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220 8. MOLECULAR PHYLOGENY OF THE GENUS LOLIUM it is reasonable to think that L. loliaceum and L. perenne with both subspecies have the common ancestor such as L. rigidum. L. loliaceum could diverge earlier and that is why it occupies somehow intermediate position between L. rigidum and L. perenne entailing certain inconsistency between dendrograms. The placement of L. loliaceum with a sub-clade with L. rigidum in the SSAP tree is a strong support for the above hypothesis. Thus, the most plausible explanation of greatest similarity between L. loliaceum and multiflorum demonstrating by some molecular data is the hybridisation that has permitted gene introgression. Thus, the inclusion of the multiflorum sequence within L. loliaceum may be a reticulate event. The beginning of diversification of L. perenne populations can be dated to about 167 000 years ago. It is worthy to note that it was a period of the Tertiary glaciations (Klimaszewski 1996) and the perennial type of growth habit might be a sign of past adaptation to harsh conditions. It would mean that perenniality is a relatively recent acquisition and that is why it is not so strong. From the present data and earlier considerations it can be postulated that L. perenne ssp. perenne evolved as a response to climatic changes and this adaptation enabled it to survive. During the interglacial period it became widespread throughout Europe and during the Quaternary Ice Ages its distribution became restricted to refugia. However, populations were not separated completely allowing gene flow due to existence of refugia also in the central part of Europe. In days of the Ice Ages it becomes widespread and extinct alternately. L. perenne ssp. multiflorum has arisen as populations more adapted to southern, warm climate but the gene flow disabled the speciation. Eventually, the unintended selection in the Middle-Ages resulted in the present domesticated form - multiflorum. Its mistaken elevation to a species rank by early taxonomists had led to much controversy that has been continued up to the present. 8.5. CONCLUSIONS (FIGURE 8.12) 1. The most probable common ancestor of the genus Lolium has the closest affinity to the Schedonorus subgenus, of which F. pratensis is the most prominent diploid representative. 2. The genus Lolium can be divided into two clades representing two different lineages that evolved independently from the common ancestor. The first clade (Temulentum) consists of L. persicum, L. remotum and L. temulentum and diverged 2.7 MYA. The second clade (Perenne) consists of L. loliaceum, L. perenne ssp. multiflorum, L. perenne ssp. perenne and L. rigidum and split from Schedonorus 2.35 MYA. Self-fertility arose independently in both clades. 3. Within the Temulentum clade, L. persicum was the first to diversify about 1.3 MYA and the separation between L. remotum and L. temulentum can be dated to 0.95 MYA. It is plausible that current forms of all species within that clade result from unintended domestication about 10 000 years ago. 4. Within the Perenne clade, L. rigidum split into a separate species about 1.4 MYA whereas L. loliaceum about 1.2 MYA. The latter originated through several point mutations that led to self-pollination.
8. MOLECULAR PHYLOGENY OF THE GENUS LOLIUM 221 5. The diversification of L. perenne populations started 167 000 years ago as a response to more harsh conditions during glaciations. In the Quaternary Ice Ages it was widespread and extinct alternately. However, its populations survived in refugia in south and central part of Europe. L. perenne ssp. multiflorum has adapted to more southern conditions. Through selection during domestication and elevation to a species rank the present L. multiflorum was created. 6. Spontaneous or artificial hybridisation between L. perenne and F. pratensis followed by introgression is responsible for incongruence in gene trees, and furthermore makes difficult interfering phylogeny from low copy sequences.
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This research was done within the E
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Acknowledgements I would like to th
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8 CONTENTS 4.4.4. Role of transposo
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10 CONTENTS 9.3.2. Similarity of ge
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12 1. INTRODUCTION
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14 1. INTRODUCTION occasionally use
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16 1. INTRODUCTION On the basis of
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18 1. INTRODUCTION The evidences fr
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20 1. INTRODUCTION overlap between
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22 1. INTRODUCTION et al. 2000; Cat
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24 1. INTRODUCTION According to the
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26 1. INTRODUCTION 1.6. APPLICATION
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28 1. INTRODUCTION of evolution wit
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30 1. INTRODUCTION are continuously
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2. GOALS The aim of this research w
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3. MORPHOLOGICAL VARIATION OF L. MU
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36 3. MORPHOLOGICAL VARIATION... fo
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38 3. MORPHOLOGICAL VARIATION...
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40 3. MORPHOLOGICAL VARIATION... Wi
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42 3. MORPHOLOGICAL VARIATION... va
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44 3. MORPHOLOGICAL VARIATION... 3.
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46 3. MORPHOLOGICAL VARIATION...
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48 3. MORPHOLOGICAL VARIATION... in
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50 3. MORPHOLOGICAL VARIATION... pl
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52 3. MORPHOLOGICAL VARIATION... di
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4. GENETIC DIVERSITY OF L. MULTIFLO
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56 4. GENETIC DIVERSITY... deed con
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58 4. GENETIC DIVERSITY... 4.2.2. D
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60 4. GENETIC DIVERSITY...
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64 4. GENETIC DIVERSITY... RAPD and
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66 4. GENETIC DIVERSITY... SSAP pol
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68 4. GENETIC DIVERSITY... It can b
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72 4. GENETIC DIVERSITY... than wit
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82 4. GENETIC DIVERSITY... power of
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84 4. GENETIC DIVERSITY... Hamrick
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86 4. GENETIC DIVERSITY... deviatio
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88 4. GENETIC DIVERSITY... individu
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90 4. GENETIC DIVERSITY... (inversi
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92 4. GENETIC DIVERSITY... related
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94 4. GENETIC DIVERSITY... cies and
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96 4. GENETIC DIVERSITY... their sp
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98 4. GENETIC DIVERSITY... 4.5. CON
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100 5. ORIGIN OF SEEDLING... Applic
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102 5. ORIGIN OF SEEDLING... 5.2.2.
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104 5. ORIGIN OF SEEDLING... 5.3. R
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106 5. ORIGIN OF SEEDLING... very l
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108 5. ORIGIN OF SEEDLING... 5.3.4.
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110 5. ORIGIN OF SEEDLING... 5.4. D
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112 5. ORIGIN OF SEEDLING... A seco
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114 5. ORIGIN OF SEEDLING... Perhap
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6. DO ANY SPECIES BOUNDARIES EXIST
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118 6. DO ANY SPECIES BOUNDARIES...
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7. ROLE OF QTL s IN THE EARLY EVOLU
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160 7. ROLE OF QTLs IN THE EARLY EV
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Page 170 and 171: 170 7. ROLE OF QTLs IN THE EARLY EV
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Page 236 and 237: 10. MARKER EFFICIENCY 10.1. INTRODU
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Page 248 and 249: 248 11. CONCLUSIONS ABOUT EVOLUTION
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Page 268 and 269: 268 13. SUPPLEMENTARY MATERIALS pla
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270 13. SUPPLEMENTARY MATERIALS
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274 13. SUPPLEMENTARY MATERIALS ANN
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280 13. SUPPLEMENTARY MATERIALS Ann
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290 13. SUPPLEMENTARY MATERIALS •
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292 13. SUPPLEMENTARY MATERIALS The
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296 13. SUPPLEMENTARY MATERIALS fie
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The average gene diversity between
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306 13. SUPPLEMENTARY MATERIALS Unb
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308 14. ABBREVIATIONS Lolcopia1 - L
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310 14. ABBREVIATIONS Drosophila wi
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312 14. ABBREVIATIONS Saccharomyces
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314 15. SUMMARY At different stages
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316 15. SUMMARY L. multiflorum mito
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REVIEWERS’ COMMENTS With a great
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