The absence of the majority of conifers from tropical forests remains an unresolved evolutionary and biogeographic enigma. Correlations between a flattened conifer leaf morphology and tropical persistence have led to the hypothesis that shade tolerance is critical for conifer survival in these environments. Yet, little is known of the physiological mechanisms constraining conifer distributions. This dissertation, using the diversity of leaf morphologies and biogeographic distributions found among conifer families, examines this hypothesis from a physiological perspective. The first chapter examines the photosynthetic process in shade-house grown saplings of conifers coexisting in the Central Highlands of Vietnam, an area of unusual tropical conifer diversity. Pinus krempfii, a flat-leaved pine endemic to the densely-canopied tropical forests of Vietnam, demonstrates an intermediate photosynthesis between needle-leaved Pinus and flat-leaved Podocarpaceae, thus providing support for shade tolerance as an important driver of conifer success in tropical forests. In chapter two, respiratory traits are examined in adult conifers growing naturally in the Central Highlands of Vietnam. Flat-leaved conifer species are found to have lower respiration than needle-leaved species. Finally, in chapter 3, the respiratory temperature responses of members from the two largest conifer families, the Pinaceae and Podocarpaceae, are compared. Consistently lower respiration in Podocarpaceae compared to Pinaceae suggests that there are phylogenetic underpinnings to the respiratory differences between the two families. These findings correlate to the biogeographic differences between the two families, with Podocarpaceae being notably more successful in tropical forests than Pinaceae. Together, all three chapters demonstrate that lower foliar carbon fluxes and flattened leaf morphologies are characteristics of a shade tolerance strategy that is necessary for conifer persistence in tropical forests. Ultimately, this dissertation makes a unique link between conifer physiologies and their current biogeographic distributions that will pave the way for a mechanistic tackling of their future survival in a changing climate.

A review of the literature revealed that there is a lack of depth in our knowledge of gymnospermous seeds with regard to the development and germination processes. The phytohormones, particularly the cytokinins have been implicated in these processes. The seeds of Podocarpus henkelii were thus selected as experimental material for studying the biochemical and cytokinin-like changes associated with development and germination. The development of these seeds was also followed at the ultrastructural level. These studies revealed that cellular detail within the female gametophyte only began .to form in December (early summer), approximately six weeks after fertilization had taken place. At this time some reserve protein was evident and the embryo sporophyte consisted of only a few pro-suspensor and pro-embryo cells. Concurrently, the cytokinin levels were fairly high in the female gametophyte but low in the epimatium. In both seed components two cytokinin- like compounds predominated which co-chromatographed with the free base cytokinin zeatin and its ribonucleoside. The second sample was taken in late January (mid-summer) by which time the embryo sporophyte had developed rapidly into a readily distinguishable seed component. The cellular detail indicated that much cell division had recently taken place and that the cells were currently increasing in size and accumulating starch and lipid. In the female gametophyte the soluble sugars were at the maximum level recorded during these experiments and the level of starch was increasing. The extractable cytokinin content of the seed was high at this time, particularly in the embryo sporophyte. In all three seed components cytokinin-like compounds which co-chromatographed with zeatin and ribosylzeatin were present. These high levels of cytokinin coincided with the rapid increase in both fresh and dry mass of the embryo sporophyte and female gametophyte. Ultrastructural studies of the third sample collected in mid-March (early autumn) showed that cellular changes were associated mainly with increases in cell size and the accumulation of food reserves, particularly starch. The cytokinin levels had decreased in all three seed components at this time. There was an increase in the cytokinin which co-chromatographed with glucosylzeatin in the female gametophyte. The seeds matured in late April (autumn) and had the unusual features of not drying out during maturation. Fresh seeds collected from the ground had a moisture content of ca. 62 per cent. The main food reserve was starch with relatively small amounts of protein and lipid also present. The seeds of Podocarpus henkelii germinated readily after scarification in the absence of water provided that their moisture content remained ca. 60 per cent. Seeds in which the moisture content had fallen below ca. 54 per cent required additional water for germination. The moisture content of the seeds fell rapidly under natural conditions and viability was lost below a moisture content of ca. 34 per cent. Unscarified seeds of 52 per cent moisture content placed under moist conditions at a constant 25°C took 23 weeks to achieve 68 per cent germination. These experiments showed that although the epimatium limited water uptake by the seeds it did not prevent moisture loss to the atmosphere. This appears to be the main factor contributing to the seed's inefficiency as a propagule. A small degree of after-ripening was recorded with the embryo sporophyte increasing in size with storage. This appeared to contribute to the increased rate of germination of the scarified seeds. An interesting feature of the seeds of Podocarpus henkelii is that they have the ability to fix atmospheric carbon, which is subsequently translocated from the epimatium to the female gametophyte and embryo sporophyte. The mature seeds were stored at 4°C for six weeks during which time little change had occurred at the ultrastructural level. Protein vacuoles in the embryo sporophyte had disappeared and in all three seed components cytokinin levels were low. Three days after scarification and the start of incubation, little change in cellular detail was apparent as limited rehydration was necessary due to the high moisture content. The cytokinin levels in the embryo sporophyte and epimatium had increased, whilst the levels in the female gametophyte had decreased at this time. In the embryo sporophyte lipid mobilization had commenced with these reserves apparently being metabolized within vacuoles. The rate of respiration measured in terms of increases in CO[2] evolution, increased 60 hours after the start of the incubation period, just 12 hours before ten per cent germination was recorded. Germination was accompanied by a large increase in the levels of cytokinins in the female gametophyte and embryo sporophyte. The cytokinins detected co-chromatographed with the free base cytokinin zeatin and its riboside, ribosylzeatin. Concurrently, marked ultrastructural changes were recorded with increases in the amounts of dictyosomes, endoplasmic reticulum and the formation of polyribosomes, all of which are indicative of increased metabolic activity. Similar increases in the female gametophyte were ofa lower order and occurred only after nine days of incubation. By this time the levels of cytokinins had decreased considerably. After 12 days of incubation 65 per cent of the seeds had germinated. As much of the food reserves in the female gametophyte and embryo sporophyte remained, it is suggested that these reserves are utilized for subsequent seedling establishment rather than for germination. The actual role that cytokinins play in the development and germination of these seeds is not clear. High levels of this phytohormone coincide with periods of food deposition and mobilization suggesting that they play an important part in these processes. The results of the biochemical, cytokinin and ultrastructural studies are discussed in relation to the developmental and germination processes and are compared to the data of other seeds. / Thesis (Ph.D.)-University of Natal, Pietermaritzburg, 1981.

Submitted by Fabricia Fialho Reginato (fabriciar) on 2015-07-03T22:44:25Z No. of bitstreams: 1 DaianaFontes.pdf: 25455596 bytes, checksum: 4dae1fc6ad9f052e0a07f54f6be98834 (MD5) / Made available in DSpace on 2015-07-03T22:44:25Z (GMT). No. of bitstreams: 1 DaianaFontes.pdf: 25455596 bytes, checksum: 4dae1fc6ad9f052e0a07f54f6be98834 (MD5) Previous issue date: 2008 / CAPES - Coordenação de Aperfeiçoamento de Pessoal de Nível Superior / A composição e a diversidade dos representantes de Podocarpaceae nas tafofloras da ilha Rei George, norte da Península Antártica, atestam a importância deste grupo de coníferas nesta região no início do Cenozóico, após uma longa história evolutiva e uma origem igualmente ligada às altas latitudes do sul. O registro fóssil demonstra uma adaptabilidade e capacidade para conquistar novas áreas, diferentemente das outras famílias modernas de gimnospermas austrais, provavelmente devido à dispersão zoocórica e a variedade de formas e hábitos que ainda hoje mantém e que garantiram sua sobrevivência em praticamente todas as faixas de temperatura e gradientes altitudinais. Sete formas relacionadas a esta família foram descritas e se caracterizam pelos dois tipos principais de ramos, ainda hoje existentes. O primeiro deles caracterizado por folhas bifacialmente aplainadas, pequenas e mais ou menos adpressas, representados por duas espécies de Lepidothamnus sp., por Halocarpus sp. e Dacrycarpus sp. O outro grupo, com folhas grandes, com disposição bilateralmente aplainada e com uma veia média destacada, está representado por formas dos gêneros Saxegothaea e Podocarpus, para o qual uma nova espécie é pela primeira vez comunicada. Os macrorrestos estão preservados como impressões e carbonizações em dois dos principais locais da ilha que expõem as litologias durante o verão, o Monte Wawel, na baía do Almirantado, e o Morro dos Fósseis, na península Fildes. Representam sucessões vulcânicas e vulcanoclásticas, englobadas em duas unidades litoestratigráficas distintas, mas provavelmente correlacionáveis, as formações Fossil Hill e Vièville Glacier. Os níveis fossilíferos são compostos por grãos de origem vulcânica, retrabalhados ou não, que atestam sua afinidade com ambientes tectonicamente perturbados e com áreas baixas, de solos rejuvenescidos, onde havia condições para a formação de pequenos lagos e deltas durante as fases de erupções. O levantamento e comparação com outras paleofloras austrais e as idades radiométricas disponíveis permitiram propor uma idade Eoceno Médio para os diferentes níveis onde as podocarpáceas estão presentes e a vigência de um clima temperado úmido que, de modo relativamente rápido, dá lugar a evidências de queda nas temperaturas. A análise dos comparativos modernos sugere que compunham uma flora única, mista e diversificada, que reúne elementos com hábitos e adaptações distintas, se desenvolvendo em diferentes gradientes altitudinais, mais uma evidência em apoio à presença de áreas altas (vulcões?) e costeiras. Quando comparada aos modernos biomas, a paleoflora da ilha Rei George também exibe uma mescla de elementos, unindo hoje os que estão distribuídos nos dois extremos do Hemisfério Sul (América e Australásia), em grande parte endêmicos, e que vivem em lugares submetidos a baixas temperaturas. Assim ao mesmo tempo em que apóia condições climáticas semelhantes para o norte da Península Antártica durante o Eoceno, evidencia o importante papel que as terras do Gondwana e sua separação tiveram, sobre a moderna distribuição disjunta dos representantes desta família. / The composition and diversity of the Podocarpaceae representatives in the fossil plant assemblages of King George Island, north of Antarctic Peninsula, indicates their importance in that region during the Early Cenozoic, after a long evolutive history linked, since its origin, to the high southern latitudes. The fossil record attest an adaptability and capacity to conquer new areas that have no comparatives in other austral conifers, probably due to their seed dispersion by distinct groups of animals and the diverse forms and habits that their modern relatives still maintain. It guarantees their survival until today and their adaptation in nearly all latitudinal and altitudinal gradients. Seven distinct forms related to this family were described, characterized by two main kinds of leaf shoots. Those with scale and short leaf, bifacial flattened shoots are represented by two species of Lepidothamnus and by Halocarpus, Microcachrys and Dacrycarpus. The other one, with bilaterally flattened and big size leaves, with a sharp midrib, was associated with Saxegothaea and a species of Podocarpus, wich was for the first time described in the island taphofloras. The macrofossils are preserved by impressions and charcoalified materials, in the two main areas of the island where the lithologies are exposed in summer seasons, the Mount Wawel, at Admiralty Bay, and the Fossil Hill, at Fildes Peninsula. Represents what was included in two probably correlative lithostratigraphic units established in previous geological works, the Vièville Glacier and Fossil Hill formations. The fossil levels are composed mainly by volcanic grains, sometimes with signals of reworking by shallow lakes and deltas and confirm an environment and deposition linked to moments of active volcanism and inter-eruption phases. The comparisons with other fossil floras from the Antarctic Peninsula and Circum-Antarctic areas, and the radiometric ages indicates a probable lower mid-Eocene age, and wet and temperate climatic conditions, which deteriorates along the succession. The nearest modern relatives of the fossil taxa preserved in the taphofloras suggests also an unique, mixed and diversified vegetation, joining elements with distinct habits and that lives today in distinct altitudinal gradients, provides further evidence in support of the presence of high (volcanoes?) and coastal areas. The King George Island paleoflora flora also comprises a mixture of endemic, eastern Gondwana and South American elements, which reflect its position close to a major floristic boundary. At the same time confirms the microthermic conditions of climate in the north of Antarctic Peninsula at the Eocene times and the important role of the Gondwana drift apart, and the derivate climatic changes, in the modern disjunct distribution of the family.