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Msc Biodiversity and Taxonomy of Plants

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Msc Biodiversity and Taxonomy of Plants MSc Summer Research Project Floral development of Cuphea (Lythraceae): understanding the origin of monosymmetry and the epicalyx in the flower Celina Barroca | August 2014 Thesis submitted in partial fulfilment for the MSc in the Biodiversity and Taxonomy of Plants RBGE | University of Edinburgh 2 MSc Biodiversity and Taxonomy of Plants Front page: Cuphea procumbens SEM picture RBGE | University of Edinburgh 3 MSc Biodiversity and Taxonomy of Plants Abstract The genus Cuphea (Lythraceae) is highly variable and floral morphology within this genus is highly diverse. It is also one of the most strongly zygomorphic genera within Lythraceae. Another notable character is the presence of an epicalyx. The origin of epicalyx within the family is unclear and debatable. Accepting Mayr’s (1969) definition, epicalyx is an “emergence of congenitally fused sepals”. The ontogeny of the epicalyx in Cuphea is very different from that observe in other families such as Dipsacaceae, Rosaceae and Malvaceae and has not been the subject of a comprehensive ontogenic study. This study has investigated the ontogenic development of the expicalyx and also the monosymmetry observe in six species of the genus Cuphea. This was undertaken using light microscopy and scanning electron microscopy. It revealed that the epicalyx arises very late in the ontogenic sequence and the monosymmetry of the flower is a result of the loss of a stamen and the development of a large nectary on only one side of the base of the ovary. The position of the nectary causes the formation of a large nectar sac on the adaxial side of the hypanthium resulting in a strongly zygomorphic flower. RBGE | University of Edinburgh 4 MSc Biodiversity and Taxonomy of Plants Table of Contents Abstract........................................................................................... 3 Table of Contents........................................................................... 4 List of Figures................................................................................ 6 List of Tables.................................................................................. 6 Introduction.................................................................................... 7 Angiosperms Background................................................................. 7 Flower...................................................................................................... 8 Perianth................................................................................................... 9 Symmetry................................................................................................. 9 Merism...................................................................................................... 10 Ontogeny.................................................................................................. 11 Epicalyx – The unresolvable structure............................................... 12 Study Group....................................................................................... 15 Aims of the study............................................................................... 25 Material&Methods......................................................................... 26 Plant Material..................................................................................... 26 Scanning electron microscopy........................................................... 27 Results............................................................................................ 28 Morphological observations............................................................... 28 Scanning electron microscopy.......................................................... 30 Epicalyx and Symmetry........................................................................... 30 Stamen, Gynoecium and Nectary............................................................. 32 Lighting Microscopy.......................................................................... 34 Petals....................................................................................................... 34 Monosymmetry........................................................................................ 36 Stamens........................................................................................ 36 RBGE | University of Edinburgh 5 MSc Biodiversity and Taxonomy of Plants Nectary......................................................................................... 38 Discussion........................................................................................ 41 Comparative analysis of different species ......................................... 41 Epicalyx origin................................................................................... 41 Evolution of the flower in Cuphea: monosymmetry, merism............ 42 Conclusions.................................................................................... 45 Recommendations.......................................................................... 46 References...................................................................................... 47 RBGE | University of Edinburgh 6 MSc Biodiversity and Taxonomy of Plants List of Figures Figure 1. Phyllogenetic hypothesis of relationships of the families belonging to 16 the order Myrtale………………………………………….……………. Figure 2. Distribution map of Lythraceae………………………………………… 17 Figure 3. Phyllogenetic tree for the genus Cuphea…………………………………... 20 Figure 4a. Cuphea cyanea flower………………………………….......................... 21 Figure 4b. Cuphea cyanea inflorescence………………………………………… 21 Figure 5a. Cuphea micropetala flowers…………………………………………… 21 Figure 5b. Cuphea micropetala inflorescence……………………………………... 21 Figure 6a. Cuphea ignea flower…………………………………………………… 22 Figure 6b. Cuphea ignea solitary flower…………………………………………... 22 Figure 7a. Cuphea caeciliae flower………………………………………………... 22 Figure 7b. Cuphea caeciliae inflorescence………………………………………… 22 Figure 8a. Cuphea hyssopifolia flowers…………………………………………… 23 Figure 8b. Cuphea hyssopifolia inflorescence……………………………………... 23 Figure 9a. Cuphea procumbens flowers…………………………………………… 23 Figure 9b. Cuphea procumbens inflorescence……………………………………... 23 Figure 10. C. caeciliae SEM figure plate.................................................................. 31 Figure 11. C. caeciliae SEM figure plate.................................................................. 33 Figure 12. C. caeciliae and C. procumbens SEM figure plate…………………… 35 Figure 13. Stamens of fixed dissected half mature flowers plate………………….. 37 Figure 14. Nectary of fixed dissected half mature flowers plate…………………... 39 Figure 15. Floral diagram of Cuphea micropetala............................................................... 40 List of Tables Table 1. Plant material collection details for each species…………………………. 28 RBGE | University of Edinburgh 7 MSc Biodiversity and Taxonomy of Plants Introduction Angiosperms Background With at least 260,000 living species classified in 59 orders and 413 families (APG III, 2009), the Angiosperms, or flowering plants, are one of the major groups of land plants and the most diverse extant group, comprising around 90% of existent plant biodiversity. However, this diversity in not equally distributed and around 75% of all species belong to the eudicot clade (Soltis et al., 2005, Judd et al., 2007). Angiosperms are far more diverse in vegetative form and in the structure of their reproductive organs than any other group of land plants (Friis et al. 2011). According to the fossil record based on distinctive angiosperms pollen grain fossils have been dated from early Cretaceous, around 135 million years ago, time of angiosperms origin (Friis et al. 1987, 2011, Judd et al. 2007). Therefore, angiosperms diversified during the Cretaceous period. Many authors suggest different hypothesis and fossil record evidence to date the divergence of angiosperms, evolution and later their lineages e.g according to Friis et al. (2005) the estimated timing of major diversification events in angiosperms range from around 175 million years ago – during Jurassic to around 60 million years ago – at the end of the Cretaceous. What is certain is, by the end of the Cretaceous, angiosperms have diversified enormously and become widespread, becoming the dominant terrestrial plants on the planet (Magallón and Sanderson, 2001). Angiosperms have a few synapomorphies (i.e. shared derived characters) that unify and distinguish them from the other groups of plants. These features include: 1) ovules that are enclosed within a carpel (i.e. a structure that consists of an ovary, which encloses the ovules, the style, a stalk portion as a slender region specialized for pollen tube growth and the stigma, a structure which receives the pollen and where pollen germination takes place; 2) double fertilization, which leads to the formation of an endosperm (a nutritive tissue within the seed that surrounds and feeds the developing plant embryo); RBGE | University of Edinburgh 8 MSc Biodiversity and Taxonomy of Plants 3) stamens with two pairs of microsporangia (i.e. pollen sacs); 4) features of female and male gametophyte reduced in size structure and development; 5) phloem tissue composed of sieve tubes accompanied by one or more companion cells (see Doyle and Donoghue, 1986; P. Soltis and D. Soltis, 2004; Judd et al. 2007). Strong evidence for the monophyly of angiosperms comes from shared derived morphological characters mentioned above but also from molecular studies,
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