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Although a Robertsonian translocation and autosome-to-X chromosome translocation are known to occur naturally in C. s. marica, the question of whether or not these chromosomal rearrangements cause fertility problems is relevant to the captive management of this gazelle. Translocation heterozygosity is known to reduce fertility in certain mammals, notably humans, mice, and cattle (reviewed by Dyrendahl and Gustavsson, 1979; De Boer and De Jong, J989; Speed, 1989). For most species, however, there are too few empirical data, particularly from wild populations, to determine the effects of chromosomal rearrangements on reproduction. Chromosomal pairing and segregation studies during meiosis can help to determine the effects of chromosomal heterozygosity on fertility because homologous chromosomes must pair and then segregate for nonnal gametogenesis to continue. Rearranged chromosomes may disrupt homology to the point that gametogenesis is also disrupted. Studies of gametogenesis in six male C. s. marica (three with 2n = 31, three with 2n = 32) indicated nonnal chromosomal pairing and segregation, suggesting that heterozygotes have nonnal fertility (Kingswood, 1992). Evidence of nonnal fertility in C. s. marica heterozygotes is consistent with the prevalence of Robertsonian and sex-chromosome translocations in gazelles (Kings wood and Kumamoto, 1994). This suggests that the translocations are karyotypically adaptive; an example of what White (1973) tenned "karyotypic orthoselection". Whether or not chromosomal rearrangements in gazelles have selective value to their karyotypes is a question that can only be addressed theoretically. However, the fact that observed frequencies of the three cytotypes in C. s. marica (Table 10.1) do not differ from Hardy-Weinberg expectations indicates that there is an absence of selection for anyone of the three karyotypes. This review of the systematics of C. subgutturosa has demonstrated the unique biological characteristics of this species. Recognition that C. s. marica is morphologically and genetically distinct from C. s. subgutturosa reinforces the need for conservation efforts directed specifically towards the preservation of C. s. marica. Continued protection of wild populations is extremely important. The importance of existing captive populations must also be recognized as they may represent diversity that has since been lost in the wild through population declines. Based on this review of the systematics of C. subgutturosa, the following management and research recommendations are suggested for the conservation of C. s. marica: 1) Continue to protect and maintain existing wild populations. 2) Preserve additional genetic diversity by maintaining the existing captive populations in Saudi Arabia and the United States. Maintaining these captive populations serves to protect against the possibility of continued declines in wild populations. Ideally, captive populations should be genetically pure and represented by founders of known geographic origin. The U. S. population, of eastern Jordanian origin, is highly inbred, and the introduction of new founder stock from northern Saudi Arabia is recommended. Chromosomal translocations found in both captive populations are believed to be of natural origin and do not appear to confer a reproductive disadvantage to heterozygotes. Because observed frequencies of the three cytotypes indicate that there is an absence of selection for allY one of the three karyotypes, all three karyotypic fonns should continue to be maintained through random mating. However, it would be useful to continue monitoring the inheritance of the translocation through cytogenetic and pedigree analyses. 109
3) Establish the genetic characteristics of wild populations of G. s. marica and G. s. subgutturosa, particularly of those reported in the Rub al Khali (Saudi Arabia), Iran, and the zone of intergradation in Iraq, through continued chromosomal banding and molecular studies. Identification of biochemical characteristics unique to either subspecies is important in order to screen captive G. s. marica for evidence of introgression of G. s. subgutturosa. 4) In keeping with the draft guidelines of the IUCN I SSC Re-introduction Specialist Group (Anon., 1992), individuals selected for reintroduction should be of the same taxonomic unit and, ideally, closely related genetically to those which were extirpated. All three cytotypes of G. s. marica should be considered suitable for reintroduction projects in Saudi Arabia. Allimals selected for reintroduction projects aimed at restoring populations of G. s. marica should ideally be free of G. s. subgutturosa introgression, as determined through biochemical studies. Acknowledgements Arlene Kumamoto, Suellen Charter, Marlys Houck, Bill Flavell, and Osama Badri have been actively involved in cytogenetic studies of the Arabian sand gazelles and deserve special thanks. Arnaud Greth, Colin Groves, Eric Harley, Marlys Houck, and Arlene Kumamoto gave helpful suggestions to improve the manuscript. The ongoing support of the Zoological Society of San Diego for antelope cytogenetics research and the commitment by the National Commission for Wildlife Conservation and Development of Saudi Arabia to gazelle conservation is gratefully acknowledged. References Anon. 1992. Draft guidelines for re-introductions. Re-introduction News, 4: 2-3. Baccus, R., Ryman, N., Smith, M.H., Reuterwall , C., and Cameron, D. 1983. Genetic variability and differentiation of large grazing mammals. Journal ofMammalogy, 64: 109-120. Benirschke, K., and Kumamoto, AT. 1987. Challenges of artiodactyl cytogenetics. La Kromosomo, II-45: 1468-1478. Carter, S. 1991. Goitred gazelle North American regional studbook. Sedgwick County Zoo and Botanical Garden, Wichita, Kansas, 87 pp. De Boer, P., and De Jong, I.H. 1989. Chromosome pairing and fertility in mice. In Gillies, C.B. (ed), Fertility and chromosome pairing: recent studies in plants and animals. CRC Press, Boca Raton, Florida. Pp. 37-76. Durand, L., Durand, J., and Catzeflis, F.M. 1989. Biochemical systematics of endangered Arabian Gazella (Artiodactyla: Bovidae). 5th International Theriological Congress, Poster, Rome, Italy. Abstract. Dyrendahl, I., and Gustavsson, I. 1979. Sexual functions, semen characteristics and fertility of bulls carrying the 1129 chromosome translocation. Hereditas, 90: 281-289. East, R. 1992. Conservation status of antelopes in Asia and the Middle East, part I. Species, 19: 23 25. Effron, M. , Bogart, M.H., Kumamoto, A.T., and Benirschke, K. 1976. Chromosome studies in the mammalian subfamily Antilopinae. Genetica, 46: 419-444. Ellennan, J.R., and Morri son-Scott, T.C.S. 1951. Checklist of Palearctic and Indian mammals, 1758 to 1946. Trustees of the British Museum, London, 810 pp. Granjon, L., Vassart, M., Greth, A, and Cribiu, E.P. 1991. Genetic study of sand gazelles (Gazella subgutturosa marica) from Saudi Arabia. Chromosomal and isozymic data. Zeitschrift for Siiugetierkunde, 56: 169-176. 110
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CONSERVATION OF ARABIAN GAZELLES
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• 14 Survey of Gazelle Populatio
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List of Contributors Prof. Abdulaz
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Foreword HRH Prince SAUD AL-FAISAL
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1. Recent Developments in Gazelle C
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• April, 1988: Survey of the Fara
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- May, 1991: Capture of rheem from
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References Abuzinada, A., 1987. Int
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until Groves (1983) re-examined the
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Figure 2.1 a Figure 2.1 b
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Dissatisfaction with the BSC over t
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Nixon & Wheeler (1990) are not so d
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In order to test the PSC (phylogene
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(2) FEMALES Variable gazella cora
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two adult male and one adult female
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4 4 * 4 3 2 *33 1 1 * 1 * 2 1 1 I 5
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(2) FEMALES Variable G. bennelli (
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In the two other equally parsimonio
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ufifrons benneui saudiya subguuurcs
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By branch-swapping, using Macelade,
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On my first visit to AI-Areen, in 1
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anch of the premaxilla generally to
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J.R., and Morrison-Scott, T.C.S. 19
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Aden: 12°50'N, 45°03'E (4 skins,
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species is currently the subject of
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One area where local people are ins
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4. The Relationship Between Taxonom
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'I1Ie current status of taxonomy Ad
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There is much scope for conflict be
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The Controversial Subject of Gazell
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• Taxonomic argument: The divisio
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conventional taxonomic characters t
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divergence between the mitochondria
Page 68 and 69: Morphometries: Some morphological c
Page 70 and 71: operative progranune could be set u
Page 72 and 73: museum specimens (data extracted fr
Page 74 and 75: Acknowledgements This work was carr
Page 76 and 77: Lange, 1. 1972. Studien an Gazellen
Page 78 and 79: 6. Modern Techniques in Taxonomy an
Page 80 and 81: protein electrophoresis still a val
Page 82 and 83: "molecular clock") has been establi
Page 84 and 85: Beni~hke, Nombela, J.J., Rodriguez
Page 86 and 87: 7. Chromosonlal Evolution in Gazell
Page 88 and 89: and C. gazella (Vassart et ai., Art
Page 90 and 91: analysis. Systematic relationships
Page 92 and 93: management efforts, whether managem
Page 94 and 95: Kumamoto, A.T., and Bogart, M.H. 19
Page 96 and 97: Table 8.1 Mammalian cells in cultur
Page 98 and 99: genetic diversity in a subspecies w
Page 100 and 101: amount of change between individual
Page 102 and 103: pardus, at positions 38, 61, 64, an
Page 104 and 105: Wildlife Research Center (KKWRC), n
Page 106 and 107: grown m MEM medium supplemented wit
Page 108 and 109: was involved in a V-autosome transl
Page 110 and 111: and heterozygosi ty values found in
Page 112 and 113: Viegas-Pequignot, E., and Dutrillau
Page 114 and 115: cellular and molecular biology requ
Page 116 and 117: structures in terms of visible morp
Page 120 and 121: Groves, c.P. 1969. On the smaller g
Page 122 and 123: G. bilkis and the dark forms of G.
Page 124 and 125: survey was carried out in February
Page 126 and 127: Conservation action: International
Page 128 and 129: 12. Genetics of Saudi dorcas gazell
Page 130 and 131: Genetic research can clarify the ta
Page 132 and 133: dorcas gazelles Gazella do rcas ssp
Page 134 and 135: y less than 2% from each other. How
Page 136 and 137: Groves, c.P. 1969. On the smaller g
Page 138 and 139: eflect the truly remarkable output
Page 140 and 141: • National (central) government p
Page 142 and 143: past colonial Africa and Asia, and
Page 144 and 145: • Studies of gazelle growth rates
Page 146 and 147: References Child, G., and Grainger,
Page 148 and 149: 14. Survey of Gazelle Populations i
Page 150 and 151: Sony IPS-360 Global Positioning Sys
Page 152 and 153: 1. Mahkshush The gazelle habitat co
Page 154 and 155: 5. Jabal Jandaf A small population
Page 156 and 157: A slight caveat is required to thes
Page 158 and 159: • Appoint one local auxiliary ran
Page 160 and 161: Recommendations on park zoning, man
Page 162 and 163: '9~ ~~~
Page 164 and 165: flamingos Phoenicopterus ruber; her
Page 166 and 167: salt marshes and associated sand du
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Many projects were initiated at KlS
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CG: Taxonomy is the essential backg
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WR: On a mitochondrial DNA basis, w
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CM & AG: We should not forget in si
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- AG: What could the taxon called G
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Gazella dorcas 1) Survey Al-Khunfah
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3 Male mountain gazelle (Thumamah)
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7 Female Arabian sand gazelle (Thum
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