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Mauritia flexuosa Phylogeny

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Lepidocaryeae, the subtribe classifying the clade the M. flexuosa lies in Julianna Filice
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1. Mauritia flexuosa Phylogeny
Mauritia flexuosa, the aguaje palm, and its phylogenetic relationships Julianna Filice
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1. Mauritia flexuosa Phylogeny
Mauritia flexuosa's closest relative is the Mauritiella armata, commonly called Buritirana or the "ghost palm" Julianna Filice
Contents of this annotation:
1. Mauritia flexuosa Phylogeny

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Aguaje: Tree of Life
Mauritia flexuosa, commonly called aguaje or buriti, is a tall fruit bearing palm tree dominating swampy areas of the amazon rainforest. In these micro ecosystems, they provide various ecosystem services including carbon sequestration, water retention and purification, habitat provisions, and a highly nutritious food source. It is one of the few trees able to thrive in the harsh acidic and flooded soils of the swamps, earning a role as a keystone species. Humans have also relied on the palm and its aguaje fruits for hundreds of years, utilizing every part of the tree from the roots to the leaves. The fruit is commonly eaten raw or turned into jellies, popsicles, ice cream, or drinks. The fruit is considered a superfood, with the highest known level of vitamin A, as well as Vitamin C, E, antioxidants, and phytoestrogens. Aguaje has traditionally served a medicinal role- energizing, focusing, renewing, balancing hormones, and boosting women’s fertility and figures. These medicinal roles, along with the economic, social, and ecological services the tree provides, have earned it the name “the tree of life.”

Nomenclature
Aguaje is the tree’s widespread name in Peru, where it grows abundantly. Along with being the name of the itself tree as well as the fruit it bears, the term has several meanings in Spanish related to water. One of these is “watering place,” or the tree’s swampy habitat across the Amazon. The palm is also referred to as buriti in Brazil, moriche in Colombia and Venezuela, and ita in Guyana (Koolen et al., 2018).
Aguaje was the first Amazonian palm tree scientifically published in 1781 as Mauritia flexuosa in Carl von Linnaeus’s Supplementum Plantarum (Koolen et al., 2018). Flexuosa means “full of bends” in Latin, which is most likely referencing the morphology of the tree- the way the palm leaves bend out away from the center. Alexander von Humboldt, an explorer considered the father of ecology, referred to aguaje as “the tree of life” in his manuscript about his expeditions through South America (Koolen et al., 2018). He had observed first hand the major ecological, economic, and social roles that it played, giving life to the surrounding ecosystem, the people, and their cultures. Today, the aguaje fruit is commonly consumed by women to improve fertility and is believed by locals to refresh and renew the body, giving further meaning to the term “tree of life.”

Evolutionary History
Mauritia flexuosa and its closest relative, Mauritiella armata, belong to the family Arecaceae, the palms. High antioxidant levels and the presence of flavonoids were observed in the extracts of all the analyzed structures of these two species. Together they form a clade with Lepidocaryum tenue, classified as the subtribe Lepidocaryeae (Freire, J.A.P., 2016).
Populations of M. flexuosa throughout the Neotropics rose and fell during the glacial cycling of the Quaternary glaciation, the last 2.6 million years of Earth's history. Its fossil pollen record has shown populations of high genetic diversity persisting in central Amazonia, these regions serving as historical refugia for them. The area surrounding the Roraima State of Brazil is thought to be the center of origin for M. flexuosa (Rull, 1998; van der Hammen, 1957), in transition areas between dense forests and more open areas. This region is considered an important source of establishment of different plant species that expanded their distribution during the Pleistocene geological time period. Gene flow between M. flexuosa populations follows river basins and the direction of their water flow, suggesting the rivers’ importance for seed dispersal and determining patterns of genetic variation (Sander et al., 2018).
Aguaje has become a staple calorie source for countless organisms in its environment because of its high fruit yield and the fruit’s high nutrient content. The tree provides other exceptional ecosystem services, including water quality improvement and carbon storage, which have helped it evolve into a central part of the niche ecosystem, the aguajales. It has been termed a “hyperdominant” species because population densities are 5x higher than expected, which is uncommon in an Amazon ecosystem (Steege et al., 2013). M. flexuosa is one of the few species in the Amazon with the ability to dominate and monopolize an ecosystem, being the closest the Amazon comes to a monoculture. The aguaje tree has adapted to endure harsh circumstances deemed unacceptable by most other trees. The trees have an extensive root system, which allows them to grow in high acidity soils with bad drainage, such as flooded swamp areas and peatlands (Sander et al, 2018).

Botanical Aspects (Vegetative and Reproductive)

Mauritia flexuosa, part of the Arecaceae family, is a palm tree that can grow up to 35 meter in height and 35 cm in diameter. Adult specimens are polygamous, with male, female and bisexual inflorescence on separate specimens, making it difficult to differentiate before any flowering occurs. Only the "females produce the fruits, with the pollination of the male flowers. There are an average of 10-25 costapalmate leaves, 6 meter in length, with a petiole of 1.6-4 meters in length. Its leaves compose the spherical crown of the palm tree. There are an average of 8 inflorecensce each 2-3 meter long, each containing approximately 115 flowers, which adds up to 45,000 flowers per blossom The fruit is characterized by its ellipsoid oval-shaped drupe covered by red or dark red-coloured cornea scales. It is between 5-7cm long and 4-5 cm in diameter. Both female and male agauje trees reach flowering at the age of 7-8, but only start producing commercial yields after 2 decades. (Koolen et al.,2018)(Vasquez et al., 2008)(Santos, 2005)
Ecology
M. flexuosa is the most abundant palm tree in South America, dominating low-lying flooded areas of the Amazon- unique for such a biodiverse environment. These “homogenous forests” span millions of hectares and play many important ecological roles (Koolen et al., 2018). Because of their abundance, with certain locations having higher than 1000 per square kilometer, the palms have become a keystone species with entire ecosystems evolved to depend on them. They serve as sources of carbon exchange and sequestration, and of food and habitats and for a wide variety of species. M. flexuosa habitats are associated with high species richness, particularly of fish, frogs, and toads (Virapongse et al.).
The tree is deciduous, with separate plants having staminate and pistillate flowers. After about a decade of maturation, a female aguaje palm produces 4–8 racemes, each bearing 500-2000 fruits. The yellowish flowers appear from December to April, bearing ripe fruit in June. Reported fruit yields range from 6.5-9.07 mt/ha. The high yield production of extremely nutritious fruit allowed for a wide range of wildlife, including primates, ungulates (hooved mammals), reptiles, birds, and even fish, to depend upon this species as a food source (Virapongse et al., 2017). Fallen stems or old trees serve as a substrate for the palm beetle, whose larvae are a delicacy in local communities. Seeds are dispersed by the animals eating the aguaje fruit and by water, as the fruits float. Aguaje fruit is a critical food source for birds during the dry season because the palm swamps stay flooded, while other fruits are scarce due to the dry environment. (Manzi et al., 2009). Individuals can grow up to 35m and live for over 150 years, providing many ecological services throughout their lifespan and nurturing the wellbeing of an entire ecosystem.
Cultivation Practices

Although the aguaje palm has the ability to grow where many other food trees cannot, the difficulty of identifying females, the 10 years it can take to produce any fruit at all, and its blockage of sunlight for understory crops makes it a difficult crop to integrate into farms, with limited success of past cultivation.
The fruit is typically harvested from wild rainforests and not generally exported or sold outside the local regions where it grows. Currently, fruits are harvested by rural communities and sold to middle men and distributors in more populated areas. It is mostly processed in traditional household ways, other than oil extraction for a few cosmetics companies.The harvest season is from December to July, with a maximum yield in April (Santos, 2005). The fruit is collected by either climbing the palm and cutting the branches, or by cutting down the entire tree. Due to increasing demand and large scale destructive harvesting methods, the palm has depleted throughout the Peruvian Amazon (Brokamp et al. 2011). There are now more efforts for sustainably Managing Amazonian palms for community use, and Several governmental organizations have begun promoting aguaje plantations for local markets as a cash crop. El instituto de investigación de la amazonía peruana (IIAP) has been working on selectively propagating individuals to create faster bearing semi-dwarf varieties for agroforestry. Camino Verde’s Reforestation Center in Madre de Dios, Peru, have vouched for aguaje’s versatility and functionality in multi-crop agroforestry systems. However, planting aguaje remains a foreign concept for most who live in the region. The fruit symbolizes the wild for them- it is gathered, not cultivated.

Nutritional Value and Consumption
The part of the plant most commonly consumed is the pulp. It can be eaten raw or made into products such as juice, jelly, ice cream, and popsicles. The stipe is starchy and is used to make flour for bread, while the sap can be fermented for palm wine. Aguaje is the richest known natural source of vitamin A, which helps maintain healthy teeth, tissues, bones, and skin (Manzi et al., 2009). It is also high in Vitamin C and E, antioxidants, fatty acids, and phytoestrogens, which reinforces the symbolic identity of the plant.
Along with its high nutritional content, aguaje is valued for its traditional medicinal role. It has been used as a hormonal regulator and to improve chronic fatigue, mental fog, and vision. The phytoestrogens in the fruit are naturally occurring compounds that mimic estrogen in the body. Many women eat aguaje to help boost their fertility or figure.
The entirety of the palm is used for different purposes, from its roots to its leaves. Oil is extracted from the fruit and used to heal wounds, bites, and burns, as well as to protect the skin from the sun (Koolen et al., 2012). Triacylglycerols and fatty acids are the main compound classes of the oil, which keeps the skin looking youthful by moisturizing and supporting collagen production.
Historical Beliefs of Mauritia flexuosa

The Maijina, Is one of Peru’s indigenous groups that bases their culture around the Amazon basin near the city of Iquito. They are deeply rooted within the habitat and ecology of Mauritia flexuosa, holding vast ecological knowledge. The Maijina has historically inhabited the Yanayacu River, the Napo and Putumayo rivers, and as a result have formed folklore in connection to 7 major aguajales/swamps areas. One of these aguajales, which the Maijina have named Gogobai ni cuadu, is believed to be associated with a malevolent female spirit, Gogobai. It is widely believed by the community that Gogobai reside within the aguajales, but on occasions will leave the jungle and wander around the communities, only to prey on children, lead them back to the aguajales and eat them. Gogobai appears to at times be invisible and when abducting children, take on the form of a woman similar to the mother of the child. This traditional Maijuna story highlights the dangers that Gogobai pose to Maijuna children, however, this does not prevent the entry of all family members to aguajales for the collection of the aguaje fruit. (Story information retrieved from consultant testimony in Gilmores, 2013)

Expansion and Domestication History
Although pollen records corresponding to Mauritia flexuosa within South America reveal that this palm tree has been present for the past four ice age cycles, it was held in small but widely distributed regions across the Amazon (Rull and Montoya, 2014). These sites likely restricted Mauritia flexuosa to small regions with favorable climate conditions for growth, often referred to as microrefugia. (Rull and Montoya, 2014). Mauritia flexuosa only expanded its domain with climate shift during the Holocene nearly 2000 years ago, when it underwent a remarkable expansion in most parts of northern South America (Montoya et al., 2009, 2011). This expansion from a series of small sites to what is now the most abundant palm tree across the Amazon, leads to the question of how or who instigated such an expansion. Though it has been theorized that humans might have domesticated and cultivated Mauritia flexuosa at large scales in the past (Triana & Molina, 1998), no conclusive evidence has been presented for this case. In answering the mystery that is the expansion of Mauritia flexuosa, through a genetic survey of Mauritia palm communities across several river sites in the amazon, it was found that the direction of gene flow between populations fell in line with the direction of river basin from which they grew from (Sanders, 2018), pointing to the realization of seed dispersal as the main contributing factor for the expansion of Mauritia flexuosa along river streams after the ice age.

Human Experience

Today Mauritia flexuosa palm has substantial economic importance, the aguaje fruit is regarded as an amazonian staple for its unique flavor, abundance and nutritional value. Sellers, known as "rematistas" in the area, purchase the fruit, mainly by women (single, widowed, divorced, etc.) from (15 to 82) years old. These merchants sell the fruit as mashed aguaje, unripe fruit, and processed items like popsicles at the main marketplaces, minor markets, and factories. For this reason, the fruit has gathered large popular and commercial diversity, often consumed raw in beverages, ice cream, frozen and much more. Iquito, the biggest peruvian city within the Amazon basin, has been estimated to consume 148.8 metric tons of aguaje fruit per month, which when compared to traditional methods of harvesting includes the cutting of adult female trees (Delgado, 2007). This growing demand for the fruit is not unique to Iquito, in other amazon region such as Loreto (Peru) and Maranhao (Brazil) the growing demand for agauje products have led to the overall overexploitation of aguajales (Sampaio et al. 2012). The economic incentive for cutting down aguaje palms threatens the communities who depend on the resources.
Aguaje is widely used by the communities along the rivers and swamps and has been referred to as the “tree of life”, with no small title, aguaje provides an array of resources, services and benefits as a daily food staple (Barros & Da Silva, 2013). In Brazil, the riverine people hold extensive knowledge of the palm tree, with their own cultural management practices for agaugales in order to harvest the fruit in a viable way to prevent the cutting of the palms (Sakai, 2017). Additionally, there seems to be strong generational ties within the riverine people of Brazil, where the harvesting of the aguege is done collectively by all members of the family, incorporating the young in such a manner that the traditional knowledge for harvesting gauge is appropriate by the next generation (Barros, 2021).
In Peru, where aguaje is the name for Mauritia flexuosa, its name is associated with the presence of water “agua”. There regions characterized by the growing of aguaje palms near bodies of water, either swamps, rivers of wetlands, are referred to as "Aguajales” and their associated resources have been and continue to be culturally significant for its inhabitants (Gilmore, 2013). Not only does aguaje mean a source of nutrition for humans but for the series of organisms that also rely on the fruit of aguajales for food. Approximately 20 distinct animal species consume Mauritia flexuosa fruit and are hunted in Mauritia flexuosa palm swamps, according to experts. There are 13 mammalian species, 6 bird species, and 1 reptile species, all of which are consumed and somewhat more than half (55%) are sold as game meat. (Gilmore, 2013). And though a large chunk of people depend on aguajales as a source of income, another large number of people who inhabit the areas near the rivers make most of their financial income from raising cattle and livestock, as well as hunting near these waterways (Barros, 2021).

Works Cited

Barros, F. B., de Sousa, F. F., de Andrade, J. P., Ramos, F. M. & Vieira-da-Silva, C. Ethnoecology of miriti (Mauritia flexuosa, L.f.) fruit extraction in the Brazilian Amazon: knowledge and practices of riverine peoples contribute to the biodiversity conservation. J Ethnobiol Ethnomed 17, 3 (2021).

Brokamp, G., N.Valderrama, M. Mittelbach, C. A. Grandez R., A. S. Barfod, and M. Weigand. 2011. Trade in Palm Products in North-Western South America. Botanical Review 77(4):571-606.

Castro, D. A. M. et al. Genetic structure analysis of Mauritia flexuosa natural population from the Lençóis Maranhenses region using microsatellite markers. Sci. agric. (Piracicaba, Braz.) 79, (2021).

Gilmore, M. P., Endress, B. A. & Horn, C. M. The socio-cultural importance of Mauritia flexuosa palm swamps (aguajales) and implications for multi-use management in two Maijuna communities of the Peruvian Amazon. J Ethnobiol Ethnomed 9, 29 (2013).

Koolen, H.F. da Silva, F., da Silva, V.S.V., Paz, W., Bataglion, G. 2018. Buriti Fruit- Mauritia flexuosa. Academic Press. 61-67.

Manzi, M. & Coomes, O. T. Managing Amazonian palms for community use: A case of aguaje palm (Mauritia flexuosa) in Peru. Forest Ecology and Management 257, 510–517 (2009).

Melo, W. A., Freitas, C. G., Bacon, C. D. & Collevatti, R. G. The road to evolutionary success: insights from the demographic history of an Amazonian palm. Heredity 121, 183–195 (2018).5.

Pereira Freire, J. A. et al. Phytochemistry Profile, Nutritional Properties and Pharmacological Activities of Mauritia flexuosa. Journal of Food Science 81, R2611–R2622 (2016).

Rull, V. & Montoya, E. Mauritia flexuosa palm swamp communities: natural or human-made? A palynological study of the Gran Sabana region (northern South America) within a neotropical context. Quaternary Science Reviews 99, 17–33 (2014).

Sander, N. L. et al. Rivers shape population genetic structure in Mauritia flexuosa (Arecaceae). Ecology and Evolution 8, 6589–6598 (2018).

Santos, 2005. L.M.P. Santos Nutritional and ecological aspects of buriti or aguaje (Mauritia flexuosa Linnaeus filius): A carotene-rich palm fruit from Latin America. Ecol. Food. Nutr., 44 (2005), pp. 345-358

Silva, R. S., Ribeiro, L. M., Mercadante-Simões, M. O., Nunes, Y. R. F. & Lopes, P. S. N. Seed structure and germination in buriti (Mauritia flexuosa), the Swamp palm. Flora - Morphology, Distribution, Functional Ecology of Plants 209, 674–685 (2014).

Steege, H. T., Pitman, N. C., Sabatier, D., Baraloto, C., Salomão, R. P., Guevara, J. E., ... Silman, M. R.. (2013). Hyperdominance in the Amazonian tree flora. Science, 342, 1243092.

Virapongse, A., Endress, B. A., Gilmore, M. P., Horn, C. & Romulo, C. Ecology, livelihoods, and management of the Mauritia flexuosa palm in South America. Global Ecology and Conservation 10, 70–92 (2017).