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A Study on Fern and Fern-Ally Spore Composition in Miocene Sediments to Improve Our Understanding of the Origins of Fern Diversity in Western Amazonia

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A Study on Fern and Fern-Ally Spore Composition in Miocene Sediments to Improve Our Understanding of the Origins of Fern Diversity in Western Amazonia A study on fern and fern-ally spore composition in Miocene sediments to improve our understanding of the origins of fern diversity in western Amazonia L. van den Bos – 10829148 Future Planet Studies: Earth Science University of Amsterdam Primary advisor: M.C. Hoorn Secondary advisor: C.N.H. McMichael July 3rd 2017 Index Abstract ................................................................................................................................................... 3 Introduction ............................................................................................................................................. 3 Area description ...................................................................................................................................... 5 Theoretical framework ............................................................................................................................ 6 Methodology ........................................................................................................................................... 7 Modern fern flora ................................................................................................................................ 7 Miocene spore determination ............................................................................................................ 7 Results ..................................................................................................................................................... 9 Discussion .............................................................................................................................................. 12 Relation between the different groups ............................................................................................. 12 The Pebas over time .......................................................................................................................... 12 Andean genera .................................................................................................................................. 13 The Várzea ......................................................................................................................................... 14 Conclusion ............................................................................................................................................. 14 Acknowledgement ................................................................................................................................. 15 References ............................................................................................................................................. 15 Appendix ................................................................................................................................................ 19 Appendix 1 ......................................................................................................................................... 19 Appendix 2 ......................................................................................................................................... 38 Abstract There are many different theories about the cause for the high biodiversity of western Amazonia. Andean uplift in the Cenozoic era is believed to be one of the main drivers for the high species- richness in this area. Pteridophytes are important contributors to the species-richness of the Amazonian rainforest. Their diversity and distribution patterns, related to specific soils, are a motivation to research their development through time. In this study, the fern composition of Miocene sediments of western Amazonia is analyzed and compared with modern taxa in Amazonia and on the Andean slopes, to increase our understanding of the origin of fern diversity. I used a combination of microscope analysis and a literature study. Through time, an increase in unique genera comes forth and typical Andean genera enter the dataset from the mid- to late-Miocene. The cation-rich Pebas formation, which initiates in the early Miocene, becomes more biodiverse over time. Surprisingly, the similarity of the fern composition between the Miocene Pebas formation and the modern Pebas formation does not seem to be significantly bigger than the similarity between the Miocene Pebas and the modern Nauta formation. This study also suggests that, in relation to overall sporomorph composition throughout the Miocene, the fern spores do not vary much in abundance, but the composition does change over time. I conclude that these changes are most likely due to the increasing influence of the Andes mountain range as sediment provider to western Amazonia, which generated a more enriched soil type and also brings in spores from the mountain forests. Keywords: Western Amazonia, Andean uplift, pteridophytes, Miocene, spore determination Introduction The cause of the species-richness of western Amazonia – the most biodiverse terrestrial ecosystem on our planet (Hoorn et al., 2010) – is a subject much debated over the past decades. In 1969, Haffer proposed the refuge theory to explain the diversity of bird populations. This theory states that during dry climatic periods in the Pleistocene, the Amazonian forest was divided into many smaller forests, thereby isolating animal- and plant species. During the wet climate periods the forests reunited and species dispersed. According to Haffer (1969), this happened several times and this led to the high biodiversity in western Amazonia. More recent studies suggested that Pleistocene climate change could not have caused forest fragmentation (Colinvaux, Irion, Räsänen, Bush & De Mello, 2001). Many other articles state that the origin of the high biodiversity of western Amazonia lies in the many dispersal barriers in the form of rivers and subsurface arches (Dinerstein et al., 1995; Olson et al., 2001; Lougheed, Gascon, Jones, Bogart & Boag, 1999; Patton, Da Silva & Malcolm, 2000). This paper, however, is based on the assumption that the cause of the high biodiversity finds its origin on a far larger timescale: the Andean uplift in the Cenozoic era. Hoorn et al. (2010) explain that the Andean uplift had great effects on the landscape evolution of northern South America. The Andes formed a barrier, changing river drainage patterns as well as climate. The changing environment of the past 65 Ma still divides the Amazonian rainforest on the basis of geological formations and their edaphic properties (Fig. 1). Figure 1: Paleogeographic maps of the northern South America in the Cenozoic era (Hoorn et al., 2010) Western Amazonia is characterized by different types of landforms that are related to specific types of bedrock. Each of these geological formations have their own characteristic types of soil and plant biomes (Higgins et al., 2011). Most characteristic are the crystalline rocks of the Amazon Craton, the white sand units related to rivers with their source in Amazonia, the Miocene Pebas (or Solimões formation as it is called in Brazil) which are fluvio-lacustrine sediments mostly of clay composition and with their source in sub-Andean areas, and younger fluvial sand/clay units such as the Ica and Nauta formations sourced from the Andes (Hoorn & Wesselingh, 2011; Higgins et al., 2011; Räsänen, Linna, Santos & Negri, 1995). Worldwide, there are 10,560 different known species divided into 240 genera and approximately 33% of them occur in the American tropics (Christenhusz & Byng, 2016; Tryon & Tryon, 2012). Due to the unique circumstances and edaphic properties, some pteridophytes species are endemic in distinct Amazonian regions (Tryon & Tryon, 2012). The modern fern composition in western Amazonia is well studied by Higgins et al. (2011) and Tuomisto et al. (2016) who presented an inventory of the pteridophyte species composition in their study areas. They related the characteristic pteridophyte composition to both soil type and geological formation (either Pebas or Nauta formation) and found there was a distinct difference depending on the geological formation on which a soil was formed. The aforementioned works form an interesting basis for a comparison of the spore fossils found in Miocene sediments, and can give insights in the development of pteridophytes in western Amazonia. This research aims to improve our understanding about the origin of pteridophyte diversity through a palynological study on spores in Miocene sediments. Here we compare the fern spores composition on the early- to mid-Miocene cation-rich Pebas formation (25-10 Ma) with fern composition on the associated soils of both Pebas and the late Miocene, cation-poor Nauta formation (10-5 Ma). We also compare these data with the fern composition in the modern, often inundated, várzea and the high altitudes of the modern Andes (Higgins et al, 2011). Insights into the origins of fern diversification will increase our knowledge about the development of the modern Amazon and may help us preserve this endangered ecosystem in the future. The following central question will be addressed in this thesis: Is there a close relation between the current pteridophyte composition on Nauta and Pebas soils and the pteridophyte composition of the Miocene Pebas in western Amazonia? Furthermore, we will investigate how the relative abundance of fern spores changed through time in comparison to other sporomorphs, whether the composition of the fern flora on the Pebas formation changed through time, at what point in time typical
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