14
Biological Conservation 58 (1991) 85-98 i Conservation Status of Gazelles in Saudi Arabia C. R. Thouless* Zoological Society of London, Regent's Park, London NW1 4RY, UK J. G. Grainger, M. Shobrak & K. Habibi National Commission for Wildlife Conservation and Development, PO Box 61681, Riyadh 11575, Saudi Arabia (Received 25 January 1990; revised version received 13 February 1991; accepted 22 February 1991) ABSTRACT The populations of the three gazelle species native to Saudi Arabia have been greatly reduced over the last fifty years. The Saudi gazelle Gazella (dorcas) saudiya is almost certainly extinct in the wild. Scattered populations of the mountain gazelle Gazella gazella still occur in the mountains of western Saudi Arabia and in the northern deserts. There is also one substantial population of the subspecies G. g. farasani on the Farasan Islands in the Red Sea. Sand gazelle GazeUa subgutturosa marica are only found in two areas on the fringes of the Nafud desert in the north. As a result of the protected area system being established by the National Commission for Wildlife Conservation and Development, some populations of the two remaining species of gazelle can be expected to survive despite continued hunting pressure and overgrazing over much of the Kingdom, but their future is bleak outside protected areas. INTRODUCTION Gazelles were once abundant in Arabia, but excessive hunting and degradation of rangeland through overgrazing have greatly reduced their numbers, leaving scattered relict populations. In this paper we describe their * Present address: c/o WWF Regional Office, PO Box 62440, Nairobi, Kenya 85 Biol. Conserv. 0006-3207/91/$03"50 © 1991 Elevier Science Publishers Ltd, England. Printed in Great Britain

Conservation status of gazelles in Saudi Arabia

  • Upload
    taif

  • View
    0

  • Download
    0

Embed Size (px)

Citation preview

Biological Conservation 58 (1991) 85-98 • i

Conservation Status of Gazelles in Saudi Arabia

C. R. Thouless*

Zoological Society of London, Regent's Park, London NW1 4RY, UK

J. G. Grainger, M. Shobrak & K. Habibi

National Commission for Wildlife Conservation and Development, PO Box 61681, Riyadh 11575, Saudi Arabia

(Received 25 January 1990; revised version received 13 February 1991; accepted 22 February 1991)

ABSTRACT

The populations of the three gazelle species native to Saudi Arabia have been greatly reduced over the last fifty years. The Saudi gazelle Gazella (dorcas) saudiya is almost certainly extinct in the wild. Scattered populations of the mountain gazelle Gazella gazella still occur in the mountains of western Saudi Arabia and in the northern deserts. There is also one substantial population of the subspecies G. g. farasani on the Farasan Islands in the Red Sea. Sand gazelle GazeUa subgutturosa marica are only found in two areas on the fringes of the Nafud desert in the north. As a result of the protected area system being established by the National Commission for Wildlife Conservation and Development, some populations of the two remaining species of gazelle can be expected to survive despite continued hunting pressure and overgrazing over much of the Kingdom, but their future is bleak outside protected areas.

INTRODUCTION

Gazelles were once abundant in Arabia, but excessive hunting and degradation of rangeland through overgrazing have greatly reduced their numbers, leaving scattered relict populations. In this paper we describe their

* Present address: c/o WWF Regional Office, PO Box 62440, Nairobi, Kenya

85 Biol. Conserv. 0006-3207/91/$03"50 © 1991 Elevier Science Publishers Ltd, England. Printed in Great Britain

86 C. R. Thouless, J. G. Grainger, M. Shobrak, K. Habibi

past and present distribution, the status of the existing populations and the conservation measures necessary for their survival.

Human activities have had an adverse effect on gazelle populations from prehistoric times. Gazelles were a major food source for people in the Middle East during much of the Neolithic period (Legge & Rowley-Conwy, 1987; Edwards, 1989) and were caught in large numbers in traps known as 'desert kites' (Betts, 1984). These were large stone structures with curtain walls, often extending over several kilometres, and an enclosed killing area into which the animals were driven. Desert kites were used until the early years of this century, when gazelle herds numbering tens of thousands were still to be seen in northern Arabia (Mendelssohn, 1974).

The catastrophic decline in wildlife populations in Arabia started soon after the First World War, when modern rifles and motor vehicles came into the country (Meinertzhagen, 1954). By the 1950s there were few gazelles left. According to Thesiger (1959) 'in Saudi Arabia during the last few years even gazelle have become rare. Hunting-parties scour the plains in cars, returning with lorry-loads of gazelles, which they have run down and butchered'. Habibi (1986) considered that two out of three of the native gazelle species were probably extinct, and the third was endangered.

In recent years a concern for wildlife has begun to develop in Saudi Arabia, and in 1986 the National Commission for Wildlife Conservation and Development was established under the direction of HRH Prince Saud al Faisal. Amongst the first tasks of NCWCD were the investigation of the status of endangered wildlife and setting up a series of protected areas for surviving wildlife populations. The results described in this paper come from survey work carried out by the Protected Areas Planning Unit of NCWCD and by members of the King Khalid Wildlife Research Centre and the National Wildlife Research Centre,

Gazelles of Saudi Arabia

There is still uncertainty surrounding the taxonomic status of Arabian gazelles (Groves, 1988). Much of this confusion results from the paucity of museum specimens, which are unrepresentative of the full range of geographical variation within populations. However, there are presently considered to be three species native to Saudi Arabia, which are all ecologically distinct, though their relationship to gazelles outside the country is still unsettled. These are the mountain or common gazelle Gazella gazella, known locally as 'idmi', the sand gazelle or 'rheem' Gazella subgutturosa and the Saudi gazelle or 'afri' Gazella (dorcas) saudiya. Some of our informants believe that there is a fourth type, the 'khudri', but descriptions of this animal differ according to the source.

Gazelles in Saudi Arab& 87

The mountain gazelle is long-legged with a side stripe on the body and pronounced facial markings. Both sexes have horns, but those of females are small and often malformed. Typically females live in small groups while males are solitary and territorial (Mendelssohn, 1974). Two subspecies are known from Saudi Arabia. Gazella gazella cora occurs on the mainland and G. g. farasani (Thouless & Al Bassri, 1991) on the Farasan Islands in the Red Sea. It is not clear where the dividing line between G. g. cora and the Palestinian subspecies G. g. gazella occurs, because there are no museum specimens from the northern half of Saudi Arabia. It is not even certain whether cora is a subspecies of Gazella gazella, and Groves (1989) has suggested that it might be the Arabian representative of Gazella dorcas. The gazelles from the Farasan Islands were previously known as G. arabica, which was described from a single specimen consisting of skull and skin (Groves, 1983). The specimen is a large rufous gazelle with long horns. However, there is considerable doubt surrounding the origins of this animal and the gazelles currently on the island are small and grey, with almost hornless females. They are similar to Gazella gazella and have been described as a distinct subspecies, G. g. farasani (Thouless & A1 Bassri, 1991).

The Saudi gazelle is a small pale red-brown gazelle with indistinct body markings, a black nose spot and knee tufts, short legs, long ears, and very long straight horns in both sexes. It has been considered a subspecies of the north African Gazella dorcas but is now thought by some authorities (e.g. Groves, 1989) to be a separate species. There is virtually no information on its biology.

The sand gazelle is a stocky pale-coloured animal. It is gregarious and lives in open country. Both sexes have horns, those of the females being long and thin. Typical gazelle markings, such as flank and face stripes, are generally obscure but there is some variation in coloration, particularly in the extent of white on the nose and the presence of a nose spot. Sand gazelle can be identified in the field at a considerable distance from their gait. They seldom stot, and, instead of bounding like the mountain gazelle, their typical fast gait consists of a trot with the prominent black tail held stiffly upright, rather than being wagged from side to side.

SURVEY METHODS

Data on historical distribution was collected through a literature search and interviews with old local inhabitants. Much of this information has to be treated with caution, because of the problems of identification. Most locals refer to all gazelles as 'dhubi'. It is generally only old men skilled in hunting

88 C. R. Thouless, J. G. Grainger, M. Shobrak, K. Habibi

who use the words 'rheem', 'afri' and idmi' to distinguish between sand, Saudi and mountain gazelles. A similar problem exists with written records from European travellers. Many did not distinguish between the various species, and the Saudi gazelle was not described until the 1930s (Carruthers & Schwarz, 1935). Even if the species has been identified, the report may have to be treated with caution. For example, a record of mountain gazelles in Jordan close to the Saudi Arabian border (Mountfort, 1961) is more likely to refer to sand gazelle on the basis of group size, habitat and behaviour.

Most of the information on recent distribution in this paper was collected during the course of field surveys conducted by the Protected Areas Planning Unit of the National Commission for Wildlife Conservation and Development, and during surveys for Arabian bustards in the south-west of the country. The main gazelle populations have been systematically surveyed from the air using helicopters and light aircraft, and helicopters were also used for general surveys of areas where gazelle signs had been found on the ground. Gazelles were more likely to be seen from the air than from the ground, since flight distances were typically more than one kilometre. In a number of locations gazelles were not actually seen and their presence was detected from dung and spoor. Species could usually be distinguished by dung piles, which are absent or small in areas frequented by sand gazelles. Mountain gazelle dung piles are conspicuous even in areas of low population density. Surveys also included interviews with local people, to stimulate interest in conservation, and to gather information about past and present distribution of wildlife.

SURVEY RESULTS

Mountain gazelle

The original distribution of the mountain gazelle is uncertain. Museum specimens all come from the western part of the country (Fig. 1), and Major H. St J. Philby (quoted in Carruthers & Schwarz, 1935) reported that their range did not extend beyond the Rakba plain (Fig. 2), while further east they were replaced by the Saudi gazelle. However local informants are insistent that mountain gazelle did occur over the eastern part of the country as well, and it is possible that Philby was only describing their distribution in a local area, since elsewhere (quoted in Morrison-Scott, 1939) he said that they occurred in all suitable country--'valleys of foothills, open rock or sand plains and in true sand areas, but not so plentiful in sand tracts as elsewhere'. According to Vesey-Fitzgerald (1952), the mountain gazelle was still common in the coastal plains and in the highland plains of the interior, and was usually associated with Acacia.

Gazelles in Saudi Arabia 89

35 ° E 4 0 ° 4 5 ° 5 0 ° 5 5 °

3 0 °

2 5 °

2 0 °

1 5 ° N

Fig. 1. Distribution of the mountain gazelle in Saudi Arabia. 0, recent sightings; @, locations where tracks and droppings have been found; O, records from before 1970. Additional records come from Doughty (1888), Carruthers & Schwarz (1935), Morrison- Scott (1939), Philby (1952), Harrison (1968), Anon. (1989), J. Gasparetti (pers. comm.) and J.-

F. Asmod6 (pers, comm.).

There are few gazelles left on the narrow coastal plain of the Tihama. This area was once rich grazing land, partly covered by open acacia woodland. It has, however, become severely degraded by overgrazing in recent years and the high human population places additional pressures on wildlife. There are probably 30-50 gazelles at Makshush, which is a small area of coastal subkha (salt pan), sand dunes and terrace deposits. Gazelles in the area feed on Suaeda sp. and Aeluropus massauensis, neither of which are eaten by domestic stock, and they also use adjoining degraded Acacia tortilis woodlands. Mounta in gazelles have also been reported further nor th in an area of sand dunes with scattered salt bushes. The dunes are soft, making vehicle access difficult.

There are a number of gazelle populations in the foothills of the Asirs, down towards the border with Nor th Yemen. These are mostly associated with acacia-lined wadis, often in the same area as the Arabian bustard Otis arabs. There are some gazelles in the Asir National Park, but there is no effective protection in the park. These populations are hunted and are at low densities, and habitats are generally severely degraded.

The Nor thern Hejaz are mounta in ranges north of Medina, extending up towards the Gulf of Aqaba. They are more arid than the southern mountains

90 C. R. Thouless, J. G. Grainger, M. Shobrak, K. Habibi

35°E 40 45 50 55 60

Fig. 2.

30 Jorda Iraq ~ /~---'~

Ar.:, : ' . ' ::::::::::::::::::::::::::: :.:::::: ':: ':::..:... t ~ Khunfa~:!Y::'i::::::+:::'}:'. :'.. :"'"':ii:lii~i':.. ~"L_ I Bahre ,n

",,.o,he,, i -% /

25 Jebel ! . ] ". '; ":~]::];]::]. .}. Arab K a l l a b ~ " Medina I R,yadh "-i:.ii!ii:.::.:." ' : l ! i i i ~

Jeddah~ Rak ibna Pla I i::ii~ii:i:iiiiiii:i:ii!i::iii " ili:':::Jii::ii: : ] i ~ i i i i l i i i i ~ ii~}~iiiiiiiii!i!i~i~!~ ~ ~ ! i ~ i i • Mecca i :".:::::::::Rub a ":':':

' ~ ~' ===============================================================

M.~hush ~ ' l....~:~:~:~:~:i:~:~:~?~:~:~:i:i:~:i:~:~:~~ ~-,~ ' ,,.. . :i:!i!:!~!~iiiiiii!!!~i]i ~ - i : . : : : " I x ~ ~ .:. ::::::::::::::::: ::::::::::: :.:.-. :: . . . . . . . . . •

• North ~ , ~ , ~ (

Arabian peninsula showing major geographical features. Stippled area shows major sand deserts.

and there is little permanent water. Gazelle populations have been found in four sites in the Northern Hejaz. In three of these locations only sparse signs of gazelle have been found, but on Jebel Kallab ibex and mountain gazelles were seen in good numbers. It seems that this population has been protected by the extreme difficulty of access which involves walking and scrambling through steep ravines to reach the gentler upper slopes and wide wadis favoured by the gazelles. There is no possibility of vehicular access, and although the area is used by camels, goats and feral donkeys, the vegetation is in much better condition than in surrounding areas. There are probably about 50 gazelles on Jebel Kallab.

There are extremely small populations of mountain gazelle on the periphery of the deserts of northern Saudi Arabia in the NCWCD reserves of A1 Khunfah and A1 Harrah. In both areas mountain gazelle occur sympatrically with sand gazelle and have been observed in mixed groups, though mountain gazelle are more frequently seen in the rockier areas. The presence of mountain gazelle here is surprising, since there are no trees and no permanent water.

The subspecies G. g. farasani is confined to the Farasan island group, a reefal limestone archipelago lying some 50 km away from the mainland of

Gazelles in Saudi Arabia 91

Arabia. Gazelles occur on three of the islands (Flamand et al., 1988). The largest number are on the main island, Farasan Kebir, particularly in the north-eastern part, which consists of well-vegetated ravine country inaccessible to vehicles and with difficult access on foot. In this area gazelle density is as high as 10 animals/km 2. There is also a thin scattering of gazelles over the rest of the island, which is more open. The second largest island, Segid, is flat and open, and only has small numbers of gazelles. To the south of Farasan Kebir lies Zifaf, which is uninhabited, and also has good gazelle numbers. Although hunting has taken place in the past and captive gazelles have been taken to the mainland, there is little evidence for a recent population decline, and aerial surveys indicate that there are at least 500 animals present in the Farasan archipelago.

The Farasan Island gazelle population is under no immediate threat, although planned developments for the islands are potentially disastrous. The archipelago has recently been declared a reserve by NCWCD, and rangers are stationed there. If hunting is completely prevented, there is a danger that with low and variable rainfall, and a high gazelle density, a population crash may occur as a result of the gazelles eating out their food resources in a poor year. It is possible that losses from hunting have acted as a brake on population fluctuations, and it is important for a high level of scientific monitoring to be implemented at the same time as effective protection.

Saudi gazelle

The Saudi gazelle was once common on gravel and sandy plains, especially in the north (Morrison-Scott, 1939; Lewis et al., 1965) and in the south-west (Fig. 3). According to Vesey-Fitzgerald (1952) it was confined to the interior of the peninsula where acacias grew. Saudi gazelles ranged widely over the great gravel plains lying to the east of the Hejaz mountains.

There have been no recent records of the Saudi gazelle. Occasional sightings have been reported by bedouin, but none of these have been confirmed.

Sand gazelle

The sand gazelle was once abundant in the open country of central Arabia and was the only species found on the northern gravel plains and around the central Arabian sands (Vesey-Fitzgerald, 1952). The majority of historical records are from the north of the country, mostly from gravel plains and black lava deserts to the north and west of the Nafud desert (Fig. 4). According to Philby (1923) sand gazelle were present in 'thousands', and

92 C. R. Thouless, J. G. Grainger, M. Shobrak, K. Habibi

3 0 °

2 5 °

2 0 °

1 5 ° N

3 5 ° E 4 0 ° 4 5 ° 5 0 ° 5 5 °

Fig. 3. Historical distribution of Saudi gazelle. O, specimens; O, sightings or other reports. Geographical location is not precise for these records. Information from Carruthers & Schwarz (1935), Morrison-Scott (1939), Lewis et al. (1965), Harrison (1968) and Abu

Mohamed al 'Ajma (pers. comm.).

Vesey-Fitzgerald (1952) saw them in flocks of 50-100. The presence of herds of this size suggests that they may have been migratory--fol lowing a fixed route each yea r - -o r nomadic. Philby (1923) and Raswan (1935) described mass movements of sand gazelles away from areas of drought. Migrating herds were observed in nearby Jordan in the early years of this century (Mendelssohn, 1974) and Legge and Rowley-Conwy (1987) have suggested that from the Neolithic period to recent times there was a regular migration from this area up into present-day Syria. However, bones excavated in Syria are of the subspecies subgut turosa , rather than the marica which exist in northern Saudi Arabia today.

There are only two areas in Saudi Arabia where sand gazelle are still known to be resident. One is the N C W C D reserve ofAl Harrah, close to the Jordanian border. This is an area of just over 12 000 km 2 of basalt plains and hills, which has been protected from grazing by sheep since the end of 1986. The density of sand gazelle is extremely low and during the course of an approximately 15% aerial coverage at 154m in 1989 only four animals were seen. They have been reported from several parts of the reserve, but never in large groups.

AI Khunfah Reserve lies to the south of A1 Harrah. Only a small part of it,

Gazelles in Saudi Arabia 93

35 ° E 4 0 ° 4 5 ° 5 0 ° 5 5 °

3 0 °

2 5 °

2 0 °

15ON

Fig. 4. Distribution of sand gazelle in Saudi Arabia. O, sightings since 1970; C), earlier reports. Additional information comes from Philby (1933), Thesiger (1948, 1949), Vesey- Fitzgerald (1952), Harrison (1968), Green (1986), J. Gasparetti (pers. comm.), G. Popov (pers.

comm.), J.-F. Asmod~ (pers. comm.) and NCWCD pilots and rangers.

Ghurrub, occupying approximately 2000km 2, is effectively protected. However, it probably contains the majority of sand gazelle within the Kingdom. Aerial surveys indicated that the total population was about 250 animals. Their habitat is largely gravel plains and low hills, with no trees or large shrubs. The dominant vegetation consists of chenopods of the genera Salsola and Suaeda with some Stipagrostis sp. grasses in the smaller wadis and runnels. After rains the gazelles feed on ephemeral plants but otherwise rely on dry grasses, chenopods and the bitter desert melons Citrullus colocynthis. Small numbers of sand gazelle have been recorded from adjacent areas, but those reported by Green (1986) near Sakaka and in the Nafud desert have almost certainly been exterminated.

Hunting within the reserves has been completely prevented by frequent ground and aerial patrols. The number of sand gazelle can be expected to increase fast, as the vegetation regenerates, since even in A1 Khunfah the present density is less than one animal/10km 2 and, with the ability to produce twins, they have a high intrinsic rate of increase. Two months after the 1989 lambing season there was a female to young ratio of roughly 1:1. The major problem facing these two populations is the effect that free- ranging camels have on vegetation regeneration. Although sheep grazing

94 C. R. Thouless, J. G. Grainger, M. Shobrak, K. Habibi

has been eliminated, the number of camels is well above the carrying capacity of the area.

The Rub' al Khali (the Empty Quarter) is the huge sand desert in the south of Saudi Arabia. Unlike the Arabian oryx, which was forced to leave the sands for the surrounding gravel plains in the heat of the summer in order to find shade (Grimwood, 1962), sand gazelle were apparently permanent residents of the dune country, digging scrapes under Calligonum comosum bushes for shade. Neither Thesiger (1959) or Philby (1933) saw them during their crossings of the Rub' al Khali, but it is likely that t h ~ were nomadic, moving to areas of fresh grazing after localised rain showers. During Cheeseman's (1926) travels in the north-eastern Rub' al Khali he saw no gazelles but remarked that the lack of rain for two years had driven them to other areas, and Thomas (1931) thought that the sand gazelle had virtually disappeared from the whole Rub' al Khali.

The Rub' al Khali has been subjected to huge numbers of hunting expeditions over the last twenty years and Mandaville (1986) believed that by the early 1970s the gazelles once present in the sands had been virtually exterminated by motorised hunting. Participants in these expeditions shoot at anything that moves, and we have heard of very large bags of sand gazelle obtained with the aid of automatic weapons. Although the Rub' al Khali is a very hostile environment, skilful drivers can cross most of the dunes in light-weight four-wheel-drive vehicles during the winter, when the sand is firmer than in the summer. We have seen hares driven to exhaustion in steep dune areas before being shot, and there is little doubt that the same would happen to any gazelles found. During the course of a traverse of the central sands in November-December 1988 and a biological survey of the eastern Rub' al Khali in February-March 1990, we found no evidence for the continued survival of sand gazelle. Bedouins resident in the area told us that the only gazelles still to be seen were ones which had wandered across the border from Oman.

DISCUSSION

Future prospects

A few years ago the prospects for the continued survival of any gazelle populations in Saudi Arabia looked bleak (Habibi, 1986), but with the establishment of the National Commission for Wildlife Conservation and Development the situation has improved considerably. Small but viable populations of both sand and mountain gazelle exist in protected areas. However, most of the populations outside these protected areas are likely to go extinct unless action is taken soon. Some of the areas concerned are

Gazelles in Saudi Arabia 95

suitable for protected sites and should be given priority. Makshush is a prime example, since it is a limited area with clear boundaries and preservation would create few conflicts with other interests. But in many places the gazelles are at a low density, and in areas intensively used by people. For these to survive, and for gazelles to re-establish themselves in other areas outside reserves, major changes are needed.

Hunting is still a serious threat in most areas. There are game laws, according to which gazelles are fully protected, but there is no enforcement. It is essential for government agencies such as the police and Frontier Force to be made aware of the laws and to assist with enforcement. But since the country is so large compared to the human population it will never be easy to enforce hunting laws. What is needed is for people's attitudes to wildlife and natural resources to change. NCWCD has engaged in a widespread publicity campaign and is working towards the creation of a new hunting ethic, which will emphasize traditional hunting skills, as opposed to the mechanised slaughter that has generally taken place.

However, hunting is only part of the problem. More serious in the long term is the destruction of rangeland through overgrazing. In the past this was prevented by tribal-based systems of resource protection and by the difficulties of providing water for stock in remote areas. The abolition of tribal areas in the 1950s, the coming of water tankers and livestock subsidies (initially paid for each animal in a flock and more recently for supplementary food), mean that it is now possible for stocking rates to be much higher than the range can support and for people to go anywhere in the country with their animals. Many parts of the Kingdom are now almost completely barren, with perennial grasses and palatable shrubs all gone, and acacias being lost. There has been little regeneration of acacias for many years but also now mature trees are being grubbed out, a practice which has been traditionally banned.

Although gazelles can take some food that domestic stock will not touch, they are at a competitive disadvantage since domestic stock do not have to rely on natural vegetation all year round. In order for gazelles and other wildlife to flourish and for better use to be made of the country's natural resources, drastic reductions will have to be made in numbers of free-ranging stock. Plans are underway to reintroduce sand and mountain gazelles using captive stocks in NCWCD's captive breeding centres, but these are likely only to succeed in areas where grazing has been controlled.

International significance

Some Saudi Arabian gazelle populations are of international importance. The Saudi gazelle has the highest priority for conservation, since it is

96 C. R. Thouless, J. G. Grainger, M. Shobrak, K. Habibi

endemic to the Arabian peninsula. However, it is almost certainly extinct in the wild. According to Gross (1987) there is a large herd on an island offthe coast of the northern Emirates. However, the identification and origin of these animals is very uncertain. The only captive population known to exist is that at A1 Areen Park in Bahrein, which consisted of 25 animals in 1989. Other captive populations have been reported from A1 Ain in the United Arab Emirates and A1 Wabbra in Qatar, but these do not closely resemble museum specimens of Gazella saudiya.

The species Gazella subgutturosa is widespread, occurring from Arabia through the deserts of east and central Asia as far as China, but it has disappeared from much of its range. The subspecies G. s. marica is confined to the Arabian peninsula. In addition to Saudi Arabia it is found in small numbers in Oman and in coastal areas and offshore Arabian Gulf islands including Bahrein. However, the purity of Gulf populations is uncertain since many appear to have been mixed with released G. s. subgutturosa. The population in A1 Khunfah is thus of major international importance.

Although the mountain gazelle has been exterminated from most of its range in the Middle East, the subspecies Gazella gazeHa gazella is common in Israel, and G. g. cora still occurs in thousands in Oman. The value of the Saudi animals is chiefly genetic, since so little is known about natural variation in the species.

A C K N O W L E D G E M E N T S

The authors wish to thank Dr Abdulaziz H. Abuzinada and the National Commission for Wildlife Conservation and Development for sponsoring the work on which this paper was based. We would also like to thank everyone who has given us information on gazelle distribution and helped with fieldwork, especially Jean-Francois Asmod6, Graham Child, John Gasparetti, Mohamed Salamah, Hany Tatwany and Youssef Wutaid.

REFERENCES

Anon. (1989). Conservation of natural resources in the Kingdom of Saudi Arabia. Proceedings of the Symposium on Wildlife Conservation and Development in Saudi Arabia. National Commission for Wildlife Conservation and Develop- ment, Saudi Arabia, pp. 10-17.

Betts, A. (1984). Black Desert survey, Jordan: second preliminary report. Levant, 16, 25-34.

Carruthers, D. & Schwarz, E. (1935). On a new gazelle from central Arabia. Proc. Zool. Soc. Lond., 1935, 155-6.

Gazelles in Saudi Arabia 97

Cheeseman, R. E. (1926). In Unknown Arabia. Macmillan, London. Doughty, C. M. (1888). Travels in Arabia Deserta, 2 vols. 1979 edition. Dover,

New York (first published 1888 Cambridge University Press, Cambridge). Edwards, P. C. (1989). Revising the Broad Spectrum Revolution: and its role in the

origins in Southwest Asian food production. Antiquity, 63, 225~,6. Flamand, J. R. B., Thouless, C. R., Tatwany, H. & Asmod~, J.-F. (1988). Status of

the gazelles of the Farasan Islands, Saudi Arabia. Mammalia, 52, 608-10. Green, A. (1986). Status of large mammals of northern Saudi Arabia. Mammalia, 50,

483-93. Grimwood, I. R. (1962). 'Operation Oryx'. Oryx, 6, 308-34. Gross, C. (1987). Mammals of the Southern Gulf Motivate Publishing, Dubai. Groves, C. P. (1983). Notes on the gazelles, IV. The Arabian gazelles collected by

Hemprich and Ehrenberg. Z. Saugetierk., 48, 371-81. Groves, C. P. (1988). A catalogue of the genus Gazella. In Conservation and Biology

of Desert Antelopes, ed. A. Dixon & D. Jones. Christopher Helm, London, pp. 193-8.

Groves, C. P. (1989). The gazelles of the Arabian Peninsula. Proceedings of the Symposium on Wildlife Conservation and Development in Saudi Arabia. National Commission for Wildlife Conservation and Development. Saudi Arabia, pp. 23748.

Habibi, K. (1986). Arabian ungulates--their status and future protection. Oryx, 20, 100-3.

Harrison, D. L. (1968). Mammals in Arabia, Vol. II. Ernest Benn, London. Legge, A. J. & Rowley-Conwy, P. A. (1987). Gazelle killing in Stone Age Syria.

Scient. Amer., 257, 88-95. Lewis, R. E., Lewis, J. H. & Harrison, D. H. (1965). On a collection of mammals

from northern Saudi Arabia. Proc. Zool. Soc. Lond., 144, 61-74. Mandaville, J. P. (1986). Plant life in the Rub'-al-Khali (the Empty Quarter), south-

central Arabia. Proc. R. Soc. Edinb., 89B, 147-57. Meinertzhagen, R. (1954). Birds of Arabia. Oliver & Boyd, London. Mendelssohn, H. (1974). The development of the populations of gazelles in Israel

and their behavioral adaptations. In The Behavior of Ungulates and its Relationship to Management, ed. V. Geist & F. Walther. IUCN Pubs, no. 24, 722-43.

Morrison-Scott, T. C. S. (1939). Some Arabian mammals collected by Mr H. St J. B. Philby. Novit. Zool., 41, 181-211.

Mountfort, G. (1964). Disappearing wildlife and growing deserts in Jordan. Oryx, 7, 229-32.

Philby, H. St J. B. (1923). Jauf and the north Arabian desert. Geogr. J., 122, 242-59.

Philby, H. St J.B. (1933). The Empty Quarter. Constable, London (paperback edition 1986 Century, London).

Philby, H. St J. B. (1952). Arabian Highlands. Cornell University Press, Ithaca, New York.

Raswan, C. R. (1935). The Black Tents of Arabia. Hutchinson, London. Thesiger, W. (1948). Across the Empty Quarter, Geogr. J., I l l , 1-21. Thesiger, W. (1949). A further journey across the Empty Quarter. Geogr. J., 113,

21~,6. Thesiger, W. (1959). Arabian Sands. Longmans, London.

98 C. R. Thouless, J. G. Grainger, M. Shobrak, K. Habibi

Thomas, B. S. (1931). A journey into Rub' al Khali. Geogr. J., 77, 1-37. Thouless, C. R. & Al Bassri, K. (1991). Taxonomic status of the Farasan Island

gazelle. J. Zool., Lond., 151-9. Vesey-Fitzgerald, D. F. (1952). Wildlife in Arabia. Oryx, l, 232-5.