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amount of change between individual species, represented by the proportion of shared sites, to be used not only to construct dendrograms, but to provide an indication as to when particular radiations took place. For example. restriction mapping of mitochondrial DNA from both black and white rhinoceros has helped provide an estimate of their time of divergence from a common ancestor to compare with fo ss il evidence (O'Ryan and Harley, 1993). An illustration of the use of restriction mapping in sys tematics is shown in Figure 8.2. Dendrograms were constructed from mitochondrial DNA restriction maps for a number of individual bovid species taken from four tribes, Tragelaphini, Antilopini, Connochaetini and Alcelaphini, analyzed by a distance method and by maximum parsimony. In the absence of a suitable outgroup the maximum parsimony tree was rooted using information from the distance analysis. The only difference between the topologies produced by these two very different tree-building methods is in the positions of the Tragelaphine antelopes relative to each other, and the exchange of Red Hartebeest Alcelaphus caama and Tsessebe Damaliscus luna/us. It is noteworthy that in general the topologies are consistent with current classifications. Cladograms are generally more concerned with the branching order than with branch lengths, although the number of shared derived characters supponing each node are frequently indicated, as here, and can provide an indication of relative branch length. Other statistical approaches, such as the consistency index, which here = 0.55, and the bootstrap (Felsenstein, 1985), can be used to assess the robustness or otherwise of the overall topology or of pa!1icular monophyletic groupings. The distance tree is particularly concerned with relative branch lengths, from which its estimate of the topology is derived, and if a calibration is available, can give an estimate of when lineages divided. A detailed critique of the relative merits of the two methods is beyond the scope of thi s article, but is the subject of much controversy (sometimes acrimonious). The major weakness of the cladistic approach for molecular data is the amount of unrecognized homoplasy in the data, and the major weakness of the distance approach is the degree of stochastic error when measuring small numbers of random events. A similar study using the restriction mapping method was performed on mitochondrial DNA prepared from twelve members of the Odontoceti (toothed whales). Distance analyses showed surprisingly little variation between members of the Delphinidae (dolphins). This indicated an unexpectedly recent radiation in the Delphinidae, and therefore directs further study to a reevaluation of the fos sil record, or to the need for studies on the calibration of the molecular clock in marine mammals. DNA Sequencing This method is able to provide information at any level of phylogenetic analysis. and is the method of choice for definitive systematic studies. The ability to obtain sequence data from long dead tissues provides totally novel uses for museum material and the opportunity to demonstrate the phylogenetic affinities of extinct species. For example, DNA extracted and amplified from 14,000 year-old bones of the <strong>sa</strong>bre-toothed cat Smilodon jaJalis have affirmed the placement of Smilodon within the modem radiation of Felidae. The advent of the polymerase chain reaction (PCR) technique as a means of amplifying specific regions of DNA suitable for sequencing has greatly enhanced and simplified the use of DNA sequencing, making it the method of choice for many applications. Aligned DNA 91
sequences from related taxa provide the most direct approach to the production of character sets since the base position can represent the character, and the type of base (adenine, guanine, cytosine, or thymidine) can represent the character state. Distance matrices are readily derived from the pairwise proportion of shared bases. Figure 8.3 illustrates the (at least initial) simplicity of evaluating DNA sequence data in a comparison of 100 bases of mitochondrial DNA from some Southern African Felidae, and gives an indication of the amount of change typically found at the generic and species level in this family. There are a number of features of note: there are changes at 18 positions; 10 of these are phylogenetically informative (marked with an *), meaning that they can be used by cladistic (e.g. parsimony) methods to construct a phylogenetic tree and require that each character state occurs at least twice, and not more than a number given by the total number of taxa less two (i.e. four in this example). The initial assumption of the cladistic approach is that at anyone of these phylogenetically infonnative sites one of the shared bases represents a derived character state, Or synapomorphy (the other representing the primitive state, or symplesiomorphy). The other positions represent autapomorphies and are not used by cladistic methods, (but are by distance methods). 10 20 30 40 50 F. lybica CCAACTTGCC TCCATCCTAT ATTTCTCAAC CCTCCTAATC CTAATACCCA F. sylvestris ...... A .. · ......... . .......... .......... . ......... F. nigripes ...... A .. T .......... ·C .... • .. · T .. ·· .. G .. . ......... C. caracal ...... A· •• -......... ·C· ...... · ...... ·G .. · .... · .. T· P. leo ...... A .. · .......... ·C .... ·C .. T .. T ...... .......... P. pardus ...... A .. · .......... ·C .... ·C·T .. ·T· .. G .. ...... .... * * * * * 60 70 80 90 100 F. lybica TCTCAGGCAT TATTGAAAAC CGTCTACTCA AATGAAGAGT CTTTGTAGTA F. sylvestris ......... . . ......... .. C .. · .... .......... .......... F. nigripes .......... .......... ........ T· · G .... · .. · .......... C. caracal ·T .. · .... · C .. C ...... "C"C"A' . . . . . . . . . . .......... P. leo ......... . .......... .. C .. C .... ........... ..·C...... P. pardus .......... C .. C ...... "C"C"A' .......... .......... • • • • • Figure 8.3 Comparison of 100 bases of mitochondrial DNA from some southern African Felidae. DNA was extracted from cell cultures or postmortem tissues. Pan of the mitochondrial cytochrome b region, corresponding to positions 15566-15665 in the bovine sequence was amplified using PCR, and sequenced. Dots indicate bases identical to the F. Iybica reference sequence. * signals a phylogenetically infonnative site. ---, bases absent after aligrunent. The major problem with the cladistic approach is the noise given by unrecognized homoplasies (parallel or reverse mutations) which are very commOn in sequence data. For example, the bases shared by the lion Panthera leo and caracal Felis caracal, but not by the leopard Panthera 92
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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,
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 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 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
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
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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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