur. The most westerly is around Moravski Bud5jovice. Three of the localities lie between Brno and Ostrava, respectively near Blanska, Olomouc and east of Rerov, while another population lives east of Strahice. The sixth area where Fallow live is south of Bieclav, between the confluence of the Rivers Thaya and Morava, close to the Austrian border.

Within the region of Eastern Slovakia, the main wild population, numbering around 1000, lies north of Bratislava, in the M a l i Karpaty hills. To the north-east, between the Rivers Vih and Nitra, are two further populations, near Trencin and Zilina respectively. Further east Fallow deer live in the 12 000 ha hunting preserves near Topolcany and, east of the River Hron, near Levice.

The area south of Bievclav is one of the very few parts of the country lying below 200 m and probably has the only Fallow living in lowland pastures in Czechoslovakia. All the other habitats utilized by Fallow are in hill country between 200 and 700 m above sea level. For most of the Fallow deer areas the average temperature in January is about 4 ° C and in July, about 20°C. The annual rainfall is between 500 and 670 mm.

2.4.5. Denmark In Denmark wild Fallow deer are present on the Jutland peninsula, on four of the major islands, namely Langeland, Lolland, Funen and Zealand, and four very small islands (Fig. 11). Following


" 50 km

Fig. 11. The distribution of Fallow deer in Denmark

its extinction during the Last (Wurm) Glaciation, the species was re-introduced to Denmark in the thirteenth century. Documents from the reign of King Valdmars Jordebog and dated 1231 refer to Fallow deer on sixteen minor islands (Braestrup, 1952; Mdhl, 1961). Later, in Jutland and on the larger islands, deer parks were established. These included the Jaegersborg Deer Park on Zealand, on the northern outskirts of Copenhagen, which now holds about one third of Denmark's entire Fallow deer population. Although wild Fallow are widely scattered their total number is exceeded by that of enclosed herds. The total estimate of Fallow deer in 1970 was almost 4000, of which just over 2000 were park animals (Alex-Hansen, 1970). Only wild herds are considered here.

The history, particularly during the last century, of Fallow deer in Denmark has been discussed by Alex-Hansen (1945). He also recorded the estimated size of the populations of park and wild populations at 5-year intervals between 1945 and 1970 (Alex-Hansen, 1945, 1950, 1955, 1960, 1965, 1970). During th is period the free-living deer were lost from the island of Sarns$, from Kastrup on Zealand, and from Rosenvold and Sostrup in Jutland. Since 1970 the few Fallow then present at L&enholm, east of Auning in east Jutland, have gone although Roe and Red deer remain there. Otherwise the distribution of wild Fallow remains similar to that in 1970.

The largest population of wild Fallow in Denmark, estimated at over 400, live in the north of Zealand (Alex-Hansen, 1970). They occur in the 5000 ha of the Crib Skov, a very vaned forest of coniferous, beech and other deciduous woodland extending between Hillerdd and Esrum. An abundance of food and cover provide an ideal habitat for Fallow as well as Roe deer. South-west from Crib Skov, about 100 Fallow live on the wooded Bognaes peninsula which protrudes into the southern end of Roskilde Fjord. In the centre of Zealand, between Ringsted and Slagelse, a small herd of Fallow has been living in the woodland near SorQ Academy since 1965: about ten were present in 1970. They probably originated from Naesbyholm which lies a few kilometers to the south and where about eighty Fallow are present. Southeast of Naesbyholm, Fallow also occur between Fensmark and the coast. Within this area, some seventy Fallow live on the Gisselfeld estate with its arable fields, deciduous woodland which includes beech, birch (Betula sp.), oak, and conifer plantations. From here some Fallow have moved eastwards to the boggy terrain of Holme- gaards Mose (H. J. Degn, pers. comm., 1978). Immediately east of Gisselfeld and close to Faske, about twenty Fallow occur on the Lystrup estate. Wandering animals from here frequently move eastwards to the Vemmetofte district. Early th is century Fallow were reported to be very numer-


ous in this area, especially around Vemmetofte, Lyrup, Gisselfeld, Bregentved, Jomfuens, Egede, Beldringe and Knuds Skov: often 60-70 were seen at one time (H. J. Degn, pers. comm., 1978). Lying in the Kattegat, nearly 30 km north of Zealand, is the island of Hasseld which has an area of only 70 ha.; Fallow deer were introduced there on three occasions from December 1977 to Febru- ary 1979 when sixty-four were present. Eventually the herd will be maintained at 150-300 animals and managed for meat production (D. Clausen & E. Jensen, pers. corns.) .

On Lolland, four herds of free-ranging Fallow were reported by Alex-Hansen (1970). East of Maribo Fallow are enclosed in the Krenkerup park but in addition about twenty-five deer were living free, presumably having escaped. East of Rddby, about 100 Fallow occurred in the woods between Bremersvold and Kjaerstrup. A herd of similar size was reported to the north in the Christianssaede woods where beech woods, conifer plantations and neighbouring corn and sugar beet fields provide a suitable habitat. Between Oreby and Berritgaard, on the coast north-east of Maribo, about twenty-five Fallow were present.

The narrow island of Langeland has two populations of Fallow deer, around Tranekaer in the central part and further south, near Hjortholm Manor. In both places the deer live in areas of woodland and arable fields. Although some conifers are present the woodland is mostly deciduous with beech, oak, lime (Tilia sp.), alder (Alnus sp.) and hazel (Corylus sp.). Langeland may have held Fallow continuously since the thirteenth century although new stock has been taken to the island on several occasions. This is one of the islands mentioned in King Valdemars’ register as having Fallow, and bones dated as seventeenth century were recently excavated on the island, near Humble (J. Skaarup, pers. comm., 1978).

On Funen, Fallow also occur in two areas. Northeast of Ringe lies the Revnholt estate, the largest in Denmark, which covers 100 km2 of arable land and woodland. In 1945 wild and park Fallow were present but shortly afterwards the park was disbanded and since 1950 all the deer have been free-ranging. More than 100 were present in the autumn of 1978. The areas of woodland, varied in age structure and in species of both deciduous and coniferous trees, have rich understoreys providing plenty of cover and food for Fallow and Roe deer. The surrounding cornfields and road-side avenues of Horse-chestnut (Aesculus hippocastanurn) trees add to the food available. On the west coast of Funen, on the 480 ha of the Wedellsborg estate near Husby, live a herd of Fallow deer world-famous for their gold-medal class trophy antlers. On three sides the estate is bounded by the sea. The existence of a fence on the fourth side was the reason given for disqualifying the Wedellsborg antlers at the World-Exhibition of Hunting in Budapest in 1971, antlers from ‘enclosed’ animals not being eligible (Came, 1975). In fact, the fence is not deer-proof and the animals move freely through areas of beech woods, other deciduous woodland, parkland and arable fields. To protect the corn, some fields are surrounded by high fences during the appropriate months but after the harvest, sections of these fences are removed so the deer have access to the stubble. To maintain the quality of the heads, prevent damage to trees and encourage the deer to remain on the estate, supplementary feed is provided during the winter.

Off the north-east coast of Funen lies the small, wooded island of R o m 4 with an area of about 100 ha and a population of about 100 Fallow. On another small (250 ha) island, AebelQ, situated off the north coast of Funen, an unsuccessful attempt once was made to shoot the entire Fallow population (Sporon-Fiedler, 1930) but 40 years later the deer were estimated to number 250 (Alex-Hansen, 1970).

On Jutland there are a few small, scattered herds of Fallow, mostly on the eastern side of the peninsula. In the north, a herd lives in the mixed deciduous and coniferous woodland at BQrglum, south of Glimsholt. About 15 km north-west of Grenaa a small number of Fallow have been present in the Emmedsbo Plantation near the coast for over 40 years. In 1970, Alex-Hansen reported only two animals there and the present situation is not known. Near the coast at Palsgard, about 20 km south-east of Horsens, about thirty Fallow were present in 1970. In the Limsfjord, in northern Jutland, lies the island of LivQ with an area of just over 300 ha. Fallow deer were introduced for hunting in 1876 and established a population in the mixed coniferous woodland, the remnants of an ancient oak wood and on the heather (Calluna vulgaris) and juniper (Juniperus


commune) zone along the north-west shore (N. H. Lindhard, pers. comm., 1978). New stock was introduced in 1957 and twenty deer were counted a few years later (Alex-Hansen, 1965) but about 1970 the herd was much reduced when a number of dead deer were found. A buck and two does were liberated on the island in 1975 but this buck died in 1977: by April 1978 only three remained, two adult does and a fawn of unknown sex.

All of Denmark is low-lying, with a maximum altitude of only 173 m above sea level, and regional changes in climate are small. All the areas where Fallow deer occur receive an annual rainfall between 500 and 700 mm. The mean temperature in the coldest month, February, is 0°C and snow may persist for several weeks in winter. In July the mean temperature is 16°C.

2.4.6. Finland There is some uncertainty as to whether wild Fallow deer occur in Finland. As a result of information obtained from Finland in 1974, we reported that no wild Fallow lived in the country, but that there were two enclosed htrds, at Hattoo about 50 km west of Helsinki, and at, Kytaja near Hyvinkaa (Chapman & Chapman, 1975). However, R. Harrington (pers. comm., 1977) reported that the Fallow deer at Hyvinkaa, about 50 km north of Helsinki, are entirely free-ranging (Fig. 12). The herd numbered about fifty in 1973 and are descended from deer obtained from central Europe in 1938 (Ueckermann & Hansen, 1968).

- 175 hm

Fig. 12. The distribution of Fallow deer in Finland, Norway and Sweden

The mean temperature in July is about 17OC but in February it falls to -6°C. Annual precipitation is about 500 mm and in this southern part of Finland snow reaches a depth of 400 mm and covers the ground for 4 months of the year.

2.4.7. France Wild Fallow deer are more widespread in France than was recently believed by Chapman & Chap- man (1975). Although few in number, t h i s species features in the hunting plan of the Office

Distribution of fullow deer 85

National de la Cha.se in thirteen areas. These are: Aisne, Aube, Calvados, Oise, Yvelines, Essonne, Haute Marne, Val de Marne, Bas Rhin, Haut Rhin and Moselle - all in northern France, and in Landes and Tarn in the south (R. Coombe, pers. comm., 1977) (Fig. 13). The estimated total population in the spring of 1974 was only 700 (Anon., 1976b): most of these deer were in the departments of Bas Rhin, Haut Rhin and Moselle which constitute the Alsace - Lorraine region. The departments of Aisne, Oise and Somme, north of Paris, account for a smaller number and the remaining areas, for which no details are available, have very few.

2 155 krn

Fig. 13. The distribution of Fallow deer in France v

This predominantly northern distribution contrasts with that given by Rode & Didier (1935) when Fallow were reported in Alsace, Nivernais in central France, and the southern regions of CBvennes, Dauphine, the Alpes and the Pyrennees. Now there are no Fallow in the Alpes, nor in the adjacent regions of Provence or Cbte d’Azur in southeast France (E. Le Chatelier, pers. comm., 1977). Of the localities listed in 1935, it seems that only Alsace retains Fallow deer although possibly Cevennes does too if the Department of Tam was included in this region by Rode & Didier (1935).

Alsace - Lorraine probably has had Fallow deer at least since the sixteenth century (Stoll, 1969). Now they are found along the River Ill, a tributary of the Rhine, between Erstein and Guemar, which are respectively about 20 and 40 km south of Strasbourg. The deer population centres on the Illwald Forest although small groups have become established a few kilometers north of Ned, in the Marckolsheim -Schoenau forest, and since 1973 some have occupied parts of la Harth (Waechter, 1976). In 1958 a study was begun and censuses of the Fallow population have been taken regularly. The 1969 figure was 215, with a probable 5-10% margin of error (Stoll, 1969). The 1975 census gave 228 Fallow (Waechter, 1976) although that year’s estimate by the office National des Fortts was over 500. Waechter’s 1976 figure was 235 (pers. comm.).

The deer mainly stay within 9500 ha of woodland and meadows which are subject to flooding. There are 2800 ha of deciduous woodland with oak (Quercus robur, Q. sessilifloru), ash (Frauinus elutior) poplars (Populus spp.) and Norway maple (Acer platunoides) predominating. An understorey of hazel (Corylus avellunu) privet (Ligustrum europueum), dogwood (Cornus sunguinue), hawthorn (Crutuegus oxycuntha) and blackthorn (Pnrnus spinosu) provides thick cover. Over half this woodland forms the Illwald Forest and the remainder consists of small woods


scattered among the meadows and fields of the Reid Centre-Alsace (E. Heil, pers. comm., 1973). In this region the mean temperature in January is about 0°C and in July 19°C. The annual rainfall is about 700 mm.

In the department of Somme, Fallow deer occur east of Longueval, about 30 km northeast of Amiens. The deer occupy Delville Wood in the environs of the National Memorial which honours South Africans who fell in the first World War. Fighting took place here in 1916 and 1918 and at the end of the war only blackened tree stumps remained. After the new growth sprang up, the young woodland was colonized by Fallow deer (A. F. Dolby, pers. comm., 1977). The deer may have escaped from a park although in 1888 de Kerville referred to Fallow a little to the north, in Rouvray Forest, near Orival immediately north of Arras. where they were introduced in the 1860s. The deer multiplied and spread towards Rouen (Pennetier, 1905) and also became established in the Hallates forests and towards Le Havre. The Delville wood. lying within the Paris Basin, has a climate similar t o that of the Illwald Forest. Mean temperatures for January and July are respectively 2°C and 17°C and the annual rainfall is about 700 mm.

In the de'partment of Oise and south-west of Amiens Fallow are believed to live in small woods around Ivy-Le-Temple, between Beauvais and Pontoise (J. Jomier, pers. comm., 1977) and therefore not far from Cisors in which forest de Kerville (1 888) reported the presence of Fallow deer.

North-east of Paris in the department of Ainse between St. Quentin and Reims, a small number of Fallow live in the Samoussy Forest near Laon, having escaped from a nearby enclosure. Partway between this forest and Paris lies the remnant of the great royal forest of CompiCgne. This beech and oak woodland held Fallow before the first World War but the last one was killed about 1920 (J. Jomier. pers. comm.). Reed (1954) referred to the presence of a few Fallow in the forest so some park escapees may have been living there in in 1950s.

2.4.8. German Democratic Republic (East Germany) In East Germany, free-ranging Fallow deer are widely distributed, especially in the northern and central parts. (Fig. 14). In a recent study Siefke (1977) estimated the size of the main populations as well as showing the distribution of the species and the following is based largely on his work.

Fig. 14. The distribution of Fallow deer in the German Democratic Republic (after Siefke, 1977)


The number of animals seems to be increasing. In 1977 the total estimate was 11,600 and over 5000 were shot compared with only 300 twenty years earlier.

In the extreme north of the country several hundred Fallow deer live on the eastern half of the Baltic island of Rugen and on the Darss-Zingst peninsula. About 60 km to the southeast some Fallow have become established on the island of Usedom, on the border with Poland. lmmediately west of this locality Fallow are present in the Greifswald-Anklam-Uechkermunde districts.

The largest area of almost continuous distribution of Fallow deer is approximately bounded by Demmin. Neubrandenburg, Prenzlau. Eberswalde, Oranienburg, Neuruppin, Malchow Lube and Gustrow. Within this region. which includes the lake district to which Fallow were introduced in about 1755, about fifty populations are believed to be present: a quarter of these consist of several hundred deer. In the Neubrandenburg province alone the population was estimated at 3800. Also in the north of the country populations occur near Schonberg, and near Butzow, east of Rostock towards the River Recknitz. Most of the habitat consists of Scots pine (Pinus sylvestris) and beech (Fugus sylvatica) forests with neighbouring arable land on which the deer feed (Siefke, 1972).

Further west, near the border with West Germany in the Hagenow-Ludwigslust area nearly 1000 Fallow are present. To the south small numbers occur in the Osterburg area and west of Stendal in the Altmark region, to which the species was introduced in 1713. The Madgeburg province also has many Fallow, estimated over 2000, mostly in the eastern part.

The second largest area of almost continuous Fallow deer distribution is in the centre of the country, immediately south of Berlin and extending approximately from Potsdam, to Furstenwalde, Lubben, Dessau, Zerbst and Brandenburg with a north-west extension to Havelberg. In the Potsdam province over 2250 Fallow were estimated to be present. Further west, Fallow occurs in the Halberstadt-Oschersleben area. The habitat utilized by the deer in this region is mostly pine forest.

In the southernmost third of the country the Fallow deer areas are smaller and more scattered. They include Sondershausen near Erfurt, an area on the border with Poland, south of Forst and between the Rivers Nysa and Spree, and several localities near Cottbus. In another five areas, respectively near to Jena, Werdau, Grimma, Oschatz and Torgau, Fallow began to establish themselves in the wild during the period 1960- 1970 and further introductions have been made since then (Siefke, 1977).

All the areas are less than 300 m above sea level, except for Jena and Werdau which are a little higher. The mean annual rainfall is between 500 and 700 mrn and the maximum depth of snow in a normal winter is less than 400 mm. The mean temperature in January is between -2°C and 0°C and in July it is between 18" and 20°C.

2.4.9. German Federal Republic (West Germany) Although wild Fallow deer are widely distributed in West Germany, the main areas are in the north and west (Fig. 15). The species has been kept in parks since at least the sixteenth century but did not become well established in the wild until much later following escapes from enclosures. Subsequently some animals were released deliberately in various places. The population in 1967 was estimated to be over 20,000 with the greatest numbers in the states of Schleswig-Holstein, Lower Saxony and Westphalia (Ueckermann & Hansen, 1968) and nearly 1 1,000 were shot during the 1976/77 hunting season (Table 2).

The history of Fallow deer in Westphalia is typical of that of many wild herds in West Germany (Ueckermann & Hansen, 1968). Having escaped from parks or hunting enclosures, the deer found suitable habitats in which to live, especially in the oak forests. Fallow were first reported as being feral in Westphalia in 1883 in the Kottenforst district, following their escape from an enclosure at Kommende-Muffendorf. The populations present in the Hoxter and neighbouring districts, near the River Weser, have resulted from deer which came in the 1930s from a park near the spa of Bad Dridburg. About 40 km north-west of Hoxter, the Fallow established around Detmold, on the edge of Teutoburg Forest, owe their origin to deer kept in a nearby enclosure. Some gained their liberty in 1938 and the enclosure was destroyed in 1945. Northeast of Buren and close to the River Alme

88 N. C. Chapman and D. I. Qlapman

130 krn

Fig. 15. The distribution of Fallow deer in the German Federal Republic (after Ueckermann &Hansen, 1968).

Table 2 The number of Fallow deer shot in the German Federal Republic in 1976177 (see Anon., 1978)

State

Baden-WiIrttemberg Bavaria Bremen Hamburg Hesse Lower Saxony Westphalia Rhineland-Palatinate Saarland Schleswig- Holstein

Total

29 1 295

0 25

1109 3899 1293

0 1

3808

10721

Fallow occur near Diilmen this population is descended from deer liberated from a nearby park about 1890. Fallow also occur near Wittgenstein, in the Sauerland hills in the southeast of the state. In the north-west, the lowland part of Westphalia, a number of places not far distant from Munster have Fallow deer.

Although Schleswig-Holstein is but sparsely wooded compared with most of Germany, it was the first state of Germany to which Fallow deer, from Denmark, were introduced. The species is very well established in many districts, particularly in the eastern half of the state, including Flensburg, close to the Danish border; Eckernforde and Rendsburg; Oldenburg in the extreme east and Sedgeberg, Steinberg and Stormarn in the south (Ueckermann & Hansen, 1968; Sartorious, 1970). The Fallow deer populations in two other localities in Schleswig-Holstein have been studied by Heidemann (1973a). In one area he investigated the distribution of the deer resulting from ten animals released in 1939 near Schleswig. Their distribution is not continuous but the deer are still spreading and are present in many localities between Satrup, north of Schleswig, and Dannewerk which lies 3 km to the south. In an east-west direction they occur between Kappeln and Wittbek, a


distance of about 6 km. This distribution suggests a slow rate of dispersal but the deer have for long been subject to an annual culling programme. The main concentration of deer is around Bollingstedt where, in 1961-69, the density of the population was more than ten deer per 1000 ha (Heidemann, 1973b). The other area studied was southeast of Schleswig, where Heidemann (1973a) made detailed observations of the Fallow population near Salzau, about 30 km east of Kiel and near Selenter Sees. The deer occur mainly in beech woodland but also utilized mixed woods, grassland and arable fields with crops such as wheat and rape. Fallow have been well- established in t h i s area for at least a century: in 1885/86 over 800 were counted in the Plon district and today Fallow are still present near this town.

The adjacent state of Lower Saxony also has had an increase in the number and distribution of Fallow deer since the Second World War. The 1967 estimate of 7000 may have been an under- estimate (Ueckermann & Hansen, 1968). Populations are widely dispersed from the northern plains, particularly west of the River Elbe towards the North Sea and in East Friesland and around Oldenburg, t o the south-east where they occur near Goslar on the edge of the Hartz Mountains. In the central part of Lower Saxony one of the best known areas in which Fallow deer occur, as well as Roe and Red deer, is Luneberg Heath, south of Hamburg and between the Rivers Elbe and AUer. Twenty-thousand hectares around the villages of Undelok and Wilseder are a nature reserve, declared as such to protect the heather and juniper heathland. Although some parts are forested with conifers there are also oak and birch wood (Ross-Macdonald, 1971). Further east, Fallow occur in several localities south of Domitz and close to the River Elbe where it forms the boundary with East Germany. About 40 km south-west of Hannover Fallow also live on the Deister hills (Foldes, 1972).

In Hesse the main populations of Fallow deer are in the south, particularly around Erbach near the border with Baden-Wurttemberg, in the Darmstadt area near Frankfurt-am-Main and around Wiesbaden in the Taunus hills west of Frankfurt. A smaller area around Ziegenhain in central Hesse is also occupied by Fallow deer. The main localities for Fallow in Baden-Wurttemberg are also in the south. They are found within a considerable area around Donaueschingen and Stockach on both sides of the River Danube and extending south towards Lake Constance. A smaller area around Crailsheim, in the north-east of Baden-Wurttemberg also has Fallow deer (Ueckermann & Hansen, 1968).

In northern Bavaria Fallow occur in the vicinity of Schweinfurt but the main populations are further south, either side of the River Danube, particularly besween Donauworth and Dillingen. About 40 km to the east, Stammham near Ingolstadt, is another stronghold for Fallow deer. In the Rhineland-Palatinate Fallow are widely distributed but not numerous. Although none was shot during the 1976/7 season, in the previous season forty-six were killed, about twice as many as in most recent years (Anon., 1978). The main areas where Fallow occur are west of the Rhine, in districts around Koblenz; between Saarburg and the border with Luxembourg; around Birkenfeld, and near Germersheim on the Rhine.

Of the three smallest states, only Hamburg has any wild Fallow. Bremen, which lacks any natural woodland, has none at all and Saarland, although well-wooded, has Fallow only in two enclosures. The single animal shot there in 1976/77 may have escaped from a park (Ueckermann & Hansen, 1968; Anon., 1978).

Most of the Fallow deer habitats in West Germany are less than 100 m above sea level although a few, as in Bavaria and Hesse, are nearer 500 m. The mean temperature in January in all the localities lies between -4 and 2°C and in July 16 to 18°C. The mean annual rainfall varies between 600 and 1000 mm according to the district, with the Schleswig-Holstein Fallow deer localities at the upper end of this range.

2.4.10. Greece There are now no wild Fallow deer on the mainland of Greece, the only Fallow there being two small enclosed herds at Karpenissi and Amfiklia, both in Central Greece. In 1878, de Heldreich reported the deer only from the Manina forest, in Akarnania, which extended westwards from the


river Akheloos as far as Catouna, in hilly country north-west Lake Trikhonis. He noted that they were not abundant and doubted whether they would survive much longer.

Nevertheless, Greece does have Fallow deer, on the island of Rhodes in the Aegean Sea (Fig. 6). Legend claims that the deer were brought from Asia Minor to stamp out snakes. They were introduced by the Knights of the Order of St John of Jerusalem who gained possession of the island early in the fourteenth century. The knights, who were keen hunters, preserved parts of the island as deer forest. During the period of Turkish rule (1522-1912) the deer were exterminated but were reintroduced by the Italians during the early part of their rule (1912-1945) (Dicks, 1974). Whatever their origin, the deer became the symbol of the island as exemplified by the nineteenth century bronze statues of a buck and a doe on columns either side of the Mandraki harbour. Captive deer are kept nearby, in enclosures against the mediaeval walls of the town of Rhodes.

The wild deer, estimated at 300-400 in 1973 (D. Sideridis, pers. comm.) are mainly on the Profita range of hills, most of which are between 220 and 400 m above sea level but with the highest point, Mount Elijah, over 700 m. The deer occur on the forested slopes where the main tree species are Pinus brutia, P. halepensis, Cupressus sempervirens, C. horizontalis, and various oaks including Quercus aigelops, Q. coccifera, Q. infectoria and juniperso(Juniperus macrocarpaA J. phoenica). The average temperature on the island in mid-summer is 26 C and in mid-winter 9 C. The annual rainfall is about 870mm but practically no rain falls from May to October. Frost seldom occurs and snow is even rarer (Dicks, 1974).

2.4.1 1. Hungary Although wild Fallow occur in only a few areas of Hungary (Fig. 16), this country is famed for trophy antlers (Sartorius, 1970; Szabolcs, 1975). The number of Fallow deer is estimated to be approximately 4500: 2143 were shot in 1977 (Berdlr & Mltrai, 1978). The majority of the largest antlers are from Gyulaj, a national forest and hunting area which is fenced but extends to over 7000 ha. The forest is near Tamlsi, about 50 km south of Lake Balaton, at an altitude of 100- 200 m. The main tree species are the Turkey oak (Quercus cem's) and hornbeam (Carpinus betulus) (V. Anna, pers. comm., 1977).

In the centre of Hungary, east of Budapest and near KecskemCt and Szolnok, about 600 Fallow deer range over the 15,000 ha of the unfenced hunting preserve around Pusztavacs. The deer were introduced here late last century and now each year over 200 are shot. About half the area is agricultural land but the rest is forest. Acacia (Robinia pseudoacacia) is the dominant tree (64% of the tree cover) but there is.also oak (19%) and other species including poplar and pine. The shrub layer of the forest, mainly privet, hawthorn (Oataegus sp.) and Ground elder (Aegopodium sp.), is much used by the Fallow deer for shelter and browsing (M. Papp, pers. comm., 1977).

h

v--- Fig. 16. The distribution of Fallow deer in Hungary

- 60 krn

Distribution of jallow deer 91

Further east is the 8000 ha forest near Gyula, between the river Koros and the Romanian border, where wild Fallow deer also occur. The terrain is less than 100 m above sea level and is flat, fertile land on the edge of the Great Plains. At the northern end of those Plains, within the pusztu or grassy prairies, Fallow occur on both sides of the river Tisza in the vicinity of Taktakenez and Taktaharkany.

Two other localities for wild Fallow are close together near Telki and Gyarmapuszta. These places lie between Budapest and Komrirom, on the western side of the Danube Bend and on the long Veites Plateau which supports some ancient forests. This is one of the higher parts of the country, lying between 200 and 500 m above sea level. In the south-west of Hungary, in La'bod near the border with Yugoslavia, about 300 Fallow are present (I. Heltay, pers. comm., 1978).

The climatic conditions in all the areas inhabited by Fallow are similar as there is but slight variation within the whole of Hungary. For example, there is only 3°C difference in average annual temperature between the north and south of the country. The mean temperature for the whole country in January ranges from -4" to - 0.5" C: the north-east, that is the Great Plains including the Taktakenez area, is the coldest for there is no break to stop the severe snow, yet here the total precipitation is the lowest; under 500mm. The mean July temperature on the Great Plains is about 23°C and a few degrees less in the central and more western parts of the country. Annual rainfall is about 600 mm for most of the areas where Fallow deer occur: snow may persist for considerable periods, sometimes for more than 30 days a year at Pusztavacs.

2.4.12. Italy Wild Fallow deer occur in several areas of Italy within state-owned forests or hunting reserves (Fig. 17). Some of these are fenced: others are surrounded by marshland or roads which act as barriers and prevent the movement of large numbers of Fallow, although a few move beyond the forest boundaries. The total population in 1973 was thought not to exceed 1000 (P. L. Florio & A. F. Pratesi, pers. comm., 1973).

Most of the localities are in central Italy. The exceptions are near Trieste in the north-east and in the state-owned Forests of Sila near Catanzaro in the Calabria region in the foot of Italy. The latter region is a table-land with an average height of about 1300 m which supports dense woods of oak, holly (Ilex sp.), laurel (Prunus sp.) and myrtle (Myrica sp.). Snow may remain in this area from November to March. The Fallow reported in various localities within 25 km of Trieste originated from Yugoslavia where some were released near the border with Italy in 1971. The numbers are small and poaching seems to prevent much increase in the population (Calligaris, Perco & Perco, 1976).

The other localities lie fairly close together and are all north of Rome but south of Bologna. North of the capital, near Terni, in the Umbria region, Fallow inhabit the state-owned forests of Monte Pegha and also some private hunting reserves. This is a hilly region, mostly between 200 and 2000 m above sea level, where the mean temperature is about 2OC in January and about 25°C in July.

West of Umbria lies the Maremma, a vast expanse of flat land in the regions of Tuscany and Latium. Here some of the deciduous and pine woodland in the Foreste Demaniah Camaldoli some 80 km from Arezzo, holds Fallow but their numbers were much greater earlier this century. Within the province of Pisa Fallow live in the hunting reserve of Migliarino. To the south, some occur in the state-owned forest of Feniglia, in the province of Grosseto, and in the hunting reserve of Sant' Agostino in the province of Viterbo. Some of the thickly wooded Appenine hills, namely the state- owned forests around Pistoia, Lucca and Florence, also hold populations of Fallow deer. Towards the east coast they occur in forests near Forli and on Mount Catria, 80 km inland from Ancona.

Formerly the ltalian islands of Sicily and Sardinia both had Fallow deer. Two centuries ago Cetti (1 774) wrote a careful description of the Fallow deer on Sardinia. At that time, Fallow were widely distributed over the island but were especially numerous on the Plain of Sindia and at least 3000 were killed each year. During the present century, after the second World War the population was thought to be between 150 and 300. Early in the 1950s, 30-50 were regularly seen in the


100 km Fig. 17. The distribution of Fallow deer in Italy

Castiadas Hills, east of Cagliari in the south of the island. After legal protection was lifted the number declined rapidly and they were thought to have been exterminated by 1957 (Reinhardt & Schenk, 1969). However, an animal was seen in the Castiadas hills in August 1960 and between 1966-69 four females were seen on six occasions in another area (Reinhardt & Schenk, 1969). Hunting was still permitted and it seems that wild Fallow are now extinct on Sardinia (Massoli- Novelli, 1976). A small number of captive Fallow have been introduced to the island, from Tuscany and Yugoslavia, and are maintained in two enclosures, at Marganai, near Iglesias and near Nuoro (R. Massoli-Novelli, pers. comm., 1976). It has not been possible to ascertain when Fallow became extinct on Sicily.

2.4.13. Netherlands Wild Fallow deer live in several places in the Netherlands in addition to a number of extensive but fenced forests, most of which are situated in the central part of the country (Fig. 18). They are widely kept in parks but any escapees are not generally welcomed in the surrounding countryside so are usually shot before a feral population is established. Most of the information given here has been supplied by Dr. W. A. Weyland (pers. comm., 1974).

The earliest date of introduction of Fallow deer to the Netherlands is uncertain. There may already have been some in parks when, early in the seventeenth century, Prince Maurits imported 100 from England and released them on the dunes near the Hague. Twenty years later the deer


h' Fig. 18. The distribution of

50 km

' Fallow deer in the Netherlands

were killed as thelevel of damage to the vegetation was unacceptable. In 1647 Willem I1 introduced some Fallow to Hof te Dieren near Rheden but this population was also shot, although not until the end of the eighteenth century.

Another introduction was made by Willem 111, about 1880, when some Fallow were put into an enclosure within the royal forest at Het Loo. In 1912 the deer were released into the forest itself but now the woodland has been divided into two by the busy Apeldoorn-Amersfoort road. Both sections of the forest are fenced. The larger section (6000 ha) holds about 200 Fallow, probably the largest forest population of Fallow in the Netherlands. Only about twenty Fallow live in the 3000 ha section.

A few kilometers north-west of the Het Loo forest, and near Elspeet, is another forest where Fallow, originally from Het Loo, became established about 1915. In 1973 about fifty Fallow lived in the 300 ha of the fenced Noorderheide section of the forest. The rest of the forest, I200 ha comprising the Elspeterbos and Elspeterstruiken sections, is also fenced but for more than 1 km the fence is only 1 m high and therefore no barrier t o a Fallow deer. About eighty deer live in this section but some do leave the forest and would probably establish themselves in the neighbouring part of the Veluwe if given the opportunity. The Game Department of the Ministry of Agriculture each year issues permits for any escapees to be shot.

In the 2100 ha of Deelerwoud, north of Arnhem, Fallow were released about 1915. Before the second World War the population reached 600 but now the three fenced sections of the forest together hold about 200. When fences were damages in 1945 some deer escaped and established a herd in the Hoge Veluwe, a forest of 5000 ha, but the last animal was shot about 1954.

To the west, and just north of Ede, Fallow deer have existed since before 1940 in the De Valouwe forest where about twenty-five deer are enclosed in 500 ha. A few kilometers away wild Fallow have been living near Leersum, since some escaped from nearby parks in 1963 and in 1968.

Several of the present free-ranging herds live on dunes on the west coast of the Netherlands. In Zeeland, Fallow have been living wild since some escaped from an enclosure in 1968. A small herd is found near Oranjezon, between Domburg and Oostkapelle, ranging over 350 ha of dunes covered by Marram grass (Ammophila arenaria) and 5 0 ha of conifer plantations. Also in the late 196Os, more Fallow escaped and established a population just to the north, on Schouwen, around Haamstede. North of Leiden, Fallow have been living in the 1200 ha of Kennemerduinen since about 1958 when a few park escapees arrived there. The population in 1974 was estimated to be twenty. In 1972 a few animals from this herd moved into the dune area to the south.

All the forests and dunes experience similar temperatures: the average in January is 2°C and in

94

July about 18'C. Rainfall does not exceed 800 mm. The woodland areas in the interior of the country are between 100 and 200 m above sea level but the dune habitats are 0-20 m below sea level.


2.4.14. Norway The date of the first introduction of Fallow deer to Norway is uncertain. Some hides were transported to England in 1305-07 but may have been in transit from Denmark. Captive Fallow were recorded in a park on Kalvo island, in Hogs Fjord on the south-west coast and east of Stravanger, in 1645 but in 1674 they were transported to Larvik. Some were also introduced to Malmo, a small island on the eastern side of the entrance to Larvik Fjord, for it is recorded that wolves (Canis lupus) killed all the Fallow deer there in 1705-06 (J. A. Pedersen, pers. comm., 1973). The predation by wolves on 'the common deer' is also mentioned by Pontoppidan (1755). As he goes on to discuss Red deer, Roe deer, elk (Alces alces) and reindeer (Rangifer farrandus), he presumably regarded Fallow as the common deer. He said that these deer were formerly more frequent in the districts of Bergen and Tronheim but that many had been devoured by wolves, although some did remain on the mainland and on some islands.

In the first few years of this century several liberations of Fallow were made. Three deer from Denmark were released on Raud island, in Oslo Fjord. Although fifty were present in 191 1, subsequently the population died out. Similarly, Fallow from Hanko released at Hurdalen in 1903 also became extinct in a relatively short time. Those liberated on SkorpQ island in 1904 swam to the larger island of Strdnen. In 1909, one male swam to the neighbouring island of Rdttingen (Collett, 191 1-191 2) and the Strdnen herd died out after a few more years.

Today the only free-ranging population of Fallow deer in Norway live on Hanko, an island which lies on the eastern side of the entrance to Oslo Fjord (Fig. 12). The island is well-wooded and has a maximum length and width of 4.5 and 1.5 km respectively. The herd originated from seven deer introduced from Denmark in 1901 and 1902. By 1936 the herd numbered 300 but was much reduced during the second World War. About 100 were present in 1968 but presumably culling subsequently took place as the number in 1972 was approximately forty (J. A. Pedersen, pers. comm., 1973). This part of Oslo Fjord has relatively mild winters, warm summers, a moderate mean annual precipitation of 700 mm and the mean number of days with snow cover is 64 during the months of November to April. Although a maximum snow depth of 800 mm has been recorded, the mean depth is less than 100 mm. A minimum temperature of -26'C has been reported but the mean temperature in January is -3'C and in July, 17'C. A climate exactly like this is found in no other part of Norway (T. W. Johannessen, pers. comm., 1976). The Fallow deer are given supplementary feed in winter.

Deer taken from Hanko in 1923-24 to the Fitzoehus estate near Larvik, south west of Oslo Fjord, established a herd which is maintained at about sixty head, but these animals live in a fenced park of 180 ha (J. A. Pedersen, pers. comm., 1973).

The attempts to establish free-ranging Fallow elsewhere in southern and western Norway have failed: it is said that they are unable to tolerate the deep snow which covers much of the country in winter (J. A. Pedersen, pers. comm., 1973) and which remains for at least 120 days in many places.

2.4.1 5. Poland Fallow deer were introduced to Poland probably in the seventeenth century and are now established in the wild in many areas, particularly in the western half of the country. More recently introductions were made in some central and eastern districts (Haber, Pastawski & Zaborowski, 1977; Oko & Wodek, 1978) and Fallow are now present in at least thirty of the forty-nine voivodeships (Z. Jaczewski, pers. comm., 1978). Figure 19 represents the distribution of Fallow in 1972 according to Oko & Wodek (1978) who estimated the Fallow deer population to be about 2600 in 1972.

Fallow occur on the Lublin Tableland, or East Polish Plateau, at about 230 mm above sea level,

Distribution o f fallow deer 95

- 70 km

Fig. 19. The distribution of Fallow deer in Poland (after Oko & Wodek, 1978)

and in the tysogory, an upland area, 600 m high, in the Kielce - Radom - Tomasz6w Mazowieki area of the Little Polish Plateau.

In the centre of the country, with its flat expanses of fields and forests, Fallow occur north of Lbdi. Further north, they are reported in many localities east of Bydgoszcz and north-west of Olsztyn but are absent from the northeast corner which includes Bialystok and part of the Mazurian Lakeland. East of Warsaw Fallow live in the Wegrow and Minsk Mazowiecki district and, further east, beyond Siedlce. In the Szczecin voivodeship in the north-west of Poland, the presence of Fallow is doubtful except in a small area between Chojna and the River Odra which forms the boundary with East Germany. From near Pozna'n Fallow are scattered in woodland almost as far north as Szczecinek and southwards towards Wrodaw. South-west of Pozna'n, towards the country's border, they are reported in several districts, especially south-west of Zielona Cora and around Wegliniec. Herds also live in woodland southeast of Wrockaw, east and south of Opole to Katowice and east of there, in the vicinity of Pszczyna.

Most of the habitat utilized by Fallow in Poland is mixed woodland with meadows or arable land nearby. About 35% of the area they occupy is pine-oak forest with Scots pine, Common oak and Durmast oak (Quercus petraeu) as the main species with some Silver birch (Betulu verrucosa), beech, Alder buckthorn (Frangula alnus), Norway spruce and Padus sp. A further 29% of their habitat consists of a pine-bilberry association with Scots pine and Bilberry ( Vuccinium myrtillus) as the dominant species but with some birch, larch, beech, juniper, rowan (Sorbus uucupuriu) and hornbeam. A similar percentage (26%) of the habitat is composed of oak-hornbeam woodland with Common oak as one of the dominant species. Pines, beech, birch, Small-leaved lime (Tilia cordufu), aspen (Populus tremulu) and hazel also occur within this habitat. The remaining 4% of the Fallow deer's habitat consist of wet forest with alder (Anus glutinosu) as the dominant tree. (Oko & Whdek, 1978; Z. Jaczewski, pers. comm.).

Most of the Fallow deer habitats are less than 200 m above sea level with some of the southern

9 6

localities a little higher. Temperatures and rainfall are similar for most of the areas where Fallow deer occur. The average temperature in January is between -4" and 0°C and in July, 17-19"C, whilst the annual rainfall is 500-600 mm. Fallow are absent from the northeast of the country which has a more severe climate.


2.4.16. Portugal In Portugal, Fallow deer are not well established in the wild. Although escapees are occasionally seen in the vicinity of parks,they apparently have failed to form populations. The largest enclosure, at Mafra, north-west of Lisbon and near the coast, has about 700 Fallow. Three other enclosed herds are found east of Lisbon: near Arraiolos, at Mora and at Vila Vigosa near the Spanish border. A fifth enclosure is at Alacer de Sol, southeast of Lisbon (J. F. Bugalho, pers. comm., 1976).

Sometimes Fallow are seen crossing the Spanish border in three areas (Fig. 20). Adjacent to the Spanish province of Badajoz they are seen north and south of Barrancos and, further north, in the vicinity of Campo Maior and Castelo de Vide. The third area is adjacent to the more northerly Spanish province of Caceres, where Fallow are occasionally sighted north of the River Tagus, towards the Sierra de las Mesas(J. F. Bugalho, pers. comm., 1976). All these areas have a low annual rainfall, usually under 70 mm. In January temperatures are rarely below 8°C and in July are at least 2OoC, often 30°C.

r-9 Spain

Fig. 20. The

13, -- i V

130 km

distribution of Fallow deer in Portugal and Spain

Unconfirmed reports of Fallow deer tracks in open conifer woodland on the sand dunes south of Ovar, which is south of Porto, were received in 1974 (C. B. Gooch, pers. comm.).

2.4.17. Republic of Ireland The Forestry Division of the Department of Lands have conducted a survey which indicated that Fallow were breeding in thirteen of the twenty-six counties within the Republic (Fig. 21). Much of the information given here is taken from the results of this survey and from Crichton (c. 1974). The total population for the Republic plus Northern Ireland was estimated to be approximately 3000 of which 85% lived in or had access to 30,000 ha of State Forests within the two countries (F. Mulloy, pers. comm., 1970, 1974).

The State forests are generally small, mostly less then 600 ha, but the total area they cover in the Republic is increasing by about 10,000 ha a year. Although 96% of the State woodlands are


Republic of Ireland

100 km Fig. 21. The distribution of Fallow deer in Ireland (after Crichton, c. 1974)

coniferous, a high proportion are young plantations which still have an understorey of mixed vegetation which provides food and shelter for Fallow deer: often a belt of deciduous trees borders at least one side of a plantation. In many places the deer also feed on neighbouring agricultural land. The most frequently grown conifers are Sitka spruce (Picea sitchensis) and Lodgepole pine (Pinus conrorta) which together account for 70% of the trees planted: other pines, larches and firs make up most of the remainder. Of the few hardwoods that are grown by the State, beech and birch are the most important. In addition, most of the 24,000 ha of woodland in private ownership in the Republic is deciduous (McCracken, 1971).

In the south of the Republic, Co. Waterford has Fallow through much of the county except the eastern corner. The main localities are the woods south of the River Suir near Kilsheelan and Clonmel, along the River Nier, north of Cappoquin and near Villierstown. Across the border in Co. Tipperary Fallow are also widespread. They live in plantations on the slopes of the Knockmealdown Mountains near Clogheen and Ballyporeen, the Glen of Atherlow where the valley and lower slopes of the Galty Mountains are forested, and smaller woods around Bansha, Dundrum, Rear Cross, Newport, the Silvermines Mountains and near Kilcooley Abbey, close to the border with Co. Kilkenny.

On the north side of the River Shannon, in Clare, Fallow occur in the woods which are scattered in the Bunratty to Tulla area. Due north from there, they are found on either side of the border between Clare and Co. Galway, in the vicinity of Lough Cutra. Fallow deer also occur in a number of other localities in eastern Galway including the plantations around Derrybrien on the Slieve Aughty Mountains and the neighbouring but lower-lying woods at Kylebrack, Woodford and Portumna. Portumna Forest, situated on limestone soil at the northern edge of Lough Derg, supports a high density of Fallow deer: 370 deer were estimated to live within 560 ha (Mulloy, 1970). The deer have access to the whole forest and the golf course within it but fences have been erected to prevent their entering neighbouring privately-owned fields. Further north they are also reported around Ahascragh, both sides of the Clonbrock River and near Ballygar.

In the central lowland part of the Republic, the counties of Offaly and Leix have Fallow in many localities. In the west of Offaly they occur near Birr Castle and also Clonfinlough: in the centre of the county the main areas are north, west and east of Tullamore. The latter two areas are contiguous with the Fallow localities in Leix, namely Clonaslee, the Slieve Bloom Mountains, Camross, Mountrath, Ballyfin, Ballybrittas and Stradbally. In the east of the country Fallow are


widely dispersed in Co. Wicklow, a county of high relief and little arable land where the Wicklow Mountains occupy a large proportion of the county’s area of 2000 km2. Fallow have been present in this county at least since 1244 (Deane, 1972). The forested mountain slopes provide a habitat for the Fallow. On the western side of the Mountains they occur from the Kiltegan district northwards to the Lackan Reservoir. East of the Reservoir they extend across the mountains towards Delgany and southwards from there through the woods around Carrick Mountain and Wicklow. On the southern border, and across it into Co. Wexford, Fallow occur south of Aughrim.

Co. Dublin is without herds of wild Fallow on the mainland but Lambay Island, 3 km off the coast from Portrane, has had Fallow since the species was introduced by Count Considine in 1888. The herd, maintained at about thirty and mostly of the black variety, roam free over the island whch has an area of approximately 1000 ha (Lord Revelstoke, pers. comm., 1974).

In Westmeath, Fallow occur in the Central part of the county, north and south of Mullingar. In Co. Roscommon the only Fallow recorded in the Department of Lands survey were those near Boyle, a feral herd long having been established there following escapes from the former Rockingham Park. The adjacent county of Mayo also has only one Fallow deer area, on the slopes of Partry Mountain close to Lough Mask. In the north-west of the Republic, Co. Sligo has Fallow in the woodland between Collooney and Lough Gill and on the other side of this Lough, especially in Slish Wood. In the north of Co. Monaghan, Fallow occur in the Glaslough district, on the border with Co. Armagh and Tyrone which are in Northern Ireland. In the south-west of Co. Monaghan, on the border with Co. Cavan, they occur in the woods between Rockcorry and Cootehill.

Although occasional sightings are reported, Fallow are not frequent in Co. Kildare, Meath, Co. Longford and Leitrim. In the southeast of the Republic records are very few for Co. Wexford, Co. Carlow and Co. Kilkenny. No Fallow are reported from Co. Donegal in the north-west nor from Kerry, Co. Limerick and most of Co. Cork in the south-west. A few do occur on the eastern boundary of Co. Cork, south of Tallow. Although Fallow are still widely dispersed in the Republic of Ireland, there seems to have been some reduction in their overall distribution since Whitehead (1 964) reviewed the situation and related the presence of wild populations to park herds. For instance, there were then small numbers in the Killarney district of Kerry and in Co. Donegal but none is reported now.

Most of the Fallow deer habitats in the Republic of Ireland are less than 200 m above sea level and even those on mountains are below 600 m. The annual rainfall in most of the localities is between 750 and 1000 mm but may reach 1500 mm in Co. Waterford, Co. Wicklow, Co. Galway and Co. Sligo. Snow is rare except on the Wicklow Mountains where the slopes inhabited by the deer may have snow on 10-20 days a year. The average temperature is around 5°C in January and 15°C in July in all the Fallow localities.

2.4.18. Romania Fallow deer are widely distributed throughout Romania. Wild populations are established in at least sixteen areas, mainly in the peripheral plains and in the plateaux which together comprise two-thirds of the country, but also at some higher altitudes in the foothills of the Carpathians (Fig. 22). The wild Fallow population was estimated to be approximately 3800 in 1975 (I. M. Bodea, pers. comm.). As in most European countries, t h i s is very few compared with the estimates for other species of deer. In 1968 Cotta estimated the populations to be: Roe 180 000; Red 27 000; Fallow 2700 with a further 850 Fallow within three enclosed parks (Sarlota, Valea Lunga and Hateg) in the west of the country, near the River Muresul. The distribution outlined by Cotta (1968) gives more localities for Fallow deer than those shown by Bunesco (1959) and the following is based on the former survey.

The largest populations of wild Fallow, possibly around 600, occur in the Tisza Plains which occupy the north-west corner of the country, comprising mainly the provinces of Crisana and Maramures. Much of this flat or undulating region is forest steppe vegetation, having grassland with scattered trees. Although much of the native vegetation has been removed for agriculture an oak forest extends the length of the Plains, (Matley, 1970).


Fig. 22. The distribution of Fallow deer in Romania - 80 km

In the northeast of the country, on the Moldavian Plateau, populations of Fallow are found neat Suceava, Iasi and Bacau. Further south, Fallow occur near Calati. The southeast corner of Romania, between the River Danube and the Black Sea, forms the Dobrogea Plain. The northern part is tableland supporting some oak forest: about 200 Fallow were believed to be established in this region.

Within the Great Wallachian or Romanian Plain, which occupies a broad belt of southern Romania, Fallow are found in four areas. North of Bucharest about 300 were estimated to be present and further north, near Ploesti a small herd existed. In the central part of the Plain, in the Arges district, about 200 were reported whilst further west, in the Oltenia region, the population was nearly 500. A large forest, predominantly oak, extends across the northern part of the Plain and some of the Fallow deer localities lie within this area, but elsewhere on the Plain, much of the natural forest steppe vegetation has been cleared for crops.

In central Romania, Fallow occur in three areas in the Transylvanian Plateau, which comprises tableland, low hills and undulating plains supporting mixed forests of oak and beech. In places this native vegetation has long been cleared giving a steppe-like landscape. Small herds of Fallow are found near Cluj and Tirgu Mures and also further south, near the River Otlul, towards Brasov. The Transylvanian Plateau is surrounded on three sides by the Carpathian Ranges which average 1300 m. The lower slopes support mainly beech forests which give way to mixed beech/conifer woodland at higher altitudes. In the foothills of the Southern Carpathians lies Hunedoara where a wild population of Fallow deer is established. The district is equidistant from the enclosed herds in the Valea Lunga and Hateg Parks.

Romania generally has a humid continental climate with hot summers and cold winters. The figures for Bucharest, near a Fallow deer area, are fairlyotypical of the other deer localities. The average January temperature is -2°C but in July it is 23 C. The Tisza Plains and Transylvania are slightly cooler. The annual rainfall in all the localities is between 500 and 700 mm and snow may persist for a considerable period, for example, SO days near Bucharest.

2.4.19. Spain Although free-ranging Fallow deer are present in several provinces of Spain (Fig. 20) there are few large concentrations, but they appear to be increasing and becoming more widespread (Hingston, 1975). They are more widely distributed than reported in 19 14 when Cabrera knew of them only in Caceres province, and in 1893 when Chapman & Buck reported wild Fallow in a few places in the neighbourhood of royal estates in the vicinity of Madrid.


The main localities where wild Fallow occur include five national reserves. In the north-western province of Oviedo a herd occurs at Sueve and another lives further east, at Quinto Real in Navarra province. In the more central part of the country, they are found in Serrania de Cuenca and further south at Lugar Nuevo and Cazorla-Segura, both in Jaen province (J. de la PeRa Payl, pers. comm., 1975).

Probably the largest and most studied population of Fallow in Spain is that, numbering more than 1000, at the Cot0 DoHana Biological Station Reserve in the south-west, between the Gulf of Cadiz and Seville, in the province of Huelva (Alvarez, Braza & Norzagasay, 1976). The Reserve comprises 45 000 ha of shore, dunes, woodland, scrub, and marsh and mudflats of the delta of the River Guadalquiver. The Fallow deer were introduced in 1917 (Sartorius, 1970) by the Duke of Tarifa and by 1924 they, together with Red deer, Wild boar and Northern lynx (Felix lynx) were being hunted by Alfonso XI11 (Fernlndez, 1975).

Unlike the Red deer which live in the scrub zone, the Fallow deer mainly frequent the marsh (Plate 5), the meadows and the Cork oak (Quercus suber) woods between the marsh and the scrub. The marshland, Las Marismas, is subject to great seasonal variation. In winter 40 000 ha are under water with only small islands and two areas of mudflats above the water level. In spring, as the water disappears, a dense carpet of sedges grows but in the hot, dry summer the area, apart from a few pools and mudholes, becomes a desert of cracked earth although the series of lakes beyond the marsh, in the woods and scrub, retain their water. With the arrival of the autumn rains, the dry mud turns into a quagmire in which Fallow deer have been known to perish (Ferna'ndez, 1975). The meadows on the edge of the marsh are wet in winter but become dry in summer and the Fallow fawns are born there, amongst the reeds. The park-like clumps of Cork oaks are remnants of a once great forest and form a discontinuous border between the marsh and the scrub (F. Alvarez, pers. comm., 1975). Attempts to introduce Fallow in the estates to the north of Cot0 Dofiana were unsuccessful because of poaching (Valverde, 1960).

The Cot0 DoRana is one of the lowest-lying areas of Spain, less than 100 m above sea level, but the other localities where Fallow deer occur are mostly at an altitude between 500 and 1000 m. The annual rainfall at DoAana is between 300 and 500mm: the other locations receive 500- 1000 mm except Sueve and Quinto Real which receive 1000 to 2000 mm annually. The mean Jancary temperature at DoAana is around 10°C and in summer about 30°C. The valley of the Guadalquiver has been described as the hottest place in Europe in summer (Branigan, 1974), maintaining a high temperature for several months. The other Fallow deer areas are mostly subject to temperatures of around 0°C in mid-winter and 22°C in summer.

2.4.20. Sweden In the southern third of Sweden..wild Fallow deer occur in the provinces of Sklne, Halland, Smaand, Vistergotland, Dalsland, Ostergotland, Sodermanland, Narke, Vastmanland and Uppland (Siivonen, 1976). The species has also been observed or killed in Varmland, the province on the north side of Lake Vanern, and in Dalarnia (Kopparberg) which is north-west of Uppland (Fig. 12). The total Fallow population was estimated to be between 4500 and 5000 in 1971 (K. Curry- Lindahl, pers. comm., 1973) and about 1100 are shot each year (Anon., 1974b). The deer were introduced to Sweden in the sixteenth century: the exact date is now known but those sent from England in 1579 were not the first (R. Gerell, pers. comm.).

In SkIine, the most southerly province, Fallow are widely distributed (Gerell, 1977). In the south of the province they occur in a triangle approximately within the points of Svelala, Sjobo and Lake Vomb with the densest populations on the Hackberga and Ovedskloster estates. Red deer also occur within the same area. Ahlen observed that a very high density of Fallow seemed to exclude a high density of Roe deer but did not influence the number of Red deer (I. Ahlin, pers. comm., 1979). The habitats are mostly forests in the proximity of fields with crops such as rye (Secale cereale), potatoes (Solanum tuberosum), rape, carrots (Daucus carota), corn, beets (Beta spp.) and legumes. The forests are varied and include Scots pine plantations on former heathland where the ground vegetation may include Deschampsia flexuosa, Carex arenaria and Rubus idaeus.


There are also Norway spurce plantations, mixed spruce-beech woods and broad-leaved, deciduous woods. The latter are mostly beech or beech with ash, hazel, hawthorn, birch and oaks (Quercus robur and Q. borealis) (AhlBn, 1965). In similar habitats, Fallow occur north-west from Lake Vomb to Angelholm on the west coast with the greatest number around Wrams-Gunnarstorp. Further populations are found east of Lake Ring, which is in the centre of the province, particularly at Maltesholm between Horby and the east coast (R. Gerell, pers. comm., 1979).

In the other provinces there are many small populations of Fallow. The main areas where they occur in Vastergotland are around Lake Mjorn which lies northeast of Goteberg, on the south side of Lake Vanem, particularly around Gossater, and on Luro, an island, in that vast lake. Ostergotland has two main centres of populations of Fallow, at the northeast end of Lake Ascunden and on the north side of Lake Roxen, which lies between Lake Vattern and Norrkopping. Narke, the small central province within the Central Trough of lakes and plains, has Fallow deer mainly between the north end of Lake Vattern and Lake Hjdmaren. On the south side of the eastern end of that Lake more Fallow occur in the province of Sodermanland and near Stockholm there are free-living Fallow on the small island of Lidingo. Of the four main areas for Fallow in Uppland, three lie immediately north of Lake Malaren around Vasteras, south of Sigtuna and near Vaxholm. A fourth population is centred north of Uppsala at about 60"N (Sartorius, 1970). Claims have been made for this population to be the most northerly in the world (Anon., 1974b) but Hyvinkia in Finland at 60'38' is slightly further north. Parts of Varmland and Dalarnia lie further north, extending to 62"20', but the Fallow deer localities in these provinces are not defined and the species is reported there only occasionally.

Oland is situated in the Baltic Sea, a few kilometers from the southern end of the east coast of the mainland. The island is about 140 km long and 10 km wide but Fallow deer now occur only at the southern end in fenced birch-oak woodland within the 1000 ha Ottenby Nature Reserve (Plate 6). The herd is maintained at about 150 deer by culling sixty to seventy animals each year (R. von Schultz, pers. comm., 1976).

Most of the districts occupied by Fallow deer are less than 200 m above sea level. The annual precipitation averages between 500 and 800 mm. The duration and depth of snow cover varies. In Uppland snow usually persists from mid-December until the end of March. Further north it is increasingly severe but in Sklne and the coast of Halland it is only intermittent. The coldest month is February when the coolest of the Fallow deer areas experiences mean temperatures just below freezing point (e.g. -3°C near Stockholm) and in July the same areas average about 17'C.

2.4.21. United Kingdom The four countries which constitute the United Kingdom all have wild Fallow deer but only in England are they dispersed throughout almost the whole country. In Wales, and more especially in Scotland where they occur in only seven of the thirty-three counties, the populations are more widely scattered (Fig. 23). In Northern Ireland Fallow occur in several localities in the eastern half of the country (Fig. 21).

Fallow deer were present in England in the Second (Hoxnian) Inter-glacial and Last (Ipswichian) Inter-glacial periods but apparently became extinct during the Last (Wiirm) Glaciation. Their re-introduction in historical times has been attributed to various peoples but was most probably by the Normans in the eleventh century (Chapman & Chapman, 1975).

The hunting of Fallow deer became very popular and many of the nobility enclosed large tracts of woodland and wasteland in which to pursue this sport. In Essex alone, there are documentary references to more than 130 parks in existence at some time between 1200 and 1390 (L. M. Cantor, pers. comm., 1976). Although Saxton's maps indicate about 700 parks in England in the sixteenth century, their number had begun to decrease before the middle of that century. As the decline continued, the concept of a deer park changed during the eighteenth century from an expanse of untamed countryside in which to hunt t o the landscaped parkland surrounding and enhancing the owner's mansion. About ninety-four such parks remain in England now, with smaller numbers in Scotland, Wales and Northern Ireland.


‘Y - 180 km

Fig. 23. The distribution of Fallow deer in England, Scotland and Wales

Although Fallow deer remain in a few forests where the species has been wild for centuries, to a very large extent the present distribution of Fallow deer in the United Kingdom is associated with the former presence of deer parks. Although disbandment. of parks and escapes by deer have happened for centuries, it is the parks that were disbanded during th is century that can be most clearly correlated with the present distribution of wild deer. For example, in the county of Essex several of the main centres of Fallow populations are in the vicinity of one of the seven parks disbanded since 1914 (Chapman, 1977). Further examples can be seen in almost every county and many are cited by Whitehead (1964) whose county-bycounty description of the history and distribution of deer in Great Britain and Ireland is still the most detailed information available. Since Whitehead’s account, only brief summaries of the distribution of Fallow throughout the United Kingdom have been published (Chapman & Chapman, 1975; Taylor Page, 1971). The Biological Records centre of the Nature Conservancy Council holds the records submitted during the National Deer Survey and Mammal Distribution Survey, both begun in the 1960s. Provisional, incomplete maps have been published showing the I0 km square for which records were received between 1967 and 1972 (Arnold, 1978). Despite these surveys, it is likely that Fallow are even more widespread than the records suggest.

2.4.21.1. England Fallow are the mostly widely distributed of the six species of deer feral in England. They are regarded as common animals in many counties and have been recorded from almost all of the present forty-six counties (Fig. 23). The five exceptions are the metropolitan counties of Greater Manchester and of Merseyside, the small and industrialized county of Tyne and Wear, Cleveland, and the Isle of Wight, an offshore island which seems not to have had Fallow, other than in a zoological collection, for more than 160 years (Whitehead, 1964). Fallow even occur within the bounds of Greater London. Although most of th is county is urbanized, on its fringes there are a few semi-rural areas such as between Harold Hill and the Essex boundary. Here, 28 km from the

Distribution of fullow deer 103

centre of London, Fallow are regularly present in the fields and woods west of Weald Brook (Chapman, 1977).

All the counties which border the south coast have Fallow. In the west of Kent they are well established, from Eynsford and Sevenoaks eastwards to Chatham and the Maidstone area. Fallow are found in many localities right across East Sussex from the Northam-Robertsbridge-Battle area, through the much-wooded countryside to, and beyond, Ashdown Forest (Conisbee, 1977). In West Sussex too, Fallow are very widespread through the many areas of mixed and deciduous woodland from St Leonard’s Forest, near Horsham, to the South Downs. On these chalk downs in the east of the county they occur around Steyning, Clapham and Arundel. The western and more heavily wooded half of the Downs has many Fallow, particularly on the south side, in the vicinity of Charlton, West Dean and Uppark. Nearby the ancient yew ( T a u s buccutu) forest in Kingley Vale near Chichester provides cover although little food, but the resident Fallow deer feed on the ash and bramble (Rubus fmcticosus agg.) at the woodland edge and in the adjacent fields (Williamson, 1978). North of the Downs Fallow also live near Midhurst and across the Rother Valley towards the borders with Surrey and Hampshire (Came, 1970a).

Hampshire, too, has much woodland and Fallow occur over most of the county. Their localities include the mixed and coniferous plantations of Bramshill Forest in the northeast, Petersfield, Emsworth and the Forest of Bere with its oaks, beech and conifers in the southeast. Near Andover, Harewood Forest has Fallow as does the Winchester area, particularly around Farley Mount and Hursley, and west of the River Test and over the county boundary towards Salisbury. The largest population of Fallow in Hampshire is in the 27 000 ha of the New Forest. One third of this Forest consists of deciduous and coniferous plantations: the rest is composed of natural deciduous woodland, heather moor, wet heath, grasslands and agricultural land and is the largest area of semi-natural vegetation remaining in lowland Britain. Fallow are dispersed throughout the Forest and probably number more than 1000. Smaller numbers of Roe, Sika, Reeves’ muntjac (Muntiucus reevesi) and Red deer are also present (Jackson, 1977). Much of the 3600 ha of unenclosed woodland of the New Forest is largely of beech and oak (Quercus robur & Q. petraeu) with holly (llex uquifolium) as an understorey but many other species are also present (Tubbs, 1968).

Dorset has Fallow deer where it borders the New Forest, in the southeast between Lulworth Cove and Wool, in the Maiden Newton-Bridport area and in Cranborne Chase (Came, 1967a). The 2000 ha of the Chase form a forest of mixed hardwoods and conifers on chalk downland south of Shaftesbury and support a large population of Roe deer as well as Fallow deer (Prior, 1968).

In Devonshire scattered herds of Fallow occur between the Dorset border and the Otter valley (Came, 1967b). West of the River Exe the forested Haldon Hills and surrounding agricultural land are a stronghold for Fallow and some occur in the Whitestone Woods west of Exeter (Carter, 1967). On the eastern edge of Dartmoor, deer which originated from Whiddon Park, live in the heavily wooded Teign Valley, especially between Dunsford and Fingle Bridge (Scott, 1967). They occur within plantations of Douglas fir (Pseudotsugu tuxifoliu), Norway spruce (Piceu ubies) and Japanese larch (Lurix leptolepis), as well as in deciduous woods with birch (Betulu pendulu, B. pubescens), rowan (Sorbus aucupuriu), oak, hazel (Corylus uvellunu), beech and holly. All of these types of woodland are attractive feeding areas as they have sufficient understorey or ground cover, with bramble, various grasses and thyme (Thymus drucei) as the most common species. Clearings covered by bracken are used as resting places (Dempster, 1967). In the south of the county, Plym Forest, northeast of Plymouth, provides a suitable habitat for the Fallow which originated from Blatchford Park, a few kilometers away (Penistan, 1967). Just across the Devonshire border and into Cornwall, Fallow occur on Torpoint near the former deer park at Mount Edgecumbe. Further west, some are present south of Bodmin Moor, near Lostwithiel and near Falrnouth and also in the vicinity of Truro. Park escapees are sometimes seen in the Launceston area too.

Of the other southern counties of England, Somerset has Fallow in the north-west, in and around the coniferous plantations on Croydon Hill which lies south of Dunster, and around Birds Hill at Combe Sydenham. Formerly in this area there were three parks containing Fallow but these were disbanded during the 1940s and 1950s (Jackson & Moore, 1976). Within the county of Avon,

104

recently wild Fallow were reported for the first time in the vicinities of two parks, Berkeley near the Severn estuary and Dyrham in the east of the county (Symes, 1976). In Wiltshire, Fallow are found east of Salisbury and in the oak, beech and mixed woodlands of Savernake Forest and the neighbouring districts near Marlborough. Fallow are scattered through much of Berkshire from Maidenhead in the east, around Reading and northwards through the Vale of the White Horse up to the boundary with Oxfordshire. In Surrey they occur in the Farnham area.

In Oxfordshire Fallow are widespread. They are present in the Woodstock-Charlbury area, in Bernwood Forest east of Oxford, and on the border with Buckinghamshire where the Chiltern Hills (Plate 7) around Stonor and east of Aston Rowant and Chinnor are thickly wooded with conifers and beech. Some Fallow occur in Buckinghamshire where this county adjoins Bedford- shire and Hertfordshire. In this region many Fallow occur on the Ashridge Estate which lies mostly in Hertfordshire but extends into Bedfordshire. This National Trust property consists of 800 ha of arable farmland plus 1600 ha of which 75% is woodland and the rest is commons and thorn scrub. The woodland is mostly deciduous with beech, ash, oak, sweet chestnut and Wild cherry (Prunus aviurn) as the main species. The commons support bracken, gorse (Uex sp.) and birch. The history of the deer on the Estate has been discussed by Kingsbury (1 958) and Wilson (1964). Other parts of Hertfordshire which have Fallow are the Watford area, including Whippendell Woods near the edge of the town; Cuffley and Northaw which are close to the Greater London boundary; Hitchin and along the boundary with Essex, especially between Nuthampstead and Bishop’s Stortford (Clark, 1974).

A survey of deer in Essex, conducted between 1963 and 1971, showed Fallow to be confined to the north and west of the county with the exception of a small area to the north-west of Clacton-onSea. Rarely were Fallow reported south of the London-Colchester road. They occur in many districts but the eight main areas were centred on Epping Forest, Navestockside, Mill Green, Roxwell, Hatfield Forest, Strethall, Markshall and Clacton-onSea. Included in the Epping Forest area are the woods and fields on the southern edge of Harlow New Town in which Fallow survive despite much hurnan pressure (Chapman, 1977).

Within East Anglia, Fallow are widely distributed (Chapman & Chapman, 1973). In west Suffolk they are found in the King’s Forest, which is mainly pine plantations but with some deciduous species including oak, beech and sweet chestnut, and is bordered by agricultural land. To the southeast of this Forest Fallow occur in many deciduous woods in parishes near Bury St Edmunds including Saxham, Whepstead, Lawshall, Long Melford, Hawkedon, Cavendish and, extending westwards towards the Cambridgeshire border, Dalham, Ousden and Lidgate (Cranbrook & Payn, 1970a). East of Bury St Edmunds they are found across the southern half of the county, through Felsham, Chelsworth, Ashbocking, Helmingham, south of Ipswich and between the Rivers Orwell and Deben to Kendlesham Forest and Tunstall Forest. These Forests, which together cover about 2400 ha, are mainly coniferous and lie between the coast and heathland (Edlin, 1972). Near Southwold, Fallow are found around Blyford and Henham (Cranbrook & Payn, 1970b). In Norfolk, they occur mainly in the King’s Lynn, Sandringham and Swanton Novers area. They are also present in some of the many, scattered mixed and coniferous woods between Cromer and Norwich, particularly north of Horsford (Goldsmith, 1972).

Much of Cambridgeshire is covered by tree-less expanses of flat, intensivelycultivated fenland but on the periphery there are some places which provide suitable habitats for deer. One such area is in the south-west corner of the county, around Gamlingay, where there is a cluster of small deciduous woods including Hayley Wood, the ecology of which has been described by Rackham (1975). Fallow deer move amongst this and the neighbouring woods which are surrounded by arable land (Symonds, 1979). In the north-west corner of Cambridgeshire Fallow are well established in the region formerly called the Soke of Peterborough, especially in the triangle formed by

’ Peterborough, Stamford and Wansford. This is an area of agricultural land in which are dispersed small deciduous or mixed woods and includes Castor Hanglands, a National Nature Reserve covering 90 ha of oak and ash woodland, grassland and scrub adjacent to plantations of spruce, oak and beech. Up to forty-five Fallow have been reported within the Reserve but the number fluctuates as


Distribution of fellow deer 105

the deer move within a greater area (Collier, 1970). Recently there has been an increase in the distribution of Fallow deer in Lincolnshire. In the south-west corner of the county they have been established for some years in the Stamford-Bourne-Kirkby Underwood-Grantham area. Now there are reports from the centre of the county, gn the western side of the Wolds, from Linwood and Willingham Forests and around Woodhall Spa and Wragby (Johnson, 1976). In the adjoining county of Humberside, in 1977 Fallow were liberated on Read’s Island which lies in the River Humber, west of Barton-upon-Humber. Most of the 100 ha of this flat and exposed island are covered by grass, especially Cock’s foot (Dactylis glomerafa) but a marshy area supports tall reeds (R. J. Nickerson, pers. comm., 1977).

Within the midland counties there are several areas where Fallow have lived for centuries although, as elsewhere, the nature and extent of the forests may have changed. In Northampton- shire they occur between Corby and Oundle, in the broad-leaved and coniferous woods which comprise Rockingham Forest, and also in Whittlewood Forest, which extends over the southern border of the county into Buckinghamshire. In Nottinghamshire, some are present in Sherwood Forest and on both sides of the M-1 motorway in the Underwood-Annesley Woodhouse area (D. R. Warren, pers. comm., 1977). Cannock Chase in Staffordshire, situated between Birmingham and Stoke-on-Trent, also holds Fallow. The Chase comprises 2400 ha of plantations, mostly coniferous but some hardwood, and expanses of Calluna heath and scrub (Burton, 1973). Elsewhere in Staffordshire Fallow occur in the Abbots Bromley area, in Needwood Forest and near Cheadle. In Leicestershire, Fallow live in the deciduous woods which constitute Charnwood Forest in the north of the county. Fallow also occur in the vicinities of Spondon and Matlock, both in Derby- shire (C. B. Bourchier, pers. comm., 1976; R. Jamie, pers. comm., 1976). In Warwickshire Fallow live around Edge Hill to the southeast of Stratford-upon-Avon. The adjacent county of Hereford and Worcester has Fallow, which escaped from the park at Elmley Castle, living in the hawthorn scrub, meadows and woods on Bredon Hill which rises to 320 m. They also occur near to both Hereford and Leominster and around Ross-on-Wye, which is close to the boundary of Gloucester- shire where the 13 000 ha of the Forest of Dean is the main centre for Fallow deer. Although conifer plantations are replacing much of the ancient woodland where oak predominated, there are extensive areas of mixed hardwoods within this Forest. One such place favoured by the deer is Highmeadow Woods, west of Coleford. The second main area for the deer is near Speech House, west of Cinderford (Venner, 1970). The deer of this ancient Forest, from early times until the present, have been discussed by Hart (1971). In Wyre Forest, which straddles the boundary between the county of Hereford and Worcester and the county of Salop, Fallow are well established, particularly in the areas of oak and beech (Hickin, 1971). Also near the boundary between the same two counties is Mortimer Forest, south-west of Ludlow, where many Fallow live, including some of the unique long-haired variety (Springthorpe, 1969). The Forest includes areas of deciduous woodland but most of the 3600 ha, which extend to an altitude of about 300 m, are covered by conifers.

In the county of Merseyside it seems that the Fallow formerly in the Ince Blundell district, south of Formby and near the northern suburbs of Liverpool, were exterminated by about 1975 (N. Dewhurst, pers. comm., 1978).

The northern counties of England have fewer populations of Fallow than the southern or midland counties but herds are scattered as far north as the North Yorkshire-Co. Durham border and Cumbria. In South Yorkshire a few Fallow occur to the north-west of Barnsley (Robinson, 1975a). In North Yorkshire small numbers of Fallow, as well as Roe and Red deer, occur in Stang Forest which lies at about 500 m above sea level and is a few kilometers south of Barnard Castle. The Swale Valley, about 8 km west of hchmond, also has a few Fallow. Further south, they have been reported near Swinton Park in the Masham and West Tanfield districts. A few Fallow are found in the Laver valley, not far from their place of origin, Studley Royal Park. A greater number occur north-west of Harrogate in the woods of mid-Nidderdale. East of Harrogate they are present in the vicinity of the former Allerton deer park near Knaresborough (Robinson, 1975b). West Yorkshire has some Fallow southeast of Wakefield, originating from Nostell Priory where


there was formerly an enclosed herd (G. M. V. Winn, pers. comm., 1975). The Fallow in Cumbria are in the south of the Lake District, near the boundary with Lancashire, where they occur in the dense ash and other broad-leaved woods on the limestone hills near Arnside (Delap, 1970). In Northumberland Fallow are present near Alnyick. A herd of about sixty live mostly within a former deer park. Although it is only partly walled there is little evidence of the deer moving out of, or into, the park (W. F. P. Hugonin, pers. comm., 1975).

2.4.21.2. Northern Ireland Fallow deer are present in four of the six counties in Northern Ireland: Tyrone, Co. Armagh, Down and Antrim (Fig. 21) (Crichton, c. 1974). The largest number are found where Co. Armagh and Tyrone meet each other and Co. Monaghan, which is in the Republic of Ireland. The main districts are around Caledon and Claslough, near the River Blackwater. In Down, Fallow are established in the vicinity of former deer parks on the Clandeboye Estate between Bangor and the northern end of Strangford Lough. A more extensive area for Fallow lies on the northern side of Lough Neagh near Randalstown in Antrim. The herds there originate from the former park at Shane’s Castle. A new 9 ha park, holding about twenty-five Fallow, has been established within Randalstown Forest primarily for educational purposes (C . S . Kilpatrick, pers. comm., 1976). No reports of Fallow deer were received in recent years from Co. Londonderry nor from Fermanagh (Crichton, c. 1974), although when Whitehead (1964) discussed the history and distribution of deer in Ireland, a small number of Fallow were present near Colebrooke in Fermanagh.

2.4.21.3. Scotland Despite the abundance of native Roe and Red deer, Fallow have been introduced to Scotland on a number of occasions as both wild and park animals. Stirling Park was in existence at least by 1283 and others were established later (Whitehead, 1964), but only five parks remain at present. The free-ranging Fallow populations are confined to the lowland parts of the country or valleys on the periphery of the Highlands (Fig. 23).

In southern Scotland, Fallow are established in two counties, Kirkcudbrightshire and Dumfries- shire, and have a tenuous hold in one area of Wigtownshire where an attempt at re-introduction was begun in 1971 but only six deer were believed to be present early in 1977 (‘Gallowegian’, 1977). The Kirkcudbrightshire Fallow are near Newton Stewart which lies at the southern end of the 6200 ha of the Galloway Forest which has some mixed broad-leaved and Sessile oak woodland amongst the extensive conifer plantations. The Dumfriesshire wood around Raehills, near Moffat, have held Fallow for two centuries: more recently some of the deer moved into the nearby Forest of Ae where spruce predominates, but willow (Salix sp.) and alder grow in the wetter situations. In 1975 the number in the Raehills area was believed t o be about 200 (P. A. Hope Johnstone, pers. comm., 1975). In the 1960s wild Fallow were also present in Roxburghshire, having escaped from Ancrum Park, but there have been no recent reports (D. Bridges, pers. comm., 1974).

Further north, Fallow are established in both Stirlingshire and Dunbartonshire in the vicinity of Loch Lomond (Idle & Mitchell, 1968). Within the Loch itself Fallow live on at least two islands, Inchlonaig and Inchcailloch. The deer were introduced to Inchlonaig. which is about 1 km long, in the seventeenth century when alder and bog plants predominated. Now bracken is the dominant plant. By 1873 the island held 150 Fallow but after this date hunting was discontinued. In 1967 the resident population was about thirty but there may be some movements from this to other islands and to the shores of the Loch. On Inchcailloch a small number, usually not more than ten, are present in the semi-natural deciduous woodland (A. Allison. pers. comm., 1974). East of Loch Lomond, lying across the borders of Stirlingshire and Perthshire. is the Queen Elizabeth Forest Park, which covers 17 000 ha and is composed of the three Forests of Rowardennan, Loch Ard and Achray. Roe and Red deer are numerous there but a handful of Fallow were also present early in the 1970s. The main area for Fallow in Perthshire is on the east side of the Tay valley between Dunkeld and Ballinluig. Smaller numbers also live between Dunkeld and Blairgowrie (A. Allison,


pers. comm., 1974). The Dunkeld populations are probably the most northerly ones established in Scotland, because the white herd which were in the Berriedale district of Caithness some years ago (Carne, 1970b) are no longer free. The herd had escaped from Langwell Park into the adjacent woodland but when this was replanted in 1963 most of the herd was reemparked and the remainder shot (M. R. M. Leslie, pers. comm., 1975).

At various times Fallow have been introduced to, or reached by swimming, at least eleven of the islands off the west coast of Scotland. Now they survive on only three islands, namely Mull, Islay and Scarba, all of which are within the county of Argyllshire. On Mull they occur in the Loch Ba’ area. A census in April 1974 on Scarba, by far the smallest of the three islands, with an area of about 1200 ha, indicated a Fallow population of twenty-one animals while on Islay over fifty live on the Kildalton estate (P. A. Joynson, pers. comm., 1974).

2.4.21.4. Wales Fallow have been kept in parks in Wales for centuries and the populations now established in the wild owe their origins to escapes (Whitehead, 1964). Now there are five park herds and feral Fallow are reported from six of the eight counties (Fig. 23). They are absent only from the small and more urbanized counties of South Glamorgan and Mid Glamorgan although many are present close to the border of the latter, in Margam Forest in West Glamorgan. This extensive forest near Port Talbot is mostly of pines but does include larch plantations, and areas of old oak woodland and other broad-leaved woods.

The much larger county of Dyfed in south-west Wales has Fallow deer in the woodlands which comprise the Forests of Brechfa and Caeo on the hills between Llandeilo and Lampeter. Brechfa Forest has some oaks among its conifer plantations and Caeo has a high proportion of broad-leaved trees, especially oak, beech and sycamore. Also in Dyfed, Fallow occur south of Carmarthen where there are a number of small woods. In the county of Powys, they are found around Welshpool and the Long Mountain, close to the English border. Gwent also has Fallow along its boundary with England which follows the Wye valley. The valley is well-wooded with both native hardwoods and conifer plantations (Edlin, 1974) and provides a suitable habitat for Fallow that originated from the Forest of Dean or from parks such as Wyastone Leys where there was formerly an enclosed herd (C. Hart. pers. comm., 1974).

In Clwyd in North Wales, the Fallow deer localities are south of Wrexham and in the Denbigh- Abergele-Colwyn Bay area. In the adjacent county of Gwynedd, a mass escape of ninety deer from Nannau park near Dolgellau in 1963 resulted in the establishment of several feral populations (Vaughan & Carne, 1971). Some animals remained in the vicinity of the Park but others moved northwards and deer now flourish in the mainly coniferous Coed-y-Brenin Forest. Others went towards the Dovey Valley, finding suitable habitats in the valleys and on the lower mountain slopes where there are some oak woods as well as conifer plantations. In the north-west of Cwynedd, they occur near Bangor, close to the Menai Straits, having originated from animals kept in Yaynol Park. Across the Straits, on the island of Anglesey, a small herd lives in the pine plantations on the sand dunes of Newborough Warren and move freely on and off the Bodorgan estate from which they originated (M. J . Green, pers. comm., 1974).

For much of the United Kingdom the mean temperature in January lies between 2 and 6°C. In July most of England has mean temperatures around 16°C. Scotland, Northern Ireland and the northern counties of England are usually a few degrees cooler. Many of the Fallow deer localities in the Midlands and eastern counties of England receive a mean annual rainfall of less than 750 mm. In the south-western counties and the border between Wales and England the annual rainfall may reach 1OOOmm and a few of the deer areas, mainly in northeast England, Wales and Scotland, receive an annual mean precipitation of 1500 mm. Almost all of the Fallow deer localities lie within areas which experience lying snow on an average of less than 20 days a year. Very many of the places, in southern England, near the west coast of England and Scotland, South Wales and Northern Ireland usually have snow on less than 10 days per year.

108 N. G. Chapman and D. I, Chapman

2.4.22. U.S.S.R. Wild Fallow deer now occur in small numbers in a few localities in the western part of European Russia (Fig. 24). Although this species has never been widely distributed in this country it has been more numerous in the past.

As early as the sixteenth century some Fallow were present in the forests of Lithuania and Belo-Russia (Heptner, NasimoviE & Bannikov, 1966). At the beginning of the seventeenth century, large numbers were observed on the Ukrainian steppes (Vereshchagin, 1967) and early in the eighteenth century some were living in the Bialowieza Forest, then in Russia and now on the

- 1000 km

Fig. 24. The distribution of Fallow deer in the U.S.S.R.

Polish-Russian border, which was the Czar’s hunting reserve. In 1857, this forest, where hornbeam (Capinus sp.), oak (Quercus spp.) and ash (Fruxinus s ~ . ) predominate, held 7000 Fallow, 7000 Red Deer, 5000 Roe deer and 1900 European Bison (Bison bonasus) (Curry-lindahl. 1964). Further introductions of Fallow were made, yet by 1920-1930 the species had become extinct (Heptner et al., 1966; Lindemann, 1956).

Lithuania was one of the Baltic areas in which Fallow deer were bred in parks during the last century. Animals from these parks were used to stock various forests in that Republic. In the 1950s. half of Russia’s total Fallow population, estimated at only 200, were in Lithuania and some are probably present there now. Park animals were also taken to the Gatchina game reserve south of Leningrad and the forests of Belo-Russia where in addition, Fallow from Germany were released in 1932 in the Voloshin lvenez Molodetscho area near Minsk (Heptner et al., 1966). but by 1974 none remained (V. G . Heptner, pers. comm.).

Similarly, the herd on the island of Khortiza in the River Dnieper near Zaporozhye in the Ukraine had ceased to exist by 1974 but. further south, the population on the Biryuchiy peninsula in the Azov sea numbered about 100. Also in the Ukraine, Fallow are found within the 867 ha of the Askania-Nova reserve on the broad plains between the lower courses of the rivers Dnieper and Molochnaya. However. these deer, bred here since 1918 to supply stock for other reserves, are kept in extensive enclosures on the steppe pastures (Treus, 1970). To the north-east, still just in the Ukraine, and on the southern edge of the Central Russian Uplands, are the extensive Donbass Forests. These lie north-west of Khakov, adjoining the vast forest which bears that name, and include the forests of Tepensky, Majaksky and Nickolsky. These comprise 20 000 ha of mixed forest in which deciduous species predominate. About 20 years ago Fallow were observed there for the first time. Their continuing presence was encouraged by the provision of supplementary winter feed (Zubets. 1959) but their subsequent fate is unknown. The introduction of Fallow to the broad-leaf forests of the C a r p a t b n foothills in the Ukraine wassuggested by Tatarinov (1966)but it is not known whether this occurred.

Distribution of fall0 w deer 109

Close to the border with Romania, in Russian Moldavia, Fallow live in the woods near Kishinev. The herd numbered about forty in 1974 (V. G. Heptner, pers. comm.) and may have resulted from a liberation in 1961. -62 (Treus & Lobanov, 1966).

In 1888, fifty-four European Fallow were liberated on the mountain slopes of the Borhomi game reserve in the Georgian Republic, 150 km west of Tbilissi, in the southern foothills of the Caucasus mountains. More were released in the following 2 years and by 191 8 several hundred wandered within the 500 ha enclosure but during the next 2 years all the herds were exterminated by the local people (Vereshchagin, 1967). Later attempts to acclimatize Fallow in the Caucasus have failed (Aliev, 1970) although the shrub-forest zone of the Stavropol plateau, the Dagestan piedmont, eastern Armenia, Nakhichevan A.S.S.R. and the Nagorno-Karabkh uplands were all considered suitable areas for their release by Vereshchagin (1 967).

The parts of the U.S.S.R. in which Fallow occur fall within the zones of deciduous forest, wooded steppe and grassland steppe all, apart from Lithuania in the central west, in the south-west corner of the country. Most of the localities are lower than 200 m and even the upland areas are not more than 500 m above sea level. These areas are subject to a moderately continental climate. The mean temperature in January is a few degrees below freezing point, the coldest of the Fallow deer areas probably being about -7"C, near Kharkov. In July the mean temperature is around 20°C. The annual rainfall is between 450 and 700 mm. Snow is not severe in any of the areas and on the coast of the Sea of Azov, in which the Biryuchiy herd of Fallow live, the maximum depth of snow is under 100 mm.

2.4.23. Yugoslavia Most of the localities where Fallow deer occur in Yugoslavia lie in a southeasterly band across the northern half of the country (Fig. 25). The successful trial importation of this species to hunting grounds in Montenegro, in the central western part, suggests that conditions are favourable so their distribution may be extended by further introductions (VuCkovid, 1970). Whereas Red deer and Roe deer are numerous, with estimated populations of 18 000 and 350 000 respectively in 1970, Fallow are few (IsakoviC, 1970).

Within Slovenia, in the north-west of the country, about 80 km north-west of Zagreb and only about 25 km from the Austrian border, lie the Pohorje hills, the most westerly outcrop of the

Fig. 25. The distribution of Fallow deer in Yugoslavia - 90 krn

1 10 N. G. Chapman and D. I. Chapman

Alps. On the partially wooded, gentle slopes between the towns of Kozina and Sezane a free- ranging population of Fallow are established. Another wild population lives in the Velenje and Posavje hunting reserves on both banks of the River Sava between Litija and Trbovje, east of Ljubljana (B. Andrej, pers. comm., 1974).

The Republic of Croatia has Fallow in the Slavonia region, much of which is relatively flat. Southeast from Zagreb, in the vicinity of Bosanski Novi and Dubica near the River Una, a further population of about thirty exists. North-west of Belgrade Fallow are established in the Fruska Cora hills. This chain of hills, mostly under 500 m high, lies between the Sava and Danube Rivers, between Sremska Mitrovica and Novi Sad and extends westwards to Kunjecvia in the Vinkovci district.

About 20 km south-west of Belgrade, in the Republic of Serbia, a population of about seventy Fallow live near Lipovica, having been introduced in 1958. About 160 km to the southeast of the capital, at the Majdan -Kucajna reserve, is a small enclosed herd which originated from ten animals introduced in 1965. A similar number was also released in the same year into the reserve at Boranja, near the River Drina, south of Loznica. Between these two enclosed reserves and north of Gornji Milanovac, is a third enclosed herd at Rudnik where the animals introduced in 1961 increased to seventy by 1965 (Bojovik, 1967).

Although three-quarters of Yugoslavia is mountainous, almost all of the Fallow habitats are less than 500 m above sea level, and most are below 200 m, and lie within the forested flood plains of the major rivers. Much of the lowland woodland is deciduous, with beech and oak as the dominant species.

There are also two island sites for Fallow in Yugoslavia. Cres is narrow but about 72 km long and lies in the Kvarner Gulf of the Adriatic Sea. Brioni, only about 6 km long, lies off Pula where the Gulf of Venice meets the main body of the Adriatic Sea. The population of Fallow on Brioni is around 500 and animals from this island have been used to stock most of the other localities where the species occurs in Yugoslavia (B. Andrej & I. IsakoviC, pers. comm., 1974).

Meteorological records from the neighbouring mainland resort of Opatija show the lowest and highest mean monthly temperatures ever recorded to be 9 and 26OC, respectively, although Cres itself is more subject to cooling breezes. Snow is unknown in this area. The annual rainfall for most of the Fallow deer localities is between 600 and 800 mm, but in the Ljubljana district it reaches 1400 mm. The mean temperatures in July are between 20°C and 24OC and in January they range from - -2°C to 8°C.

2.5. NORTH AMERICA Despite the abundance of native deer in North America, Fallow have been introduced and now occur wild in one province of Canada and in many widely spread areas within the United States of America (Fig. 26).

2.5.1. Canada I

British Columbia has the distinction of being the only part of Canada to which Fallow have been introduced (Fig. 26). According to Carl & Guiguet (1958) the deer were released on James Island about 1895 and subsequently, during the 1930s, some were transferred to Saltspring and Pender Islands and to the Alberni District of Vancouver Island. However, J. H. Todd (pers. comm., 1974) believes that Fallow deer were introduced to James Island in 1907, having been imported from the Chatsworth estate in Derbyshire, England, so this may have been the second introduction to the island. About 1954 some Fallow swam the 2 km from James Island to Sidney Island and became established there.

The deer are now present only on Saltspring Island, James Island and Sidney Island, not having been reported on Pender Island since about 1930 nor on Vancouver Island since 1952 (G. W. Smith, pers. comm., 1974). These islands are situated at the southern end of the Strait of Georgia, within a few kilometers of Vancouver Island. In this Gulf Island Zone the winters are

Distribution of fallow deer 11 1

James Island

- 400 km

Fig. 26. The distribution of Fallow deer in North America.

mild, with an average December temperature of 4°C: summers are moderate with an average July temperature of about 17°C and precipitation is 760-1000 mm per annum, with 300-700 mm of snow in winter.

Saltspring Island is the largest with 18,000 ha of hills, forests, lakes and mountains, but has only ten to twenty Fallow deer (G. W. Smith, pers. comm., 1974). On James Island, the smallest with an area of 280 ha, the Fallow number about 125. A large population lives on the 713 ha of Sidney Island where the habitat comprises woodland, open fields and rocky knolls. The trees include Douglas fir, Red alder (Alnus nrbru), Broadleaf maple (Acer mucrophyllum), Carry oak (Quercus gurtyuna) and madrone (Arbutus menziesii). Ground vegetation includes salal (Guultheriu sp.) , ferns, Stinging nettles (Urticu dioicu), grasses and mosses. In November 1977, the Fallow population was believed to number several hundred. The deer have flourished on this island since freshwater ponds were dug: before then Fallow which swam to the Island did not survive. The island also supports various other introduced species of birds and mammals, including Mule deer (Udocoileus hemionus), (J. H . Todd, pers. comm.).

2.5.2. United States of America Fallow deer are the most widely introduced of the exotic ungulates in America (Presnall, 1958). Although they did not become established in some areas, wild Fallow are now present in at least eight States, from the east coast to the west coast and on some off-shore islands (Fig. 26).

The main period for the release of exotic game species began later in the United States then in Australasia, most introductions being made during this century. The earliest known introduction of Fallow deer to the U.S.A. was from England in 1878 (Wehausen, 1973). Since then the species has been kept in many zoological collections and parks, and stock released into the wild have usually come from such sources rather than being imported direct from Europe. Although a controversial subject amongst biologists and conservationists, further introductions are still being considered in some States to fill unstocked or under-stocked areas with game species. Consider-

1 12

ation has been given recently to the release of Fallow in southern Indiana but the authorities fear there may be too much conflict with agricultural interests. In the same State, Fallow released at Bluffton were shot after straying into the suburbs (Johnson, 1972; pers. comm., 1973). A recent attempt to establish them in Louisiana for archery hunting failed because the animals were poached (J. W. Farrar, pers. comm., 1974).

N. C. Chapman and D. 1. Chapman

2.5.2.1. Alabama In 1925, twelve Fallow, from an enclosed herd at Prattville, were released in the Black Warrior National Park in Wilson and Laurence Counties but did not survive (Allen, 1965). About the same time an enclosed herd was established at Miller's Ferry, Wilcox Co. and in the early 1930s some of the deer escaped. During the next 30 or so years the population reached an estimated 200-300 and spread up to 32 km from the point of escape (Allen, 1965), a rate of spread of about 1 km per year. Since then there appears to have been little increase in their distribution in Wilcox Co. and Dallas Co, (Fig. 26). The deer h a b i t the alluvial river plains along the Alabama River, known as the Black Belt, which is an area of dark lime or basic soil once covered by salt water. The land is used for pasture, forestry and crops such as Soy beans (Clycine sp.), cotton (Gossypium sp.) and cereals (R. H. Allen, pers. comm., 1974). The main tree species in the wooded areas include Shmgle oak (Quercus imbricaria), Overcup oak (Q. lyrata), other Quercus species, Nutmeg hickory (Carya myristicaeformis) and Red Cedar (Junipems virginiana).

The ultimate fate of th r ty white Fallow which escaped in Mobile Co. in 1946 is uncertain: up to 1965 some were thought possibly to remain in the area. Captive Fallow which escaped at Chapman in Butler Co. failed to become established (Allen, 1965).

2.5.2.2. California Fallow deer occur in several areas in California (Fig. 26). The largest population is that at Point Reyes Peninsula, Marin County. The Peninsula protrudes into the Pacific Ocean about 50 km north-west of San Francisco and is bounded on the east by Inverness ridge of which the highest point is M t Wittenberg (460 m). A survey carried out in March 1977, on the ground and from a helicopter, concluded that the Fallow numbered over 500, ranging over 6400 ha. Mule deer, domestic cattle and about 400 Axis deer were also present (Elliot 111, 1977).

Although some common, black and menil colour varieties of Fallow are present, the white phase predominates: 75% of the stock introduced to Point Reyes from San Francisco zoo were white. The first fifteen deer, all white, were liberated in 1942, another eleven were released in 1947 and two more in 1954. No shooting of the deer took place until 1956 and since then hunting has been limited to adult males apart from some does and fawns taken for the purposes of study- ing parasites, health and food preferences (Brunetti, 1976).

The main habitats at Point Reyes are forest, scrub and grassland. The scrub was formerly extensive but much of it was cleared during the 1920s to 1950s to create more pasture for live- stock. This grassland is the main habitat of the Fallow deer. Much of it has not been grazed by cattle for several years and the vegetation is generally knee-high. The dominant grasses are Bromus rigidus, replaced in the drier places by Fesruca dertonensis, Bromus mollis and, in the moister situations, Holcus lanarus, but at least six other species of grass are also grazed by the Fallow deer. The deer do occasionally utilize the scrub which varies from dense to very open. Before and during the fawning period, does are seen in the scrub more often and the deer also seek shelter there when the wind is strong. In some parts of the range where there is less grassland the deer feed on the grasses and herbs in the open scrub. They browse some scrub species such as Scotch broom (Cyrisus scoparius), blackberry (Rubus virifolius) and Ceanorhus rhyrsifloms and also willow (Salk sp.) and Red alder whch grow along streams (Wehausen, 1973).

The average annual rainfall is 460 mm, most of which falls between November and April but ocean fog is frequent in summer and its condensation enables the growth of perennial grasses. The average temperature in January is 10°C, rising to only 12°C in July.

The fifty-one white Fallow introduced on the Ridgewood Ranch south of Willets, in Mendocino


County in north-western California, in 1949 have become well established although remaining within 11 km of the point of introduction. Despite hunting and poaching, in 1970 at least 130 deer were estimated to be present within 7300 ha of grassland and oak woodland (Plate 8) center- ing on the Walker Valley, which lies at an altitude of 400 m and is a few kilometres north of Lake Mendicino. Highway-101, a four-lane road, bisects the area. The terrain is rolling foothills and ridges, some over 800 m above sea level, between the Laughlin Range and other mountains, interspersed with flat areas in the bottoms of valleys. The most abundant grasses are Bromus mollis, Slender wild oat (A venu burbufu) and Dogtail grass (Cynosunts echinutus). The commonest species of tree in the oak woodland and oak savanna are Carry oak and California black oak (Q. kelloggii), but there are some Valley oak (Q. lobutu), Interior live oak (Q. wislizenii) and Blue oak (Q. douglasii). Dense thickets of madrone and manzanita (Arctostuphylos munzunitu) are scattered within the woodland and grassland. No adverse effects have been observed on the native Mule deer which live in the same habitat. The Fallow's range is to some extent restricted by the surrounding coniferous forests where Douglas fir predominates; the chaparral, a shrub association reminiscent of the maquis of the Mediterranean where there is scant growth of grass and herbs; and the redwood (Sequoia sempervirens) forest. The annual rainfall is about 130 mm. Average snowfall is about 75 mm and covers the ground for only a few days. The m e a n o m i n i m ~ and maximum temperatures in July are 11" and 30°C respectively and in December 0 and 9 C, but the extremes of 41" and -14" have been recorded (Jurek, 1977).

Also in Mendocino County, within the past few years a small number of Fallow deer has been released from the Hopland Field Station of the University of California but their present status is unknown (H. W. Elliot 111, pers. comm., 1978).

The deer originally supplied to the Ridgewood Ranch came from a private zoo on the Rancho F'iedra Blanca (Hearst Ranch) at San Simeon in San Luis Obispo County. Fallow were kept there from the 1930s until 1953 when eighty-five escaped: in the mid-1960s it was believed that only one animal remained.

Near San Francisco Bay Fallow occur in two localities. At the south end of the Bay in Santa Clara County, there are some in the Mount Madonna County Park. The herd originated from two pairs of Fallow released in 1934 but only about twenty were present in 1976 as culling takes place annually (H. W. Elliot 111, pers. comm.). The annual rainfall is around 500 mm: the mean temperature is about 9°C in January and 16°C in July. In the adjacent county of San Mateo, on the southwest side of the Bay, another herd lives on the Pomponio Ranch. About fifty were present in 1965 but the size of the population now is unknown (H. W. Elliot 111, pers. comm.). The annual rainfall is about 700 mm. The mean temperature in December lies between 10 and 16°C and in July it is between 21 and 27°C.

Fallow may also be living wild in Tehama County. A captive herd of about thirty was maintained on a ranch 12 km south of Red Bluff but about 1976 vandalism to fences enabled the deer to escape and disperse along the Sacromento River between Tehama and the confluence of Thomes Creek. A few individuals are thought to have survived despite poaching. Rainfall is similar to that in San Mateo County but the average temperature in December is slightly lower (4-10°C) and in July it is a few degrees higher (27-30°C).

White Fallow were released in 1967 and 1970 at Lone Pine Ranch in south-west Trinity County, north of Mendocino County. By 1971, the population numbered between twenty and thirty. Less successful were the thirteen white Fallow liberated in 1960 on the Butts Ranch near Call Mountain in San Benito County. By 1970 only two females had escaped the attention of hunters. Similarly, by the mid-1960s Fallow had disappeared in Siskiyon County although a dozen had been released near Yreka a decade earlier (Jurek, 1977).

2.5.2.3. Colorado Presnall (1958) reported about seventy-five wild Fallow in Colorado, resulting from introductions in the 1920s to 1940s in Conejos, Larimer and Rio Blanco Counties. In 1973 the only recent

1 14 N. G. Chapman and D. I. Chapman

record known to the Colorado Division of Wildlife was of one Fallow shot near Fort Collins (A. E. Anderson, pers. comm.).

2.5.2.4. Georgia Early in the 1950s Fallow were introduced to Little St. Simons Island, which lies off-shore from Brunswick (Fig. 26). The deer flourish on this low-lying island with its mild climate (mean winter temperature 13OC: mean summer temperature 27°C) and annual rainfall of 1270 mm. By 1974 the population was estimated to be between 500 and 600 within an area of 4000 ha. The deer utilize all the habitats available, namely zones of Live oaks (Quercus virginiana), of pines (Pinus spp.), an intermediate zone of oaks and pines with some understorey of palmetto (Sabal sp.) and myrtle (Myricu sp.), and salt marsh. The deer feed mainly on the salt marsh, although acorns also form an important part of their diet. In contrast, Fallow released in 1957 on the mainland of Georgia, near Tocoa in the Lower Mountains, were so vulnerable to predation by dogs and poaching that they were exterminated within a year (T. Kile, pers. comm., 1974).

2.5.2.5. Kentucky Probably the largest population of wild Fallow deer in North America is in western Kentucky, between the Barkley and Kentucky Lakes, an area known as the Land Between the Lakes (Fig. 26). This peninsula of 68 850 ha is being developed for public recreation, including hunting, and environmental education. The Fallow population, estimated at about 700-800 and derived from twenty deer released in 19 18, has been studied by Terpening & Hawkins (1975). White-tailed deer were released at about the same time and are even more numerous.

The habitat includes rolling uplands with steep hillsides and ranges from 120 to 170 m above sea level. Woodland covers much of the ground (Plate 9). The main tree species include various oaks such as Shingle oak, Cherrybark oak (Quercus falcata var. pagoduefolia), white oak, (Q. a h ) , Scarlet oak (Q. coccinea), Post oak (Q. stelluta), Northern red oak (Q. rubra), Flowering dogwood (Cornus florida), Black walnut (Jugluns nigra) and hickory (Carya spp.). The meadows support a rich flora of fescues, including Festuca elutior, other cultivated grasses, legumes including white clover (Trifolium repens) and perennial herbs. The annual rainfall is about 1220 mm. The mean minimum and maximum temperatures for January are -2 and 8°C respectively and for July, 19 and 33°C.

Some Fallow were introduced to the Fort Knox and possibly also to the Fort Campbell Military Reservations a few years ago but it now seems that they are extinct (R. W. Nall, pers. comm., 1974). In Carter County a few Fallow have been released at the Grayson Lake Wildlife Area (J. S . Durrell, pers. comm., 1974).

2.5.2.6. Maryland Fallow were introduced between 1920 and 1945 to Worcester Co. and Talbot Co. in Maryland and a few were believed to have survived until the 1950s (Presnall, 1958). Mills Island in Chincoteague

Bay also formerly had a herd of Fallow (Flyger, 1959). By 1974 it appears that all three herds had become extinct (R. L. Miller, pers. comm.).

2.5.2.7. Massachusetts The island of Martha’s Vineyard, 32 km long and with a maximum width of 16 km, lies off the coast of Massachusetts 8 km from Cape Cod (Fig. 26). Two introductions of Fallow were made during the 1930s and small numbers were released in 1968 and 1969 (Codin, 1977; Keith, 1967 J. J. McDonough, pers. comm., 1973). The most recently introduced animals are thought to have been poached and the 1976 cull was the first for over 20 years to fail to report Fallow deer although a few deer of the black variety are believed to remain in the West Tisbury and Gay Head areas, at the western end of the island. See note on p. 138 added in proof.

Fallow were introduced to Nantucket Island, which lies east of Martha’s Vineyard (Godin,


1977; Uloth, 1971). However J. J. McDonough (pers. comm., 1977), the Game Biologist concerned with deer research projects, has not known of any Fallow on this Island during the past 28 years.

2,5.2,8. Nebraska The only population of wild Fallow in Nebraska owes its origin to the introduction, around 1939-40, of about seventy white deer from Pioneer Park, Lincoln which were released 16 km northwest of Petersburg in the Beaver River Valley in Boone Co. (Fig. 26). This broad valley has extensive hayfields and cornfields and, along the river bank, cover is provided by shrubs, particularly willows (Salix sp.) and cottonwood (Populus palmeri) (Packard, 1955). By 1946 Fallow were reported between places 3 km east and 25 km north-west of the point of release. Their range had increased but little by 1955, the population still being centred in Boone Co. and the adjacent Wheeler Co. Although there were unconfirmed reports from the neighbouring counties of Madison, Holt and Hall. An aerial survey in 1957, claimed to give a total count, recorded 157 Fallow: in subsequent years up to 1965 the counts varied from 105 to 157. The population, although maintaining itself, is said to have a low reproductive performance with an average, over a 9 year period, of only twenty-one fawns per hundred does. The authorities do not wish the numbers to increase and in 1966 a deliberate culling policy was begun: by 1973 the population had been reduced to 50-60 which was regarded as a suitable level to avoid conflict with agricultural interests and competition with the White-tailed and Mule deer (K. Menzel, pers. comm., 1973).

The mean minimum and maximum temperatures in January are -13 and 1"C, respectively and in July 16 and 32°C. The mean annual precipitation is 635 mm but is very variable from year to year and more than 100 mm may fall in one continuous period. The average annual snowfall is about 700 mm and severe storms and strong winds also occur in winter.

2.5.2.9. New Mexico A wild population of Fallow deer is established in the Sacramento Mountains in east central New Mexico (Fig. 26), having originated from animals which escaped from a park. A few are sighted or shot each year (W. A. Snyder, pers. comm., 1973) but no further details of the population are known.

2.5.2.10. Oklahoma The presence of Fallow deer in the Camp Gruber area of Muskogee County was mentioned by Presnall(l958) but J. Hammond Eve (pers. comm., 1976) reported that none is there now.

2.5.2.1 1. Tennessee Fallow are not known to be established in this State but a single animal has been reported on at least one occasion. A buck was shot in the Natchez Trace area, possibly having wandered from Kentucky or escaped from an enclosure in the Jackson Area (Anon., 1968).

2.5.2.12. Texas Texas has a large number and wide variety of exotic game including thirty-nine species of artio- dactyls. The most numerous are Axis deer, Mouflon-barbados sheep (Ovis sp.), blackbuck (Anfilope cervicapra), Fallow deer, Auodad sheep, Sika deer and nylgai (Armstrong, 1975). The first introduction of Fallow was during the 1930s on King Ranch, whtch extended from the Rio Grande in the west to the Gulf of Mexico in the east, within the South Texas Plains (J. L. Cooke, pers. comm., 1973). Although some 500 Fallow were said to have been introduced, by 1938 only twenty-two were thought to survive (Halloram & Howard, 1956), presumably as a result of excessive hunting. Fourteen of these deer were taken to other ranches after King Ranch was bought by the Government, but details of these translocations and subsequent introductions are not available.

In 1974, a State survey recorded a Fallow deer population of almost 4500. The majority were

1 16 N. C. Chapman and D. I. Chapman

Table 3 The zones o f Texas in which Fallow deer occur

1. 2. 3. 4. 5. 6. 7. 8. 9.

10.

Zone

Pineywoods Gulf Prairies and Marshes Post Oak Savannah Blackland Prairies Cross Timbers and Prairies South Texas Plains Edwards Plateau Rolling Plains High Plains Trans-Pecos, Mountains and Basins

Percent of Fallow deer population

0.4 0.6

15.3 0- 1

12.0 10.1 61.2

0.3 0 0

on ranches which had deer-proof fences, although only 142 out of 316 ranches with exotic game reported having such fences. Although only sixty of the Fallow counted were free-ranging, this species is known to be free in many counties, especially on the Edwards Plateau, but such herds were not included in the survey unless a reliable estimate was available. The total estimate was, therefore, a conservative one and the true figure would include more free-ranging Fallow. The report of this survey (Armstrong, 1975), and of the earlier one of 1971 (Young, 1972), does not distinguish between free-ranging Fallow and those on fenced ranches. Consequently, the map of the distribution of Fallow in Texas (Fig. 26) indicates the presence of both wild and enclosed deer. Although the wild Fallow formed a relatively small proportion of the total population, it is thought that if they continue to increase, they together with free-ranging Sika and Axis deer, could present considerable competition to the native White-tailed deer. The Fallow population has increased by more than 70% since 1971 (Armstrong, 1975).

Could (1969) divided Texas into ten zones based upon their vegetational features. In the 1974 survey, Fallow were recorded from seven of these zones (Table 3). By far the most Fallow occurred in the .Edwards Plateau in central Texas but only 2.2% were free-ranging. The proportion of free-ranging animals in the other zones was not given (Armstrong, 1975). The Edwards Plateau comprises 9 700 000 ha of Hill Country ranging from 30 m to over 900 m (Plate 10). The vegetation is grassland or open savanna. The grasses include Needle grass (Aristida sp.), bluestems (Andropogon spp.), gramas (Bouteloua spp.), Texas winter grass (Stipu leucotrichia), Buffalo grass (Buchloe ducryloides) and Curly mesquite (Hilaria belungeni). The rough, rocky areas support Live oak, Shinnery oak (Q. brevilobu), juniper (Juniperus ushei) and thickets of mesquite (Prosopis chilensis). The chief land use is the ranching of beef cattle, sheep and Angora goats with cultivation in the valley bottoms and near towns. The annual rainfall varies from less than 380 mm in the west to over 840 mm in the east: in Kerr, the county with most Fallow, the average rainfall is about 630 mm but severe droughts occur in some years. No Fallow were recorded in the 1974 census in either the High Plains or the Trans-Pecos, Mountains and Basins zones, both in the west of Texas, although a few animals were reported there in the 1971 survey, in the counties of Hudspeth and Carson (Young, 1972). Their absence in the later survey may mean that the herds had become free-ranging but could not be accurately assessed.

2.6. SOUTH AND CENTRAL AMERICA Despite the presence of eleven native species of deer in South America, Fallow have been introduced to Argentina, Chile, Peru and Uruguay (Fig. 27). In Central America, they are present on the Leeward Islands of Barbuda and Cuana in the Caribbean Sea (Fig. 1).

Distribution of fall0 w deer 1 17

Fig. 27. The distribution of Fallow deer in South America

2.6.1. Argentina Fallow deer are present in two widely separated parts of Argentina. One region lies within the Pampas Zone in the Province of Buenos Aires in the east of the country. The other, some 1000 km to the south-west, is in the Lakes Region in the Andean foothills (Fig. 27).

The Buenos Aires populations are found within a circular area bounded by Buenos Aires, Mar del Plata on the coast and Bahia Blanca with possibly an extention north-westwards into Santa FB Province. Although the first introduction of Fallow deer was to the Park Pereyra Iraolo early this century (J. M. Gallardo, pers. comm., 1974) most of the present populations owe their origin to animals imported by Ernest0 Tornquist in 1905. He obtained Fallow from Spain and Poland and enclosed them near hls home in the Sierra de la Ventana in the southern part of Buenos Aires Province. About 1930, his sons released the deer into the surrounding low mountainous country where they have prospered ever since. The deer now range over 300 000 ha in the region of Sierra de la Ventana, Cura-Malal, Saavedra and Tres Picos. A census in the first two of these places in 1972 concluded that approximately 5600 Fallow deer were present on forty estates within an area of 195 000 ha. About 3800 wild goats ( C a p hircus), 6000 guanacos (Lama guanicoe) and some feral horses (Equus caballus) were also present (S. A. Safontas, pers. comm., 1976). Within this area is a wildlife refuge where the fauna, including some 200 Fallow deer, is protected. For most of the year the Fallow remain in the rolling open plains and foothills at about 400 m, but sometimes they wander to higher ground which extends to over 1100 rn. A ground cover of grasses and herbs provides excellent pasture but there are almost no trees or shrubs (Plate 11). The annual rainfall in this usually windy region is about 860 mm. Temperatures in summer reach about 4OoC but in winter may drop to -9°C but there is little snow (E. E. Willard, pers. comm., 1977).

To the north and north-east of this region, Fallow deer are widely dispersed around Daireaux, Olavarria, Bolivar, Tapalquen, General AIvear, 25 de Mayo and further east to Tandil, Ayacucho,

1 18

Mar del Plata, Loberia and General Pueyrredon. The greatest number is around the Estancia Huetal, near the towns of 25 de Mayo and General Alvear. This estancia, 50 000 ha in extent when under the ownership of Mrs Concepcion Unzuk de Casares, is now divided into several properties. Fallow were introduced early this century and by 1950s the population was estimated at over 20 000. About 1958,4400 were shot, causing the remaining herds to scatter into the neighbouring agricultural land, used mainly for raising cattle. The present population in the area is estimated to be between 10 000 and 15 000. The region is very flat with no forests but the Fallow deer inhabit creeks and the grassy borders of lagoons (H. R. Rivadavia, pers. comm., 1977).

South of Sierra de la Ventana, some Fallow live in the Villarino district including the Verde Peninsula in Bahia Blanca Bay. Further south along the coast another wild population occurs on Isla Gama at Bahia San Blas (J. E. Jackson, pers. comm., 1979).

The west of Argentina, adjacent to the Chilean Province of Osorno, has several populations of Fallow deer. The greatest number, estimated at about 3000 in 1977, is on Isla Victoria, the largest island in Lake Nahuel Huapi which lies across the border between the provinces of Neuquen and Rio Negro. The Lake is within the Parque Nacional Nahuel Huapi, which is situated between the Andean forest and the Patogonian plateau, and lies at about 760 m above sea level. The habitat of the island includes False beech (Nofhofugus sp.) bamboos, and conifers, especially alerce (Fifzroya pafugonica). Feral Fallow are also living on the nearby Estancia Huemel which is on a peninsula of the Lake. Both populations were introduced by Sr Aaron de Anchorena, a big game hunter who had many species of deer on his Estancia, ‘La Barra de San Juan’ in Uruguay (H. R. Rivadavia, pers. comm., 1977).

About 40 km east of San Carlos de Bariloche, in Neuquen province, Fallow have been established for over 40 years at the Estancia La Primavera at Lago Trafal near Confluencia. The population in 1977 was estimated at 180 (H. R. Rivadavia, pers. comm.). Herds are also found north of the Lake at San Martin de 10s Andes and, 40 km further north, at Junin de 10s Andes. At San Martin, which lies at the eastern end of Lake Lacar, about forty Fallow are present in Parque Diana (B. Affolter, pers. comm., 1974).

The environment of the Lakes Region, in the foothills of the Andes, contrasts with the Pampas conditions experienced by the Fallow in Buenos Aires Province. Rainfall is greater with an annual average of 1035 mm. The summers are cooler and the winters colder with considerable snow. The rougher and varied terrain support a wide range of plants including coniferous forests.


2.6.2. Chile The earliest authenticated introduction of Fallow deer to Chile was made in 1887 when animals from Europe, possibly Spain, were taken to the national park near Lota, on the coast of central Chile. When their increasing numbers caused serious damage, they were transported a few kilometres to the north to the El Escuadron Reserve near Coronel, but some escaped into the surrounding Nahuelbuta mountains. The remainder were taken to the nearest small island, Santa Maria. There they thrived for 15 years, the population exceeding one hundred. A plan to round-up the deer resulted in many of them falling over the cliffs. A few of the survivors were taken to farms in the mountains near Collipulli, between Concepcion and Osorno (Schneider, 1936). None is believed to be there now (M. Urbina, pers. comm., 1976).

Following an importation from Hamburg Zoo in 1925; an enclosed herd was established in Valparaiso province where about thirty deer are still maintained. Later animals were taken from there to other provinces, especially Bib-Bib and Osorno, to stock other enclosures (J. Rottmann, pers. comm., 1973).

Although most Fallow in Chile live on farms or other privately-owned land where they are enclosed, there are free-ranging populations on at least two islands (Fig. 27). AltuB-Huapi Island, which has an area of 2500 ha and lies within Lake Ranco, near Rio Bueno in Valdivia province, is one of the islands which has Fallow. The annual precipitation is almost 2500 mm. The average temperature in January is 17OC and in the coldest month, July, it is 8°C. Just to the south, but in the province of Osorno is Lake Rupanco. On an island within this Lake a census taken in 1975

Distribution of fallo w deer 1 19

recorded a total of 200 Red deer, Fallow and Mouflon (Ovis musimon). Both islands are mostly covered by forest in which the dominant species are olivillo (Aexroxicum punctarurn), ulmo (Eucryphiu cordifolia) and Laurelia phdippianu (J. R. Rau, pers. comm., 1978; M. Urbina, pers. comm., 1978).

2.6.3. Peru Fallow occur in only one area of Peru (Fig. 27). Some were introduced in 1948 to the Huacraruco Ranch in the district of San Juan in Cajamarca Province, some 560 km north of Lima in northern Peru. After a period of acclimatization, the deer were released nearby in the Chicamo river basin. The deer became established and apparently did not decline during the 1963-64 epidemic of foot-and-mouth disease which greatly decreased the population of the native White-tailed deer. Recently, however, the Fallow have been much reduced by poaching; the population was estimated at only twenty-five in 1974. Some of these Fallow are to be maintained as a special attraction within the Sunchabamba Game Reserve which adjoins the Huacraruco Ranch (C. F. P. del Prado, pers. comm., 1974).

The mean temperature is between 10 and 15°C in July and about 20°C in January. The annual rainfall is between 100 and 400 mm.

2.6.4. Uruguay The number of Fallow deer in Uruguay is small and very little appears to be known about them except that they occur in two areas in the south-west of the country (Fig. 27).

During the 1930s, Sr Aaron de Anchorena enclosed a number of deer, mostly Axis but also some Fallow, within 500 ha on the estancia ‘La Barra de San Juan’ in the Department of Colonia, on the mouth of the San Juan River and about 200 km from Montevideo (D. J . mcks, pers. comm., 1978). Fallow deer from this estancia are believed to have been used to stock Isla Victoria in Argentina (H. R. Rivadavia, pers. comm., 1977). The enclosure no longer exists but Axis deer are widely established and some Fallow have been reported in the heavily wooded area, which con- tains some grassland, on the river banks in the vicinity of the former enclosure.

North of Colonia lies the Department of Soriano. Here, on the edge of scrubland bordering the Negro River, at least one Fallow buck is known to be living with a small herd of axis deer (A. 0. Martinez, pers. comm., 1978).

The Fallow deer habitats in Uruguay are low-lying, less than 130 m above sea level. The mean annual rainfall is about 1000 mm. In January the mean temperature is about 22°C and in July it is about 10°C.

2.6.5. Leeward Islands In the West lndies two of the Leeward Islands, namely Barbuda and Guana, have populations of free-ranging Fallow deer (Fig. 1).

Barbuda, which has an area of 155 km2 , lies 40 km north of Antigua. In the eighteenth century the island was granted to the Codrington family who introduced the deer at an unrecorded date (Aspinall, 1954). Aletter written from Antiguain 1740 to Lady Codrington,in England, (Gloucester- shire Record Office, D1610 C5) refers to ‘about a thousand head of deer’ on Barbuda. Three Fallow skins, two common colour and one black, are in the British Museum (Natural History) [Ref. 1857. 1.14.1-31, having been presented by Charles Darwin who provided the following information: ‘Domesticated and naturalized about 100- 150 years ago in the Island of Barbuda’. Darwin never visited the West Indies but probably acquired the skins when some were sent to England in an unsuccessful attempt to establish a trade. A letter dated 18 Nov. 1834 (Gloucester- shire Record Office, D1610 C40) to Sir C. B. Codrington refers to a sample skin and also thirteen others sent previously, in 1831, which resulted in an out-of-pocket loss of 4s.7d. because they were much perforated by shot.

It therefore seems that Fallow were taken to Barbuda early in the eighteenth century. Sir Hans Sloane (1 725) , recording his voyage to some West Indian Islands, mentioned that Fallow deer


were sometimes brought to Jamaica. Their place of origin was not stated but could have been Barbuda, since there were 1000 head only 15 years later. The 1740 letter discussed the shipment of Fallow, caught by netting, to Antigua where they would be sold for 2s.6d. per pound. Nine- teenth century writers refer to deer on Barbuda but do not name the species (Coleridge, 1826; Southey, 1827). Millais (1906) referred to a ‘recent’ introduction of Fallow deer to a West Indian Island but gave no further details.

The only deer mentioned by Harris (1965) in his account of the fauna and flora of Barbuda were White-tailed, but his information was hearsay (D. R. Harris, pers. comm., 1978). In 1974 Fallow were reported to be present (J. L. Robinson, pers. comm.) but no details were available. Local residents still believed the deer to be White-tailed in 1978 when Mr. David Watters began enquiries on our behalf. The descriptions, photographs and specimens that he sent to us proved beyond any doubt that Fallow do indeed live on Barbuda. At least two colour varieties, black and common, are present. They occur in all the vegetation zones described by Harris (1965) including swamp and grassland, but mostly live in the xerophytic, evergreen woodland and scrub, particularly in the so-called Highlands: the highest point on the island is only 34 m above sea level (Brasier & Mather, 1975). This area is said to be ‘barely accessible’ so has been less intensively used by man for his shifting agriculture, charcoal-burning and grazing of domestic stock. The dominant evergreen tree in this locality is White cedar (Chamaecyparis sphaeroidea): other species include whitewood (Bucida buceras) and satinwood (Chloroxylon swierenia) and a few deciduous trees are also present. Much of the scrub woodland consists of thorny trees and shrubs such as acacia, hogwood (Comocladia dodonaeu), cankerbury (Solanum racemosum var. igneum) and Black sage (Lantana camara) and spiny succulents (Harris, 1965).

Barbuda has an annual rainfall of about 1150 mm of which over half falls from August to November. The average temperature is about 29°C in mid-summer and falls only slightly, to 24OC, in mid-winter.

Guana (also called Guiana, Iguiana and Iguana) lies only 100 m off the north-west coast of Antigua, immediately south-west of Crabbs Peninsula. Its maximum dimensions are about 4 km and 1 km. Like Barbuda, it is low-lying, with a maximum altitude of only about 15 m. Mangroves and swamps fringe part of the coast but most of the island supports xerophytic trees and scrub. The climate is similar to that of Barbuda but with slightly less rainfall. The original stock of deer are believed to have been introduced by the Codringtons.

500 km Fig. 28. The locations of Persian Fallow deer in Iran


Until recently the Fuller family, who own the island, believed the forty or so deer to be White- tails. In 1978 Mr Morris Nedd, an experienced hunter of the deer on Barbuda, was taken to Guana and identified the deer as Fallow (D. Watters, pers. comm.).

3. DISTRIBUTION OF PERSIAN FALLOW DEER The Persian Fallow deer (Plate 1) is one of the rarest deer (Cowan & Holloway, 1978). An account of the history and biology of this animal has been published previously (Chapman & Chapman, 1975). In its natural habitat in the wild it is believed to survive in only one locality, in south-west Iran (Fig. 28).

Although this animal was not described in the scientific literature until just over a century ago (Brooke, 1875), there may have been a specimen living in the London Zoo 35 years earlier. Gulliver (1840) referred to a deer, the size of a full-grown Fallow, recently arrived from the Persian Moun- tains but as yet unidentified. Alternatively, this animal may have been a maral for the Annual Report presented in April 1840 refers to two new Persian deer, Cervus maral, from the Persian mountains (R. Fish, pers. comm., 1976). Maral may retain spots on the pelage until 5 years of age (Harrington, 1977) and, if not full-grown, could be similar in size to a Fallow deer. When Brooke described the Persian Fallow it was known from the riverine forests of Khuzistan to the oak forests of the neighbouring province of Lorestan in south-west Iran. A later report came from further north, from the Jauanrud district about 8 km north of Kermanshah (Ellerman & Morrison-Scott, 195 1). This distribution was much reduced from that of earlier times. Archaeological and historical evidence indicates its former existence in Iraq, Jordan, Lebanon, Palestine and Syria as well as Iran, and also, by introduction, to Cyprus (Chapman & Chapman, 1975; Harrison, 1968). This century there have been four reports from outside Iran. Two were from Iraq, namely Zakho in the extreme north in 1917 and, about 1943, from Jebel Baradost, a 40 km long ridge in Kurdistan close to the Iranian border (Hatt, 1959). An unverified report came from north of Latakia in north-west Syria in 1940 and in 1923 an antler of a Persian Fallow buck from Jerash in northern Jordan was offered for sale (Talbot, 1960). The other reports, never numerous, were all from western Iran and mostly from Khuristan province in the south-west.

By the 1940s the Persian Fallow deer was thought to be extinct. In 1955 a small population was reported from where the Sirwan River crosses the border between Iraq and Iran, about 50 km north-west of Kermanshah and about 350 km north-west of the previously most recently known localities. The deer were said to live in scrub on the Zagros Mountains up to 1300 m, occasionally coming down to a narrow valley near the river (Talbot, 1960).

In 1957 and 1958 Persian Fallow deer were reported from dense forest along the three great rivers, Dez, Karkheh and Karun, in Khuzistan province. From 1957 to 1963 five German expeditions resulted in the capture of one pair of deer. A male was captured near Harmala on the River Dez in 1957 and the following year a female was caught near Achmad-Molah on the River Karkheh. Both these animals were taken to Georg von Opel’s zoo at Kronberg near Frankfurt-am- Main in West Germany (Haltenorth, 1959, Pepper, 1964). Three Iranian expeditions were made from 1964 to 1967, resulting in the capture of six deer, three of each sex. One male was sent to Kronberg to replace the original male that had died in 1960. The others were taken to Dashte-Naz, a reserve created for them east of Sari and near the Caspian Sea in northern Iran. The dates and details of the capture expeditions and the subsequent success of the captive animals have been given by Mueller (1979). At Dasht-e-Naz the deer flourished: more than fifty were present before the 1978 fawning season. The park consists of 55 ha of woodland of the Humid or Hyrcanian type which has been described by Bobek (1968). Quercus castaneaefolia, Zelkova crenata and other broad-leaved, deciduous trees are dominant. The understorey is now much-browsed but there is a rich ground flora (Personal observation, Dec., 1977).

A buck and two does were taken in 1977 from Dashte-Naz to Ashk lsland in Lake Rezaiyeh in north-west Iran, about 1OOkm north-west of Jebel Baradost where a deer was reported about 1943. The island has an area of 2300 ha with almond (Prunus amygdalus) and pistachio (Pistacia mutica) as the dominant trees. During their first winter there the deer experienced temperatures

122

around -18°C and snow to a depth of 0.5 m. In late August 1978 one female was found dead (S. Hess, pers. comm., 1978).

Two adult males and five adult females bred in Germany were sent to Dasht-e-Naz in March 1973. They and their progeny, together numbering twenty-eight in the autumn of 1977, are now established in 6 ha of fenced deciduous woodland at Semeskandeh, between Sari and Dasht-e-Naz. Formerly the arrivals from Germany were kept at Dashte-Naz, but in a section of the park separated by a high fence from the deer which originated from south-west Iran. This segregation was maintained as there was some doubt as to the genetic purity of these deer. In Germany some hybrids between Persian and European were produced and, although the zoo's veterinarian main- tains that all the deer sent to Iran were pure-bred, the genetic character of the animals sent to Iran remains suspect (Department of the Environment, Iran, 1978). In 1976 three Persian Fallow were sent from Germany to Israel and the following year a pair went to East Berlin (Mueller, 1979).

About the time that the Dasht-e-Naz park was established to breed the Persian Fallow deer in captivity, two areas were given protection by the Iranian Department of the Environment to aid the survival of the deer in the wild. Along the River Dez, 524 ha were declared a Wildlife Refuge with a further 10 633 ha adjoining as a Protected Area. Along the neighbouring River Karkheh, 3600 ha became a Wildlife Refuge with a further 9427 ha as a Protected Area. These measures were intended to protect the remaining habitat and prevent poaching. Game guards were appointed to patrol the areas and a heavy fine (c.E2000) was imposed for shooting a Persian Fallow deer.

The Dez Wildlife Refuge lies on a broad plain, west of Dezful and within sight of the western foothills of the Zagros mountains. It consists of a strip of vegetation on each bank between the River Dez and the desert. The width of the strip is mostly about 100 m but in some places is much less and elsewhere may extend to a kdometre. Nearest the river is the zone of forest thicket where poplar (Populus euphratica) is the dominant species. Parallel to the forest lies a zone of scrub with tamarisk (Tamarix sp.) as the dominant species but also other xerophytic shrubs such as Capparis spinosa, Prosopis farcta and Schanginia baccata. The scrub is bordered by desert which supports a few small, scattered, xerophytic plants. Serious loss and degradation has occurred in this habitat which is but a remnant of once-vast riverine forests. Outside the Refuge, natural vegetation is very sparse and the domestic stock from the several neighbouring settlements invade the scrub and forest, entering through breaks in the barbed-wire perimeter fence. These animals include sheep, donkeys, goats, water buffaloes and camels (Camelus dromedarius), all of which have a very marked effect on the vegetation by browsing, trampling and preventing natural regeneration (Personal observation, Dec., 1977). Areas of the once impenetrable forest, as well as the scrub, have been opened up by these animals. In 1977, to serve new sugar and paper factories, a road, a railway and a bridge to carry them over the river, were constructed through the middle of the Refuge, destroying the habitat for a width of 100 m. This area previously was favoured by bucks as evidenced by the finding of cast antlers. The numbers of Persian Fallow deer remaining in the reserve was estimated at not more than twenty-five in May 1978. There was a proposal to con- struct a new boundary fence of woven wire to exclude domestic stock and to prevent entry by wood-cutters (S. Hess, pers. comm., 1978).

The Dez Wildlife Refuge is less than 200 m above sea level. The mean temperature in January is about 6°C and in July about 30°C. Most of the annual rainfall of about 350 mm falls from Nov- ember to March; very little falls from June to September.

The habitat at the Karkheh Refuge was very similar to that at Dez (Haltenorth, 1961) but has been subject to even greater pressures resulting in loss of habitat. By 1977 it was thought no Persian Fallow deer survived at Karkheh.

In 1978 plans were made to establish Persian Fallow at another site, to the south-east, in the Zagros Mountains. Initially an enclosure would be constructed at Dashte-Barm, west of Shiraz and within the Arjan National Park. This valley is forested with oaks and probably would have been within the former natural range of the Persian Fallow deer before the loss of so much of the semi-humid, deciduous oak forest. The enclosure would be stocked with some animals from Dasht-



e-Naz and subsequently, when the surrounding land had been purchased and its natural vegetation reestablished, the deer would be liberated. The likelihood of the plans corning t o fruition, a t least in the near future, now seems remote.

Whilst the Persian Fallow deer has been saved from extinction by breeding in an enclosed reserve, its survival in the remnants of its natural habitat remains precarious.

4. DISCUSSION The present-day distribution of the European Fallow deer extends t o thir tyeight countries, spread over six continents. They occur between latitudes 61°N and 46"s and in places as diverse as the tropical Fijian island of Wakaya and the cool temperate regions o f southern Sweden (Fig. 1).

Table 4

Vegetation types and their use by Fallow deer

Type*

Extra-tropical Coniferous forests

Mixed forests

Broad-leaved forests

Tundra and Alpine vegetation

Grasslands

Woodlands & scrublands

Tropical Tropical rain forest

Tropical seasonal forests

Thorn forest, semidesert and desert

Savanna

Country/State

E. Germany, England, Georgia, Netherlands, New Zealand, N. Ireland, Republic of Ireland, Scotland, S. Australia, Sweden, Wales

Alabama, Argentina, Austria, Bulgaria, Canada, Chile, Denmark, England, Georgia, Hungary, Italy, Netherlands, New Zealand, N. Ireland, Poland, Rhodes, Rumania, Scotland, S. Australia, S. Africa, Spain, Sweden, U.S.S.R., Wales W. Germany

Austria, Belgium, Bulgaria, Denmark, England, E. Germany, France, Georgia, Hungary, Kentucky, Nebraska, Netherlands, N. Ireland, New Zealand, Poland, Queensland, Republic of Ireland, Rumania, S. Australia, Scotland, Sweden, Tasmania, Wales, W. Germany, Yugoslavia

New Zealand

Argentina, California, S. Africa, S. Australia, Spain, Texas

California, New Zealand, S. Africa, Texas, Turkey

Fiji, Leeward Islands, Queensland

New South Wales, Queensland, S. Australia, Tasmania, Victoria

Tropical mountain vegetation

* Lyre (1968)

124 N. G. Chapman and D, I. Chapman

Very few, if any, of the places in which Fallow deer live support a completely natural vegetation. Almost everywhere the vegetation has been modified to some extent either by man or his domesticated stock. The description of the habitats, therefore, presents problems because the various classifications for world vegetation are concerned with natural climax vegetation. The situation is further complicated by the fact that the deer rarely restrict themselves to one type of vegetation. Their habitats are usually mosaics of different types of vegetation utilized at various times, possibly for different purposes. In England, for example, deciduous broad-leaved forest is regarded as the deers’ typical habitat but they also spend time feeding or sheltering in arable fields (Plate 7), pasture, heathland, mixed forest and coniferous forest, thus taking advantage of whatever is available.

Fig. 29. The types of vegetation inhabited by Fallow deer

Only broad categories for the main vegetation types utilized by the deer are considered here. The classification of Eyre (1968) is shown in Table 4, together with the countries in which Fallow occur in each habitat. These categories do not take into account localized conditions where edaphic factors influence the vegetation. Consequently marsh, swamp, mangrove and heath do not fall within Eyre’s classification. Where such habitats are of importance to Fallow deer they are mentioned in the appropriate section.

In the context of this discussion on vegetation, the word ‘country’ also includes individual States of Australia and the U.S.A. Within one country, the deer may occur in several major types of vegetation. For a few places there was insufficient information available to us on which to classify the vegetation of the habitat. Finland, Norway and Uruguay have, therefore, been omitted but their inclusion is unlikely to have added to the number of vegetation types used by the deer.

Of the eleven major vegetation types, Fallow are absent from only three, namely Tropical Rain Forest, Tropical Mountain Vegetation, and Thorn Forest, Semi-Desert and Desert. Fallow occur within all six of the extra-tropical types (Fig. 29). Figure 29 has been derived by expressing the number of countries in which Fallow occur in a particular type of vegetation as a proportion of the total number of countries recorded for all types of vegetation. (See Table 4). The result is somewhat artificial in that it depends on political boundaries and takes no account of either the size of the country or the density of deer. The Figure does, nevertheless, indicate that whilst forest, particularly Mixed or Broad-leaved types, forms the main element of the animal’s habitat in most countries, Fallow are evidently able to live within a considerable range of vegetation types.

Eyre (1968) uses the term Coniferous Forests to embrace Boreal, Subalpine, and Lowland and Montane North American coniferous Forests. The term is extended here to include the coniferous

Distribution offallow deer 125

forests created by man. Fallow do not occur within the Boreal Forest characteristic of northern Eurasia and North America where winters are cold and the growing season short. In many countries, however, they live within man-made coniferous forests in which the main genera include those typical of Boreal Forest, such as Pinus, Abies and Lark.

Mixed Forests, composed of broad-leaved and coniferous trees, account for some of the Fallow deer habitats in twenty-five countries. The Evergreen Mixed Forests of the Mediterranean region, as on Rhodes, and the Evergreen Mixed Forests of the Southern Hemisphere, as in some Fallow deer areas of Argentina, Chile and New Zealand, are composed of trees in which the broad-leaved species, as well as the conifers, are evergreen. In a greater number of countries, Fallow deer live in Mixed Forests composed of conifers and deciduous, broad-leaved trees. Forest of this composition occurs naturally in Northern and Central Europe and North America, forming a Transition between coniferous and broad-leaved, deciduous forest and is known as Ecotone Mixed Forest. Some Fallow deer habitats in Poland are of this type. In many other countries, forests of similar composition have resulted from sylvicultural practices, often by the creation of conifer plantations within or in place of natural broad-leaved, deciduous forest.

Broad-leaved Forests form the major part of Fallow deer habitat in the same number of countries as do Mixed Forests. In most of the countries this type of Forest is composed of deciduous species (Plate 6). This is to be expected as broad-leaved, deciduous forest is the climax vegetation for much of Europe and the distribution of Fallow deer is predominantly European. In a few countries, such as Georgia, New Zealand (Plate 4) and Australia, the trees are evergreen.

Tundra and Alpine Vegetation consists of the grasses, sedges, mosses and lichens occurring in the Arctic, Sub-kctic, Antarctic and Alpine regions. Fallow occur within this vegetation type only in a few places in New Zealand where, at some seasons, they may leave the lower forested slopes to seek food from the grassy vegetation at higher altitudes.

Although areas of grass form part of the deer’s environment in almost all its habitats, in only a few is natural Grassland the major vegetation type. In part of Texas they occur within Prairie grassland which is characteristically composed of tall species. Several grassland formations are classified as Extra-tropical grasslands of the Southern Hemisphere. Within this type Fallow occur in the Pampas of Argentina where in hilly and dissected country, including part of the Sierra de la Ventana (Plate 1 l ) , grassland may give way locally to shrubby vegetation. In South Africa they are found in the Veld, both the Bushveld where trees and shrubs are scattered through the grassland, and the Highveld where tall grasses give way in places to bracken, scrub and eventually forest. Elsewhere the grassland generally available to Fallow is small areas of natural grass, as in a glade or edge of a forest, pasture used by domesticated stock or neglected pasture, as at Point Reyes in California. Marsh is a wet grassland (de Laubenfels, 1975) and, although outside Eyre’s classification, is mentioned here as it forms a major part of the Fallow deer habitat at Cot0 Doiiana, Spain (Plate 5).

The Woodlands and Scrublands of Eyre’s classification include the Sclerophyllous Formations of the Mediterranean marquis, the chaparral and the fynbos which respectively form Fallow deer habitats in Turkey, California and the south-western part of Cape Province in South Africa. Also included in the Woodlands and Scrublands type is the Semi-Desert Scrub and Woodland. This occurs in Fallow deer areas on the Edwards Plateau in Texas (Plate lo), and in California where evergreen species are dominant.

Of the five Tropical vegetation types, Fallow are present in only two. They live in Tropical Seasonal Forest on Barbuda and Guana in the Leeward Islands, on Wakaya in the Fijian islands and also in northern Queensland. All these areas are subject to a dry season but northern Queensland has a high annual rainfall and the forest is called monsoon forest.

The word ‘tropical’ is not necessarily applied solely to the region between the two Tropics. Consequently, the Savanna which is primarily associated with truly tropical regions, is also used to describe the association of tall grasses and Eucalyptus or Acacia species found in parts of New South Wales, Victoria and South Australia where Fallow occur.

The different types of vegetation reflect, in part, a range of climatic conditions. Of the five


Table 5 The classification of climatic zones and occurrence of Fallow deer

~ ~ ~~

Climatic zone Code Characteristics

Tropical A Rainy climate, Humid no winter,

coolest month >18"C

B Dry climate, excess evaporation over precipitation

Warmer C Warm temperate, Humid rainy climate,

mild winters, coolest month >O°C but < 18"C, warmest month > l o o c

Cooler Humid

Polar

D Rainy climate, severe winters, coldest month

warmest month <ooc,

> 1 ooc E No warm season,

warmest month 1 0°C

~

Country/State

Leeward Islands, Queensland

Fiji, New Mexico, South Africa

Alabama, Argentina, Belgium, Bulgaria, California, Canada, Chile, Czechoslovakia, Denmark, England, France, German Democratic Republic, German Federal Republic, Georgia, Ireland, Italy, Kentucky, Netherlands, New South Wales, New Zealand, N. Ireland, Peru, Portugal, Queensland, Rhodes, Scotland, S. Africa, S. Australia, Spain, Tasmania, Texas, Turkey, Uruguay, Victoria, Yugoslavia, Wales

Austria, Finland, Hungary, Massachusetts, Nebraska, Norway, Poland, Romania, Sweden, U.S.S.R.

major climate zones designated by Koppen (Boucher, 1975), Fallow deer occur within four, namely Cooler Humid, Warmer Humid, Dry and Tropical Humid. They are absent only from the Polar zone. Figure 30 shows the number of countries within each zone where Fallow deer occur. In this context 'country' again includes individual States of Australia and the U.S.A. Within one country Fallow may occur within more than one climatic zone (Table 5).

In thirty-six countries, at least some of the habitats are within the Warmer Humid zone. This zone includes the temperate regions with a rainy climate, mild winters with the average temperate above 0°C but below 18°C in the coldest month, and above 10°C in the warmest month.

Relatively few Fallow deer habitats lie within the other climatic zones. The Cooler Humid climate is experienced by Fallow in eight European countries and two states of the U.S.A. These places, which apart from Massachusetts and Nebraska are mostly in northern or eastern Europe, have a mean mid-winter temperature below freezing point. Fallow occur within the Dry zone only on Wakaya in the Fijian islands, in some areas within Cape Province in South Africa and possibly in New Mexico where their precise location is unknown.

The only Fallow deer areas subject to a Tropical Humid climate are in northern Queensland and on Barbuda and Guana in the Leeward Islands. A characteristic of this zone is the lack of a cold season, the coolest month having a mean temperature greater than 18°C. Although annual rainfall is hgh, these three places have a marked dry season.

Distribution of fall0 w deer 127

Climatic Zones

30. The climatic zones in which Fallow deer occur. A: Tropical humid, B: Dry, C: Warmer humid, D: Cooler humid, E: Polar. *Including states in the case of Australia and U.S.A.

Fig.

The widest range of temperature that Fallow deer experience in one area may be in Argentina. In 1972, the only year for which we have been able to obtain data, the maximum and minimum temperatures recorded in Sierra de la Ventana were 40°C and -9°C (E. E. Willard, pers. comm., 1977). Summer temperatures around 30°C are usual at Cot0 Doiiana in Spain, Fiji, the Leeward Islands and Rhodes, but none of these places has cold winters so the annual range of temperatures is not great. In northern and eastern European countries where average winter temperatures are sub-zero, the summers are cool with an average temperature of less than 2OoC, so in neither of these groups of countries is there normally a very wide seasonal range in temperature. If extremes rather than average temperatures are considered, there are a number of countries where, in some years, the annual range may be between 40 and 50 degrees.

The area in which Fallow experience the highest rainfall is on the Paporoa Range in the South Island of New Zealand where, in some places, 6000 mm may fall in 1 year. Over 2000 mm fall in the Fallow deer habitats in Chile and northern Queensland and more than 1000 mm is experienced in a number of widely separated places such as Alabama, Austria, South Australia, Victoria and Yugoslavia. In the majority of the deer habitats the annual fall is between 500 and 1000 mm.

The absence of Fallow from Polar climates is to be expected: despite its wide distribution by introduction around the world the species has not been released in Tundra or Ice Cap conditions. Fallow are also absent from Subarctic climates, a subdivision of the Cooler Humid zone where less than 4 months in a year are above 10°C. Such conditions, with heavy and persistent snow occur in much of Norway, most of Finland and northern Sweden. Fallow do exist within these three countries but only in the southern parts where milder conditions prevail. In Finland the Fallow deer are near the northen limit of the Continental Cool Summer climate (a subdivision of the Cooler Humid zone) with its constantly moist atmosphere, and the warmest month averaging less than 22°C. This also applies to the more northerly of the localities within southern Sweden. In the southernmost part of Sweden, Skine, the number of days with snow cover is, on average, 40 but this period is seldom continuous and the mean January temperature is 0°C. Where Fallow occur in Norway, on the island of Hanko, conditions are similar to much of west and central Europe with constantly moist conditions and the warmest month averaging less than 22°C. On Hanko food has to be supplied to the deer in winter, but this may reflect the density of the deer in the available habitat rather than the severity of the climate.

Several attempts to establish new populations of Fallow deer in south and west Norway have failed due, it was thought, to their inability to tolerate the deep snow (J. A. Pedersen, pers. comrn., 1973). In contrast, Red deer occur as far north as 60"N in Norway but further east, in Sweden, eastern Europe and the U.S.S.R. to about 55"N. Similarly, the northern limit of Roe deer reaches 64"N in Scandinavia but 60"N or less in eastern Europe and Asia (Flerov, 1952). The depth of

128 N. G. Chapman and D. I . Chapman

snow has been claimed to be a critical factor in limiting the distribution of both these species of deer. Snow depth was considered to be more important than temperature, topography or vegetation for Roe deer and a depth of 50 cm limits their distribution (Flerov, 1952). Even the much larger Red deer is said to find movement and feeding difficult in snow deeper than 25 or 30 cm and a depth of 50-60 cm may determine their distribution (Heptner et aZ.. 1966). Observations during normal and unusually severe winters in Norway and Sweden, however, led Ahlen (1965) to con- clude that the limit of the distribution of Red deer was determined by the snow conditions and associated availability of food, not merely the depth of snow.

Roe and Red deer both occur on the rough hill land of Scotland whereas Fallow do not. After the last Ice Age Roe and Red deer colonized much of Europe including Britain which they reached before it was separated from the continent of Europe about 8000 years ago. In contrast, Fallow deer are not indigenous to Britain and th is appears to be true for much of western Europe. These facts suggest that there may be fundamental differences between the climate and/or habitat requirements of Fallow deer and those of Roe and Red deer. Such differences are more likely to account for the failure of attempts to introduce Fallow to the snowier parts of Norway, in situations where Roe and Red deer occur. It is surprising if the Fallow, which is intermediate in size between Roe and Red deer, cannot cope with a similar depth of snow.

Although snow occurs in very many of the places where Fallow live, including most of Europe, it does not normally cover the ground to much depth for a long period of time. Where the snow does occur regularly and persists, the habitats include forest so the deer still have shelter.

High temperatures are not believed to have been a limiting factor on the survival of Fallow in any of the areas to which they have been introduced.

The ability of Fallow to survive within a fairly wide range of climatic conditions has contributed to their acclimatization in many countries far from their original home. In at least one country their adaptation to the local climate was, under some circumstances, superior to that of the native ungulates. Vincent (1962) referred to the ability of Fallow to withstand a period of ‘extreme cold’ when springbok and blesbok as well as sheep succumbed in the Cradock district in Cape Province, South Africa. The severity of th is cold spell was not stated but the lowest temperature recorded in Cradock was -7°C for six consecutive nights in June 1956 (J. Bothma, pers. comm., 1979).

Table 6 Dispersal rates of Fallow deer

Country Period Rate (years) km/year

Source

Australia Lake George

England Essex

German Federal Republic

Schleswig-Holstein New Zealand

Paporoa Wanganui

Alabama California

Nannau

U.S.A.

Wales

92

<30

32

10 >5 0

30 21

8

0.8

0.6

0.9

0.9 0.8

0.9 0.5

0.9

Allison (pers. comm. 1978)

Chapman (1977)

Heidemann ( 1973)

Clarke (1976) Caughley (1 963)

Allen (pers. comm. 1974) Jurek (1 977)

Vaughan & Carne ( 1 97 1 )

Mean f SD 0-8f0.15


The approximate rate of dispersal can be calculated from a knowledge of the date of the liberation or escape of the deer and the date when they were first recorded furthest from that place. Dispersal rates for Fallow are given in Table 6. These figures, taken from several countries with differing habitats and over widely varying periods of time, are surprisingly uniform and give a mean rate of dispersal of 0.8 km/year (SD * 0.1 5). Consequently, Fallow deer populations are generally found close to the places to which they were introduced or from which they or their ancestors escaped. Similar rates of dispersal (0.6-1.0km/year) have been recorded in New Zealand for wapiti (Cervus canadensis), sambar, Timor deer and White-tailed deer whilst Red deer and Sika deer achieved a rate of 1.6 km/year (Caughley, 1963).

The present world-wide distribution of Fallow deer is due almost entirely to man and it is of interest to consider some of the reasons for these many introductions. All the introductions to places outside Europe, apart from the Leeward Islands, occurred during the nineteenth and twentieth centuries. The introductions to Australia, New Zealand and South Africa are associated with the period of British colonization. The reasons for taking Fallow across the world were varied and often fulfilled several aims. The primary purpose was usually for sport, particularly where there were no comparable endemic species to hunt, as in Australia and New Zealand. Many of the settlers had a desire to make their new surroundings as much like their homeland as possible. In some instances they went as far as creating a 'little England' complete with deer, foxes and tra- ditional sports such as hunting, coursing and horse-racing (Rolls, 1969). Even for the less ambitious, a herd of Fallow deer on their properties doubtless added to their status, gave an attractive re- minder of the stately homes of England and provided a palatable addition to the diet. Ecology was an unknown science at that time and there was evidently no appreciation of the uniqueness of the native fauna. Encouragement for the introduction of exotic species was especially strong in Australia and New Zealand where there were very active Acclimatization Societies pursuing their objects of '. . . the introduction, acclimatization and domestication of all innoxious animals, birds, fishes, insects and vegetables whether useful or ornamental . . .'. Even in South Africa with its abundance of native ungulates suitable for hunting and eating, the desire for the atmosphere of an English estate prompted the introduction of Fallow. In North America Fallow were introduced, despite the wide distribution of White-tailed deer and Muledeer, to areas where there was believed to be an abundance of habitat not fully stocked by the native species.

In Europe introductions of Fallow have taken place over a long period, with examples known from most centuries from the eleventh to the twentieth. The need to introduce them to Europe at all needs an explanation because during the Last (Ipswichian) Interglacial, Fallow deer, slightly larger than those of to-day, were widespread in Europe, being known from England, Denmark, France, Germany, Gibraltar, Greece, Italy, Poland and Georgia and Armenia in the U.S.S.R.

Then followed the Wurm glaciation which lasted around 60 000 years before it ended about 10 000 years ago. During this glacial period there were many climatic oscillations. The ice sheet was not as extensive as in earlier glaciations: at the coldest stage the ice covered most of Ireland and Wales, extended to central England and the north Norfolk coast, across the North Sea to Jutland and north Germany. Further to the northeast the ice sheet joined that from the Urals region. In southern Europe, the Pyrenees and Alps had their own ice sheets. Nevertheless, the influence of the glaciation was far-reaching and affected most of Europe with discontinuous permafrost extending at least as far south as latitude 46" (Butzer, 1964). Some places such as central Spain, northern Portugal and northern Italy supported coniferous forest while temperate deciduous forest existed around the Mediterranean in southern Italy, Greece and Turkey. Further east, in the western U.S.S.R., grassland of steppe and prairie types occurred (John, 1977). Although Fallow survived in a few places during the Wurm, there is no evidence for their continuous existence in southern Europe right through this period. It has been suggested that even there they became extinct late in the Wiirm glaciation, only surviving further east in Asia Minor (Kowacski, 1967; Kur th , 1968).

Of 165 European post-glacial archaeological sites of Mesolithic age, 96% yielded bones of Red deer and over 60% had bones of Roe deer but none contained Fallow bones (Jarman, 1972). The


distribution of Fallow after the glaciation was very different from its widespread occurrence in Europe before the Wurm glaciation. Why Fallow did not recolonize Europe after the last ice age, as did Roe and Red deer, remains a mystery. It has been suggested that with the onset of glacial conditions, the Fallow would have been one of the first mammals to retreat (Sutcliffe, 1957) and hence presumably one of the last to return. We have alluded already to possible differences between the habitat requirements of Roe and Red deer and Fallow.

Whether or not Fallow moved naturally from southern Europe, if they did survive there at all, or from Turkey after Mesolithic times is unknown. In any case, the extent of their movements must have been small, otherwise the many introductions to European countries in later centuries would not have been necessary, unless they had been hunted to extinction. In Bulgaria, for example, Fallow were introduced as recently as 1904 (Sartorius, 1970). If the deer had spread naturally into Europe, one would have expected Bulgaria, being next to Turkey, to have been one of the first countries to have been colonized. It seems probable that in Europe, as in the other continents, the distribution of Fallow is largely the result of man’s introductions. The history of these introductions to various European countries would repay investigation.

In many places the free-ranging populations of Fallow deer owe their origin to park deer, usually by accidental escape rather than deliberate release. This correlation between wild herds and the proximity of enclosures or former parks is well-known in Britain (Chapman, 1977; Whitehead, 1964) but has also been reported in other countries including Argentina (H. R. Rivadavia, pers. comm.), the Netherlands (W. A. Weyland, pers. comm.), U.S.A. (Allen, 1965) and West Germany (Ueckermann & Hansen, 1968).

Although most liberations and some escapes have led to the establishment of Fallow in the wild, some have been more successful than others. In New Zealand and Argentina they have proved to be particularly successful in acclimatizing. In the Blue Mountains of New Zealand the Fallow lacked competition and, despite shooting from an early date, they prospered and presented a pest problem in areas intended for afforestation (Donne, 1924). In 1923 alone nearly 4000 were shot. Hunting continued, yet in 1957-58 within one forest of 8000 ha, 6000 Fallow were killed (Baker, 1972). In the Sierra de la Ventana in Argentina the deer share the mountain grassland habitat with native ungulates but even so, and despite hunting and the lack of woodland, the estimated population in 1972 wasnearly 6000 within 195300 ha (S. A.Sofontas,pers. comm., 1974). Where the environment is suitable, Fallow are capable of establishing very large populations.

Whereas most introductions have succeeded, a few have failed. Too much hunting too soon, before populations were large enough to sustain the reduction, caused the demise of the deer in a few places in the U.S.A. In the Northern Territory of Australia, predation by dingoes on fawns has been suggested as a reason for the failure of the Fallow to become established (Calaby & Keith, 1974). In Western Australia, the deer died out for unrecorded reasons a few years after an apparently promising beginning (Bentley, 1978). In some parts of Australia, however, the deer were so successful that they increased to pest proportions and consequently were exterminated by man.

The number of the Persian Fallow deer are in complete contrast to those of the European subspecies. Their decline is almost certainly due to the loss of suitable habitat, which has resulted from the activities of man and his domestic stock. A similar situation exists with the European Fallow in Turkey. Given a suitable habitat, such as that at Dasht-e-Naz, the Persian Fallow is capable of breeding and increasing. They also breed well in close captivity (Mueller, 1979; Walther, 1961). With the present (August, 1979) problems in Iran the creation of a further new reserve would seem very unlikely in the immediate future and the outlook for the wild Persian Fallow is poor.

5 . ACKNOWLEDGMENTS We are most grateful to very many people for the assistance they have given in various ways. Translations have been kindly undertaken by Kate Adrian, P. Belman, J. Cerhan, Heather Crowter, the Embassy of the Socialist Republic of Romania in London, Gina Estrada, Wary Footit, R. Mc- Creedy, Rosa Musgrave, Stephanie Mole, J. G. Morris, Ghizela Ringrose and Anne Yannoulis.


The following generously provided the photographs attributed to them: F. Alvarez, J. K. Fawcett, S. Hess, J. M. Jurek, G. Moore, P. Murrell, New Zealand Forest Service, R. von Schultz, Tennessae Valley Authority, Texas Parks and Wildlife Department, E. E. Willard, S. R. Worsfold. Mr P. Bangs drew the maps and Dr J. M. Lock kindly piovided advice on the classification of world vegetation. Many Meteorological Bureaux provided climatic data. The Leverhulme Trust Fund are thanked for an award of a Research Grant to N.G.C. which enabled much of the information to be obtained.

We thank the following people and organizations who either provided information about the distribution of Fallow deer or read the manuscript for the areas attributed to them:

Africa. Madagascar: R. Albignac, Laboratoire de Zoologie, Montpellier; J. M. Andriamampianina, Direction des Eaux et For& et de la Conservation des Sols, Madagascar; Sir Hugh Elliott, I.U.C.N. Morocco: A. Elkadiri, Ministere de I’agriculture. South Africa: Prof. R. C. Bigalke, Dept. of Nature Conservation, University of Stellenbosch; J. Bothma, Dept. of Transport; Dr. J. David, Dept. of Zoology, University of Cape Town; Natal Parks, Game & Fish Preservation Board; H. J. Richards, Nature Conservation Division, Transvaal Administration; South African Department of Prisons.

Asia. Iran: R. Fish, Librarian, Zoological Society of London; S. Hess, formerly Dept. of the Environment, Dasht-e-Naz. Israel: Dr. G. Ilany, Chief Zoologist, Nature Reserves Authority. Japan: Dr. T. Ito, Tohoku University.

Australasia. Australia: C. Allison, Rozelle, New South Wales; A. Bentley, The Victorian Deer Conservation Coaperative Ltd.; J. Conquest, Oaklands Pk., S. Australia; Dr. W. P. Crowcroft, Director, Taronga Zoo; B. C. Gepp, Woods & Forests Dept., S. Australia; G. W. Saunders, National Parks & Wildlife Service, Queensland. Fiji: Mr & Mrs. Bill Beckon; F. Marseu, Ministry of Agri- culture, Fisheries & Forests. New Guinea: Dr. E. Lingren, Dept. of Agriculture, Stock & Fish, Papua New Guinea. New Zealand: Dr. L. H. Harris, N.Z. Forest Service; R. Pittaway, Wanganui.

Europe. Austria: Dr. K. Bauer, Naturhistorisches Museum, Vienna W. Herbst, Haus der Natur, Salzburg. Belgium: Dr. S. de Crombrugghe, Groupe de Travail pour 1’Etude de 1’Equilibre Forgt- Gibier. Bulgaria: E. Clay, British Embassy, Sofia. Cyprus: L. 1. Leontiades, Department of Forests. Czechoslovakia: Prof. V. Mimra, Research Institute of Forestry & Gamekeeping, Trutnov. Denmark: Dr. H. J. Baagge, Zoologisk Museum, Copenhagen; Dr. D. Clausen, Stateus Veterinaeve Serum Laboratory, Copenhagen; Dr. H. J. Degn, Funen; Dr. B. Jensen, Game Biology Station, Kald; H. K. Jensen, Gamekeeper, Wedellsborg; N. H. Lindhard, Limfjordsmuseum; J. Skaarup, Langelands Museum. Finland: Dr. R. Harrington, Forest & Wildlife Service, Republic of Ireland. France: A. F. Dolby, Information Officer, Commonwealth War Graves Commission; R. Combe, Office National de la Chasse; E. Heil, Association FCdCrative Regionale pour la Protection de la Nature; J . Jomier, Office National des Forkts, Compikgne; E. Le Chatelier, Office National des For& Aixen-Provence; A. Waechter, Laboratoire de Psychophysiologie, Strasbourg. German Democratic Republic: Dr. A. Siefke, Emst-Moritz Arndl-Universitat Greifwald. Greece: D. Sideridis, Forestry Dept., Ministry of National Economy. Hungary: Dr. V. Anna, Janus Pannonius Muzeum, Ptcs; Dr. 1. Heltay, Re- search Station for Game Biology; Dr. M. Papp, Nagykunsig Forestry Estate, Szolnok. Italy: Dr. P. L. Florio, World Wildlife Fund, Rome; Prof. R. Massoli-Novelli, Cagliari; Dr. A. F. Pratesi, Conser- vation Committee, World Wildlife Fund. Liechtenstein: B. Konrad, Office National du Tourisme. Luxembourg: R. Van den Kerchove, Parc a Gibier, Hosingen. Netherlands: Dr. W. A. Weyland, Rijksinstituut voor Natuurbeheer, Arnhem; E. P. Uittenbroek, Leiden. Norway: T. W. Johannessen, Det Norske Meteorologiske Institut; J. A. Pedersen, Zoologisk Museum, University of Oslo. Portugal: J. F. Bugalho, Director of Game Dept., Lisbon; G. B. Gooch, Afife. Poland: Prof. Z. Jaczewski, Research Station of Polish Academy of Sciences, Popielno. Republic of Ireland: F. Mulloy, Dept. of Lands, Dublin; Lord Revelstoke, Lambay Island. Romania: I. M. Bodea, Directa Generala a Silviculturii. Spain: Dr. F. Alvarez, Estacion Biologica de Dofiana; J. De la Pefia Payi, Subdireccion General de Recursos Naturales Renovables. Sweden: Prof. I. Ahlen, The Swedish

132

University of Agricultural Sciences; Dr. K. Curry-Lindahl, UNESCO Field Science Officer; Dr. R. Gerell & Dr. H. Kristiansson, Dept. of Zoology, University of Lund; B-0. Stolt, Naturhistoriska Riksmuseet, Stockholm; R. Von Schultz, Gamekeeper, Oland. Switzerland: Dr. R. Schloeth, Commission FBdbrale du Parc national suisse. United Kingdom: England: C. E. Bourchier, Estate Manager, Stanton Park; Prof. L. M. Cantor, University of Technology, Loughborough; N. Dewhurst, British Deer Society, Southport, Lancs.; C. Hart, Coleford, Gloucestershire; W. F. P. Hugonin, The Northumberland Estates, Alnwick; R. Jamie, Agent, Locko Park; R. J. Nickerson, Read’s Island; D. R. Warren,Nottinghamshire;G. M. V. Winn, Nostell Estate, Wakefield. N. Ireland: Dr. J. S. Fairley, Zoology Dept., University College, Galway; C. S. Kilpatrick, Irish Deer Society. Scotland: A. Allison, Central Scotland Branch, British Deer Society; D. Bridges, Lothian Estates, Jedburgh; P. A. Hope Johnstone of Annandale, Dumfriesshire; P. A. Joynson, Manager, Scarba Estate, M. R. M. Leslie, Factor, Portland Estate, Berriedale. Wales: M. J. Green, Bodorgan Estate. U.S.S.R.: Prof. V. G. Heptner, Museum of Zoology, Moscow University. Yugoslavia: B. Andrej, Biotehniike Fakultete, University of Ljubljani.

North America. Canada: G. W. Smith, Dept. of Recreation & Conservation, British Columbia; J. H. Todd, Sidney Island. USA. : Alabama: R. H. Allen, Dept. of Conservation & Natural Re- sources. California: H. W. Elliott, University of California. Colorado: A. E. Anderson, Dept. of Natural Resources. Georgia: T. Kile, Dept. of Natural Resources. Illinois: Prof. W. D. Klimstra, Southern Illinois University. Indiana: L, Johnson, Middlebury. Kentucky: J. S. Dwell. Dept. of Fish & Wildlife Resources; Dr. R. W. Nall, Resource Projects Manager, Land Between The Lakes. Louisiana: J. W. Farrer, Wildlife & Fisheries Commission. Maryland: R. L. Miller, Dept. of Natural Resources. Massachusetts: J. J. Mchnough, Division of Fisheries & Game; R. E. Woodruff, Vineyard Conservation Society. Nebraska: K. Menzel, Game & Parks Commission. New Mexico: W. A. Snyder,Dept. of Game & Fish. Oklahoma: J. Hammond Eve, Dept. of Wildlife Conservation. Texas: A. Briscoe & J. L. Cooke, Parks & Wildlife Dept.


South & Central America. Argentina: B. Affolter, San Martin de 10s Andes; J. M. Gallardo, Museo Argentino de Ciencias Naturales; Dr J. E. Jackson, Proyecto Venado, El Museo, La Plata; H. R. Rivadavia, Buenos Aires; S. A. Safontas, Tornquist; Prof; E. Earl Willard, University of Montana. Chile: J . R. Rau, Instituto de Zoologia, Valdivia; J . Rottmann, Unidad de Vida Silvestre Corpor- acibn Nacional Forestal; M. Urbina, Museo de Concepcibn. Peru: C. F. P. Del Prado, Ministerio de Agriculture. Uruguay: A. 0. Martinez, Direcci6n Forestal, Parques y Fauna. Leeward Islands: Drs N. & J. Fuller, Guana Island; Dr. D. R. Harris, Dept. of Geography, University College, London; Miss J. Ingles, British Museum (Natural History), London; N. Morris, Barbuda; Dr. J. L. Robinson, Veterinary Officer, Antigua; B. S. Smith, County Archivist, Gloucestershire Record Office; D. Watters, University of Pittsburgh.

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NOTE ADDED IN PROOF

Massachusetts, U.S.A. (p. 114) Fallow deer are now extinct on Martha’s Vineyard (R. E. Woodruff, pers. comm., 1979).

Plate 1. Persian Fallow deer a t Dasht-e-Naz, northern Iran. Buck in summer pelage and velvet antlers showing some palmation at the proximal end of the antlers. Photo: S. Hess.

Plate 2 European Fallow deer. Buck in summer pelage and velvet antlers showing palmation at distal ends only. Photo: J. K. Fawcett.

Plate 3. Tasmania. Typical Fa!low deer habitat with scattered trees and pasture near to forest, Near Ross in the Midlands. Photo: P. Murrell.

Plate 4. New Zealand: Greenstone Valley, near Lake Wakatipu, West Otago, South Island. Beech forest (NorhoJugu.7 sp.) with grassy valley flats. N.Z. Forest Service Photo: By J . H. Johns, A.R.P.S.

Plate 5. Spain: Coto Dohana. Fallow deer on the edge of the marsh. Photo: F. Alvarez.

Plate 6. Sweden: Oland. Fallow deer in broad-leaved, deciduous forest with grass glades. Photo: R. von Schultz.

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Plate 8. California: Mendocino County. White variety of Fallow deer and Mule deer feeding on grassland with woodland beyond and Redwood Valley in the background. Photo: J. M. Jurek.

Plate 9. Kentucky: The Land Between Two Lakes. Fallow buck on the edge of broad-leaved, deciduous woodland, with Cherrybark oak on the left and Fesruca elarior as the predominate grass. Photo: Tennessee Valley Authority.

Plate 10. Texas: Edwards Plateau. Wooded habitat of Fallow Deer. Photo: Texas Parks and Wildlife Department, Wildlife Division.

Plate 11. Argentina. Fallow deer in the Sierra de la Ventana. Photo: E. E. Willard.

The distribution of fallow deer: a worldwide review

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