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Cytotaxonomic Studies in Themeda Triandra Forssk.: IV

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Cytotaxonomic Studies in Themeda Triandra Forssk.: IV S.AfrJ.Bot., 1993, 59(3): 305 - 310 305 Cytotaxonomic studies in Themeda triandra Forssk.: IV. A population cytogenetic study of a contact zone between tetraploid and hexaploid populations H. Liebenberg*, J. Lubbinge and Annabel Fossey Department of Genetics, University of Pretoria, Pretoria 0002, Republic of South Africa Received 17 October 1992; revised 20 January 1993 A population cytogenetic study was undertaken across a contact zone between Themeda triandra popula­ tions in the Transvaal Bushveld to the north of the Magaliesberg, and Highveld populations to the south. Fifteen collection points were chosen and three plants collected at each point. The northern-most collection point contained only tetraploids and those on the Highveld only hexaploids. In the vicinity of the Magaliesberg Mountain range, contact between the tetraploids and hexaploids produced pentaploid and decaploid hybrids. Meiotic pairing association analyses showed that both the tetraploids and hexaploids formed apomictic clones, of which some were widely distributed and others were more limited. This indicates that facultative apomixis occurs among both the tetra- and hexaploids and gives rise to genotypic and genomic segregants thus producing variation and forming new apomictic clones. The possible evolution of Themeda triandra in southern Africa is discussed. 'n Bevolking sitogenetiese ondersoek is onderneem deur 'n kontaksone tussen Themeda triandra bevolkings in die Transvaalse Bosveld, noord van die Magaliesberg, en op die Hoeveld suidwaarts daarvan. Vyftien versamelpunte is gekies en drie plante is by elke versamelpunt versamel. Die versamelpunt wat die verste noord was, het slegs tetraploiede gelewer, terwyl die op die Hoeveld slegs heksaploiede bevat. In die omgewing van die Magaliesbergreeks, waar kontak tussen tetra- en heksaploiede voorkom, is pentaploiede en dekaploiede basters gevind. Meiose parings-assosiasie analises het aangetoon dat beide die tetra- en heksaploiede apomiktiese klone bevat. "Party van die klone vertoon 'n wye verspreiding, terwyl ander se verspreidings beperk is. Hierdie verskynsel toon dat fakultatiewe apomiksie voorkom by beide die tetra- en heksaploiede. Dit gee aanleiding tot nuwe genotipes en genoomsegregante wat weer nuwe apomiktiese klone vorm . Die moontlike ewolusionere ontwikkeling van Themeda triandra in suider Afrika word bespreek. Keywords: Agamic complex, cytogenetic, microsporogenesis, polyploidy, Themeda triandra . • To whom correspondence should be addressed. Introduction The distribution of different polyploid levels of Themeda triandra in southern Africa show some general tendencies (Gluckmann 1951; De Wet 1960; Liebenberg 1986), al­ though no distinctive distributional pattern has been identi­ N fied. The diploids tend to occur towards the eastern and southern lower-lying coastal areas, whereas the hexaploids are concentrated on the central highveld regions. Intermedi­ De Wildt ate polyploids (4X and 5X) tend to occur inbetween. In the central Transvaal, the Magaliesberg Mountain Pretoria North + range (an east/west mountain range) appears to form a transitional region between the predominantly hexaploid populations of the higher, southern region and the northern bushveld region where diploids, tetraploids and even some Pretoria West hexaploids occur. Recent studies (Fossey & Liebenberg 1987; 1992) have shown that in the area east of Pretoria they are separated into two apparently entirely diploid (to the north) and hexaploid (to the south) populations. The distance between the two populations is a mere 25 km . The transition from Bushveld (c. 1 100 m above sea level) to Roads Highveld (c. 1 500 m above sea level) on the western side Ra11wayl1ne o Collec1ion po1n 1s of Pretoria, is more complex. Apart from the Magaliesberg Mountain range, there are at least two smaller ranges before the Highveld is reached. The aim of this study was to ascertain how and where the Johannesburg transition between the different polyploid levels occurs. We will report on a population cytogenetic study of 45 plants collected along a transect covering this transition. Figure 1 Road map indicating the collection points. 306 S.-Afr.Tydskr.Plantk., 1993, 59(3) Materials and Methods hexaploids. Regarding the decaploids, the meiosis was fairly Plants were collected along the side of the road between irregular and, with their high chromosome number, it proved Rashoop (north of Pretoria) and Brits, and over the very difficult to obtain large enough numbers of well spread Magaliesberg towards Johannesburg, a distance of c. 55 km metaphase I meiocytes from which reliable analyses could (Figure 1). The 15 collection points (numbered 1 - 15) be done and therefore no association tables could be chosen along this road included points on the plains, in compiled. valleys, on top of mountain ranges and on the northern and The data of a comparative analysis of metaphase I mono­ southern slopes (Figure 2). At each of the points, three valents and anaphase I laggards for the tetra-, penta- and plants (labelled a, b or c) were collected from an area not hexaploids, based on information from 100 meiocytes each, exceeding 5 m2 • The plants were selected by throwing an are presented in Table 4. A highly significant positive iron hoop in the air and then collecting the plant nearest to correlation exists between the number of metaphase I mono­ where it landed. Slides prepared from some plants were valents and the number of anaphase I laggards (r = 0.99 for inadequate and therefore these collection points were the tetraploids, r = 0.98 for the pentaploids and r = 0.97 for revisited. the hexaploids). This indicates that the analyses are accurate The inflorescences were fixed in Pienaar's solution, 6:3:2 and highly repeatable. (methanol: chloroform: propionic acid) (pienaar 1955) and squashed in 1% propionic carmine. One hundred Pollen Discussion Mother Cells (PMCs), each of metaphase I and anaphase I Previous studies have indicated that the Magaliesberg were scored for chromosome number, pairing associations, Mountain range, in the vicinity of Pretoria (central Trans­ laggards and any other irregularities. vaal), forms a boundary between predominantly hexaploid populations to the south, and tetraploid and diploid popula­ Results tions (Fossey & Liebenberg 1987) to the north. Hexaploids The chromosome numbers of the 45 plants are given in do, however, also occur north of the Magaliesberg. Gluck­ Table 1. At collection point 1 tetraploids were encountered. mann (1951) and Liebenberg (1986) have reported on hexa­ The region from point 2, c. 6 km north of the Magaliesberg, ploids from Warmbaths and Naboomspruit, respectively. On to point 6, at the foot of the Magaliesberg, shows a clear the other hand De Wet (1960), whose study was mainly transition from tetraploids to hexaploids. Here tetraploids, based on stomatal size, reported no hexaploids for the pentaploids, hexaploids and decaploids were found. At the Bushveld area. It would appear that the distribution patterns southern collection points (7 - 15), only hexaploids of diploid and different polyploid populations in the central occurred. and northern Transvaal is more complex and needs further The meiotic pairing associations of the four tetraploids study. The fact that the diploid population studied by Fossey (Table 2) and of the hexaploids (Table 3) have been group­ & Liebenberg (1987) and the tetraploids found in this study, ed into similar pairing association groups. The hexaploids occur at about the same latitude south, about 50 km apart, have further been subgrouped to facilitate comparisons . reflects the intricate distribution of the different ploidy Each group represents plants with similar association pat­ levels in the Transvaal Bushveld. terns, whereas the subgroups represent finer subdivisions The collection localities of this study resulted in the within the groups. The groupings are based on the number identification of a contact zone between tetraploids and of pairing association types found and the relative frequen­ hexaploids. This contact zone has proven to contain hybrids cies. In the case of the pentaploids, the pairing associations between tetraploids and hexaploids, resulting in pentaploids proved to be so variable that it became impracticable to (collections 2b, 4a, b & c). These pentaploids apparently present them in tables such as those used for the tetra- and undergo chromosome doubling from time to time, producing decaploids (collections 5c & 6c). This is the first time, as far as could be ascertained, that decaploid T. triandra have been reported to occur. Other ploidy levels, not yet found in southern Africa, are the octo-, nona- and endecaploids (8X, Height 9x & llX) found in India (Birari 1980). It would seem that Above Sea further south (collection points 7 - 15) only hexaploids Level occur. These findings are supported by those of Gluckmann 1600 1550 15 (1951), Liebenberg (1986) and Fossey & Liebenberg (1987) 14 1500 that the T. triandra populations south of the Magaliesberg, 1450 • 10 1400 in the Pretoria vicinity, are predominantly hexaploid. Gluck­ l.BO mann (1951), however, found pentaploids in Johannesburg 1300 lHO and the eastern Transvaal Highveld (Standerton and Devon). 1200 A significant finding uf this study, however, was the 1100 occurrence and distribution of apomictic clones (the apospo­ 1050 ric progeny of the same apomictic lineage, having the same MI:,::.'-------.-I--.--,---.--,----,----.---.--,----,--;I~ 5 10 15 20 25 30 35 40 45 50 55 genotype
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