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6. Modern Techniques in Taxonomy and their Reliability Eric H. Harley and Marc Vassart Abstract: Cytogenetics is still a useful 1001 for taxonomy especially when modern techniques are used (single or double thymidine synchronization for example) 10 obtain high resolution banded kJJryotypes. In the present communication. examples of the recognition of species and subspecies by cytogenetic means and protein electrophoresis are given, and techniques which were in use at the National Wildlife Research Center (NWRC) in Taif, Saudi Arabia, are described. In recent years biochemical methods of DNA analysis have developed a degree of versatiliry and sophistication that enables their use in a whole range of new applications. One of the most rapidly developing of these is in addressing problems of relationships between organisms, and the versatility of these techniques is demonstrated by their use at levels ranging from differentiation between subspecies to interrelationships between phyla. Characteristics of the types of DNA used, an ourline ofmethods, and their reliability, will be discussed. Keywords: cytogenetics, protein electrophoresis, DNA analysis, taxonomy. Introduction Problems of relationships between species are not only of great concern for taxonomy but also from the point of view of conservation. Conventional cytogenetic techniques still offer a great deal of information to scientists, since the chromosomal constitutions of natural populations of wild mammals are poorly known. However. modern techniques based, for example, on single or double thymidine synchronization appear to be a more powerful tool than conventional ones and are greatly informative for taxonomy and conservation. Other methods involving deoxyribonucleic acid (DNA) analysis also ftnd a whole range of applications in addressing problems of taxonomy . Different methods and their possible applications are presented in this article. Is cytogenetics still a useful tool for taxonomy Cytogenetic techniques describe the number and structure of chromosomes. This was the aim of classic animal cytogenetics. but today, cytogeneticians also use the modern tools of molecular biology to locate precisely the position of genes on chromosomes. Fluorescence in situ hybridization is now the most used tool for this task. Cytogenetic laboratories around the world mainly receive funds to work on domestic animals and are studying the gene maps of cattle and pigs. Cytogenetic techniques such as resolution banding methods are also available. and can be used to describe more precisely the structures of chromosomes. It is now possible to obtain very long chromosomes by the II5e of thymidine synchronization on fibroblast cell culture or the use of actinomycin. For example. a conventionally banded karyotype of the Corsican deer Cervus elaphus corsicanus distinguished a maximum of 400 bands compared to 600-900 bands with high resolution techniques (Rubini et al., 1991). Precise descriptions of chromosomes promises the recognition of a higher degree of chromosomal differentiation between closely-related species or even subspecies. Such results have 69
een obtained with Gazella gazella species. Among Bovids, centric fusions have been the predominate chromosomal rearrangements detected by conventional techniques. By examination of high-resolution banded chromosomes, complex re-shapings of the chromosomes, such as pericentric or paracentric inversions have been found in goat, cattle and sheep (Hayes et al., 1991). Cytogenetics and the recognition of species - Bovids: It was formerly believed that one karyotype was a characteristic of a particular species. However, with increases in sample sizes, chromosomal polymorph isms have been frequently found, e.g. in Madoqua guentheri and M. kirki (two dik-dik species) (Ryder et ai., 1989); Arabian oryx, Oryx leucoryx, (Cribiu et ai. , 1990; Cribiu et al., 1991; Vassart et al., 1991); waterbuck, Kobus ellipsiprymnus (Ryder et ai., 1990); goral, Naemorhedus goral, Sika deer, Cervus nippon, Bactrian wapiti, Cervus elaphus bactrianus, and tufted deer, Elaphodus cephalophus (Shi, 1987); impala, Aepyceros melampus; and blackbuck, Anti/ope cervicapra (Effron et ai., 1976). Benirschke et al. (1984) found 12 different karyotypes in Soemmerring's gazelle Gazella soemmerringi because of different centric fusions and Arryo Nombela et ai. (1990) have described a centric fusion in Gazella dama. Centric fusion leading to chromosomal polymorphism has also been found in G. subgutturosa (Kings wood and Kumamoto, 1988; Granjon et ai., 1991; Vassart et al., 1993) . To be a valid criterion for the recognition of a new species the chromosomal difference between two forms has to be large enough to lead to their reproductive isolation. - Rodents: Conventional techniques are still very useful for recognizing different species inside one genus, as for example, in the case of Mastomys from Senegal (Duplantier et al., 1990). Specific assignations can be attributed partly on the basis of chromosomal characteristics: Mastomys erythroleucus (diploid chromosome number 2n = 38; chromosomal arm number or fundamental number FN =52), M. huberti (2n =32, FN =44) and M. natalensis (2n =32, FN =54). Each of these karyomorphs has its own ecological characteristics; the first is a generalist species, the second is restricted to humid biotopes and the last is commensal. Similarly Tranier et al. (1973) found that Taterillus congicus (Gerbil1idae) was specifically different from T. lacustris, the first having 54 chromosomes and the second 28 chromosomes. Cytogenetics has also been used to recognize sibling species in the genus Hylomyscus (Iskandar et al., 1988). Cytogenetics and the recognition of subspecies One example of the use of cytogenetics to distinguish subspecies comes from a study of the black spider monkey, Ateles paniscus. Two subspecies, A. p. paniscus and A. p. chamek, can be distinguished by their chromosome number: 2n = 32 in the former and 2n = 34 in the latter (De Boer and de Bruijn, 1990). The difference is most probably the result of a tandem fusion between chromosomes 4 and 13 of the original Ateles karyotype (2n = 34) to form a unique metacentric chromosome in A. p. paniscus. Even intra-subspecific karyological variation in A. p. chamek was detected. Cytogenetics has also been used to differentiate nine different subspecies from one single species of night monkey Aotus sp., previously described from its phenotype as a single species (Hershkovitz, 1983). 70
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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
Page 28 and 29: (2) FEMALES Variable gazella cora
Page 30 and 31: two adult male and one adult female
Page 32 and 33: 4 4 * 4 3 2 *33 1 1 * 1 * 2 1 1 I 5
Page 34 and 35: (2) FEMALES Variable G. bennelli (
Page 36 and 37: In the two other equally parsimonio
Page 38 and 39: ufifrons benneui saudiya subguuurcs
Page 40 and 41: By branch-swapping, using Macelade,
Page 42 and 43: On my first visit to AI-Areen, in 1
Page 44 and 45: anch of the premaxilla generally to
Page 46 and 47: J.R., and Morrison-Scott, T.C.S. 19
Page 48 and 49: Aden: 12°50'N, 45°03'E (4 skins,
Page 50 and 51: species is currently the subject of
Page 52 and 53: One area where local people are ins
Page 54 and 55: 4. The Relationship Between Taxonom
Page 56 and 57: 'I1Ie current status of taxonomy Ad
Page 58 and 59: There is much scope for conflict be
Page 60 and 61: The Controversial Subject of Gazell
Page 62 and 63: • Taxonomic argument: The divisio
Page 64 and 65: conventional taxonomic characters t
Page 66 and 67: 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 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 118 and 119: Although a Robertsonian translocati
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
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12. Genetics of Saudi dorcas gazell
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Genetic research can clarify the ta
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dorcas gazelles Gazella do rcas ssp
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y less than 2% from each other. How
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Groves, c.P. 1969. On the smaller g
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eflect the truly remarkable output
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• National (central) government p
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past colonial Africa and Asia, and
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• Studies of gazelle growth rates
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References Child, G., and Grainger,
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14. Survey of Gazelle Populations i
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Sony IPS-360 Global Positioning Sys
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1. Mahkshush The gazelle habitat co
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5. Jabal Jandaf A small population
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A slight caveat is required to thes
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• Appoint one local auxiliary ran
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Recommendations on park zoning, man
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'9~ ~~~
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flamingos Phoenicopterus ruber; her
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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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