Colombian plants in Kew's Economic Botany Collection

Catalogue of Useful Plants of Colombia Edited by Raquel Negrão Alexandre K. Monro Carolina Castellanos-Castro Mauricio Diazgranados Catalogue of Useful Plants of Colombia Catalogue of Useful Plants of Colombia Edited by Raquel Negrão Alexandre Monro Carolina Castellanos-Castro Mauricio Diazgranados Kew Publishing Royal Botanic Gardens, Kew © The Board of Trustees of the Royal Botanic Gardens, Kew 2022 Text © The Board of Trustees of the Royal Botanic Gardens, Kew, and the Instituto de Investigación de Recursos Biológicos Alexander von Humboldt Images © The Board of Trustees of the Royal Botanic Gardens, Kew, unless otherwise stated The authors have asserted their right to be identified as the authors of this work in accordance with the Copyright, Designs and Patents Act 1988. All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form, or by any means, electronic, mechanical, photocopying, recording or otherwise, without written permission of the publisher unless in accordance with the provisions of the Copyright Designs and Patents Act 1988. Great care has been taken to maintain the accuracy of the information contained in this work. However, neither the publisher nor the authors can be held responsible for any consequences arising from use of the information contained herein. The views expressed in this work are those of the authors and do not necessarily reflect those of the publisher or of the Board of Trustees of the Royal Botanic Gardens, Kew, or Instituto Humboldt. First published in 2022 by Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK, in association with the Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Calle 28A #16-29, Bogotá, Colombia. www.kew.org ISBN 978-1-84246-774-9 eISBN 978-1-84246-775-6 Distributed on behalf of the Royal Botanic Gardens, Kew in North America by the University of Chicago Press, 1427 East 60th Street, Chicago, IL 606037, USA. British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library. Design: Nicola Thompson, Culver Design Production Manager: Jo Pillai Proofreading: Sharon Whitehead Printed in Great Britain by Halstan & Co Ltd For information or to purchase all Kew titles please visit shop.kew.org/kewbooksonline or email publishing@kew.org For Instituto Humboldt publications visit http://repository.humboldt.org.co/ Kew’s mission is to understand and protect plants and fungi, for the wellbeing of people and the future of all life on Earth. Kew receives approximately one third of its funding from Government through the Department for Environment, Food and Rural Affairs (Defra). All other funding needed to support Kew’s vital work comes from members, foundations, donors and commercial activities, including book sales. Corresponding author: Mauricio Diazgranados, Herbarium, Royal Botanic Gardens Kew, Richmond TW9 3AE, United Kingdom E-mail: M.Diazgranados@kew.org Front cover photo credit: Néstor Plata © RBG Kew; back cover background photo credit: Mauricio Diazgranados © RBG Kew; back cover plants photo credits: Lupinus bogotensis (Mauricio Diazgranados © RBG Kew), Espeletia jimenez-quesadae (Mauricio Diazgranados © RBG Kew), Typha domingensis (Ori Fragman-Sapir ©), Passiflora cumbalensis (David Granados © RBG Kew), Leopoldinia pulchra (Mauricio Diazgranados © RBG Kew), Cardiospermum halicacabum (Wolfgang Stuppy © RBG Kew), Anacardium occidentale (Wolfgang Stuppy © RBG Kew), Draba litamo (Mauricio Diazgranados © RBG Kew), Datura metel (Laura Kor © RBG Kew), Cattleya trianae (Mauricio Diazgranados © RBG Kew) Funded by: Implemented by: Contents Preface 7 Executive Summary 11 Foreword 15 Acknowledgements 17 List of Contributors 19 Chapter 1 The Useful Plants and Fungi of Colombia (UPFC) project: delivering botanical knowledge to support conservation and sustainable development 21 Chapter 2 Assessing extinction risk coverage and identifying geographical hotspots to guide conservation action for Colombia’s useful plants 35 Chapter 3 Endemic Medicinal Plants of Colombia 49 Chapter 4 The hidden food basket of Latin America: an overview of Colombian edible plant diversity and its distribution 63 Chapter 5 Insecticide Plants of Colombia 79 Chapter 6 Useful Plants of the Colombian Amazon 93 Chapter 7 Colombian plants in Kew’s Economic Botany Collection 103 Chapter 8 Bioeconomy: a sustainable and responsible option for making the most of Colombian plant resources 113 Chapter 9 Sustainable value chains and development pathways for natural ingredients in Colombia: the case of naidí (Euterpe oleracea Mart.) 121 Chapter 10 A taxonomic summary of useful plants in Colombia 135 Chapter 11 Notes on the geographic distribution of the useful plants of Colombia 149 Chapter 12 Annotated checklist of useful plants of Colombia 165 Checklist 171 Laminae 475 Index of Families 529 Index of Genera 531 Index of Synonyms 541 Index of Vernacular/Common Names 989 Espeletia nemekenei Cuatrec (Asteraceae =Compositae) used as Medicinal in Colombia. 6Mauricio Diazgranados CATALOGUE OF USEFUL PLANTS OF COLOMBIA Preface Raquel Negrão, Alexandre Monro, Carolina CastellanosCastro, Mauricio Diazgranados “Existence begins as sunlight Much of life on earth is a solar being From the plant that drinks its golden rays To make the sugars that nourish all animal life We are all powered by energy from the sun”1 (…) “In this room, the ingredients of life are made possible In a ceaseless cycle of transformation” 1 (…) “In a glade and in a thicket In a desert and woodland They are part of a legacy that imbricates all life on earth: Their exhale with your inhale Their inhale with your exhale”2 (…) “We are all environments Walking oceans Teeming with life United by an unending rhythm That blurs the boundaries between all breathing bodies And submerges us in torrent of being”3 Excerpts from a poetic narrative by Daisy Lafarge presented at Observations on Being (developed by Marshmallow Laser Feast for Coventry City of Culture 2021): 1 “The World’s First Breath”; 2 “From Breathing Cell to Breathing Planet”; 3 “The Tides within us”. Daisy Lafarge is a poet from the UK and studied at the universities of Edinburgh and Glasgow. Her collection of poetry Life Without Air (Granta Books) was published in 2020 and shortlisted for the T.S. Eliot Prize 2020. Plants and fungi fulfil our everyday needs, and it depends on us to use them sustainably. The consequence of nature’s entangled evolution is that our future, from our first to our last breath, depends on all life forms on the planet and in this network, plants and fungi are found at the base. They offer us everything we need to survive and to enjoy life with comfort and joy: from the basic needs of oxygen, food, fuels, clothes, housing, water cycling, and erosion control, to the thermal comfort of shade, medicines for the body and mind, gratifying properties such as fragrances, the aesthetic and mental health values of flowers and landscapes, the entertainment or delight provided during cultural-social celebrations, spiritual tools for use in rituals (such as aguardiente, tobacco, ayahuasca, mambe, rapé) and daily pleasures such as a coffee or a chocolate bar. There are also several examples of how plants have played a remarkable part in our cultural and social history, influencing largest-scale human population shifts, and allowing written as well as visual and oral communication. The primitive hardware for much of our intellectual development was made of natural resources: clay tablets, animal skin, and plant fibres especially have been used for thousands of years. Paper or paper-like materials made from plants were developed mainly from papyrus, rice, insect-based silk, hemp, bamboo, Kapa bark cloth (paper mulberry), and cotton-based materials, providing a portable and durable means to “spread the word” and allowing shifts in societal organisation. After centuries of traditional travellers disclosing paper producing cultures (from Egypt, China, passing by the Middle East and entering Europe from the Italian city of paper, Fabriano), vegetable-based writing materials (e.g., cotton sheets) started to be transformed on a much larger scale. Eventually, printing techniques were upgraded for the large-scale production of books, magazines, and newspapers, facilitating the diversification of communication, languages, knowledge, and art. From the same production centre and using the techniques developed to transform cotton sheets, the Italians also developed a more sophisticated, “lighter!” monetary system. This was the first and still the main producer of paper money (it is worth visiting the paper museum [Museo della Carta e della Filigrana] in Fabriano). You may argue that we are already in the age of digital money and “intelligent” virtual systems. Still, CATALOGUE OF USEFUL PLANTS OF COLOMBIA 7 PREFACE we would argue that we are not even partially independent of plant-based resources as primary resources for a continuous evolutionary process. Amid the biodiversity crisis, more than ever, our survival requires that we sustain the natural world and plants more than our short-term wealth (especially the wealth of those who are already economically comfortable). In other words, our origin, persistence, and future on this planet rely fully on plants and fungi, which must be protected. Conservation and the sustainable use of biodiversity necessarily incorporates rural livelihoods. Since 2015, the United Nations has been implementing the 2030 Agenda for Sustainable Development, which comprises 17 Sustainable Development Goals and 169 targets that complete the Millennium Development Goals. The 2030 Agenda aims to integrate and balance the three dimensions of sustainable development (economic, social, environmental) and to reinforce human rights. The inclusion of livelihoods in conservation is a particularly important development in the post-2020 Global Biodiversity Framework, as it defined new global targets and actions to support a sustainable, healthy future for people and nature”. Also, expressive activist movements responding to climate change, extinction, and ecocide, have increased the accumulation and diffusion of scientific knowledge and public awareness. The post-2020 Global Biodiversity Framework also emphasises the key role of indigenous and local communities using traditional knowledge to protect nature and develop sustainable strategies. This book is our contribution to dealing with one of the challenges of the Anthropocene Age, how to use plants and plant-dominated landscapes sustainably. RBG Kew’s Science Strategy 2021–2025 “sets out a series of ambitious commitments to urgently help stop biodiversity loss and accelerate understanding of the potential of plants and fungi to help address challenges such as food insecurity, climate change and deforestation”. Recent publications (Bridle, 2020; Marteau et al., 2021) have shown that a quarter of greenhouse gas emissions come from food and transport. Behaviour changes by individuals, commercial entities and policy makers that incentivise people to eat more locally grown produce and have more meat-free days could help us to fight climate change. Our Catalogue of Useful Plants of Colombia demonstrates the great diversity of plants available as local foods, medicines, fuel sources and, materials, valuable resources that could help us to achieve net zero in all domains. It will hopefully contribute to Colombia’s ambitious plan to develop a green economy working with traditional communities and local initiatives in an inclusive approach that, especially considers the voices of traditional communities, and their needs and rights. Our more practical goal with this publication is to inform you about the richest natural resources available, thereby potentially helping us to achieve a better society in which natural resources are valued and used and avoiding sustainably further habitat and biodiversity loss. We want to offer you (scientists, students, and academics in general) a comprehensive guide to the sample of this biodiverse beautiful planet that is found in Colombia. With 8 almost 29,000 plant species and thousands more yet to be described, Colombia is one of the most biodiverse countries in the world. The Catalogue of Useful Plants of Colombia showcases a successful partnership between Colombia and the UK (see Chapter 1), which has enabled us to document the strengths (on uses, biological diversity, geographical distributions, and business interests) as well as a lack of knowledge about less well known uses and species conservation (chapters 2, 3, 4, 5, 6, 10 and 11). Chapters on the history of Kew’s Economy Botany Collections (chapter 7) and the development of bioeconomy in Colombia (chapter 8) look to the past to understand the present and point us towards better future, as does the successful example of ‘Colombia’s Bio’ – green growth development model in the form of naidí (Euterpe oleracea) (chapter 9). Finally, we explain how the checklist was assembled and what kind of resources are presented for all of the 7,472 species of useful plants in Colombia (chapter 12). Following the checklist, there is a section showing 500 images as examples of species and uses. USING THIS CATALOGUE AND THE COLPLANTA ONLINE PORTAL A large amount of the information on Colombia’s plant diversity present in this Catalogue is aimed mainly at an academic audience. To access additional information, users can follow the hyperlink on the species name, read the species QR code, or type the scientific, common name, genus, family, or descriptor of a plant of interest or its uses in the search bar on the home page of the ColPlantA portal (http://www.colplanta.org) (see Chapter 12). The search results will show the users relevant plant species profiles, which they can further explore. Each plant profile encompasses the classification of the plant, the status of its name (whether accepted or considered a synonym), and information on habit, description, ecology, distribution, uses, common names, and conservation status (see Chapter 12). Images of wild plants and herbarium specimens are also featured where available, alongside additional information about the species. The ColPlantA portal has advanced search and filtering options that allow users to build customised queries based on their needs. Both the Catalogue of Useful Plants of Colombia and the ColPlantA portal are open-access resources built by experts, using accredited information on the useful plants of Colombia. Both resources are designed to disseminate information on useful plants to the community, supporting research, learning, development, and innovation for sustainable growth, economic development, and well-being. The basic data can be used as a source of information about Colombian plants. Also, both resources represent essential baseline information that will underpin conservation actions, environmental management, legislation, and policy. Although so many botanical and conservation tasks appear to be carried out by scientists, ecologists, government bodies, and non-governmental organisations, there is a CATALOGUE OF USEFUL PLANTS OF COLOMBIA PREFACE multitude of key people behind the scenes, involved in documenting, communicating, and conserving plant diversity, and playing single or multiple roles: from the teachers who show children, for the first time, bean seeds germinating on cotton, to horticulturalists, gardeners, reserve managers, field biologists and helpers, data compilers, curators and many others. There are still many unknown challenges related to climate change. Very painfully, the Covid-19 pandemic has shown that it is better for human societies to be more collaborative instead of competitive, sharing benefits and commitments, accessing the diverse knowledge available, and taking emergency actions to protect all living beings when needed. References Bridle SL (2020). Food and climate change without the hot air: change your diet: the easiest way to help save the planet. UIT Cambridge Ltd, 256 p. Marteau TM, Chater N & Garnett EE (2021). Changing behaviour for net zero 2050. BMJ, 375, n2293. https://doi.org/10.1136/bmj.n2293 CATALOGUE OF USEFUL PLANTS OF COLOMBIA 9 Orthrosanthus chimboracensis (Kunth) Baker (Iridaceae). In Colombia, this species presents Medicinal, Material and Environmental uses. 10 Mauricio Diazgranados CATALOGUE OF USEFUL PLANTS OF COLOMBIA Executive Summary Colombia is the second most biodiverse country in the world. Its biological resources are key assets in the country’s efforts to transition into a more sustainable, competitive, and inclusive economy. However, to increase the contribution of the bioeconomy to national development, a crucial step is to consolidate the current state of knowledge of the rich diversity of useful plants in the country. Arising from the collective efforts of the Alexander von Humboldt Biological Resources Research Institute (Colombia) and the Royal Botanic Gardens, Kew—under the Useful Plants and Fungi of Colombia Project (UPFC)—this Catalogue is presented as a source of baseline information about the great diversity of useful plant species in Colombia. Before this project, Colombia did not have a updated national list of useful plants, with only a few lists of species by use category (i.e., medicines, edible plants, and fibres), often with conflicting taxonomy and incomplete geographic coverage, being available. The Catalogue presents a comprehensive panorama, including the results of studies developed across the country by several scientific groups from Colombia and the UK. The substantial progress, represented by this catalogue, resulted from a successful partnership between Colombia and the UK, showcased in the introductory chapter (Chapter 1). The authors highlight the achievements of the Useful Plants and Fungi of Colombia (UPFC) project, which has provided a framework to develop and promote a market for useful indigenous species and their high-value products, whilst protecting the surrounding natural resources. With a multinational team, the project compiled and generated knowledge on over 36,000 species of plants and fungi, and developed pathways for tackling socio-environmental challenges, contributing to Colombia’s green transformation. To date, it has produced over 100 outputs, including books, booklets, scientific journal publications, technical reports, websites, online portals, and educational tools. Drawing on the Checklist of Useful Plants of Colombia (CUPC), Chapter 2 proposes conservation actions based on (1) taxonomic gaps in extinction risk assessments and (2) geographic hotspots for native useful plants of Colombia (UPC). From the analysis, 544 species of conservation concern—threatened with extinction and/or endemic to Colombia—were identified. However, the analysis also underlined that 45% of the native useful plants of Colombia lack extinction risk assessments. The proportion of unassessed species was significantly higher for 13 of the 256 useful plant families, results that can be used to prioritise extinction risk assessments. There are also variations between categories of uses, and 78 units of analysis (covering 89,901 km2) were identified as hotspots for native useful plants of Colombia. In this chapter, the authors discuss how the hotspots can contribute to the identification of Important Plant Areas in Colombia, as well as the importance of stakeholder engagement to develop and implement conservation actions and sustainable uses of plants. One of the main findings of this checklist is that medicinal species represent the most diverse group. Chapter 3 presents a long-term study (from 2014 to 2021) on plant species that are used as medicines in Colombia and their relationship with traditional, scientific, and technological knowledge. Before the checklist was compiled, the authors had documented a total of 3,005 medicinal plants used in Colombia, of which 1,719 are native (including 204 endemics), 558 exotic, and 728 species of uncertain geographical distribution. The lack of knowledge on the conservation status and sustainable uses of most of the Colombian endemic medicinal species was verified: only a few restricted species had sufficient information available to allow their categorisation. Chapter 4 addresses the second largest use category, the edible plant species, and their conservation status. The chapter concludes that the high diversity of Colombian edible plant species is not currently matched by adequate in-situ and ex-situ conservation actions. Chapter 4 also identifies significant knowledge gaps regarding the conservation status of some of the most important edible plant genera. The biogeographic analysis highlights the ‘Andes’ and ‘Páramo’ bioregions as diversity hotspots for edible plants, emphasising the urgent need for their CATALOGUE OF USEFUL PLANTS OF COLOMBIA 11 EXECUTIVE SUMMARY preservation. The authors concluded that further research is critical to understand the complex interactions between people, food, and the natural environment, whilst at the same time, conservation efforts need to consider the intrinsic value of edible plant diversity and the full potential for socio-economic development at the local and national levels. A special category of use is presented in Chapter 5, which reports an insightful long-term study on insecticide activity. To date, no publication has detailed this category of use in Colombia and besides identifying and characterising the plant species used as insecticides, this chapter also assesses the state of traditional and scientific knowledge on these species. The authors demonstrate that 632 species used as ‘insecticides’ were identified in Colombia. For none of these species, however, is there sufficient scientific knowledge to underpin their application as natural insecticides. Chapter 6 presents a synthesis of the current state of knowledge about the Useful Plants of the Colombian Amazon, based on information gathered from field expeditions and records added to the Colombian Amazon Herbarium (COAH) database managed by the Amazonian Scientific Research Institute SINCHI. Useful plants were classified into 12 categories of uses, by origin (native or endemic), and according to their extinction risk assessments. In total, 2,286 useful plants were recorded in the Colombian Amazon. Similar to the results found from the CUPC, Chapter 6 found that the medicinal (883) and food (690) categories presented the highest numbers of species documented for the Colombian Amazon. Challenges to achieving the sustainable use of species in this region are discussed from the economic botany perspective. The history of Kew’s Economy Botany Collections is presented in Chapter 7, looking to the past to understand the present. This collection was first founded as the Museum of Economic Botany in 1847. Among a total collection of 100,000 plant raw materials and plant products, 320 specimens originate in Colombia. Recent research has enabled botanical names to be updated and provenance data retrieved through archival research. The Colombian specimens mirror the changing scientific relationship between Colombia and Kew from the mid-nineteenth century to today. For a megadiverse country such as Colombia, sustainable bioeconomy requires a completely different circular way of producing natural products, with intensive use of science to add value to biological resources and biological processes, as is presented in Chapter 8. Basic concepts of circular bioeconomy, value chains and biotechnology are described, and some examples of bioeconomy-focused biotechnological developments from the Research Centre for Agro-industrialisation of Tropical Aromatic and Medicinal Plant Species CENIVAM are presented. The authors concluded that integrating bioprospecting results with those from analytical chemistry and molecular biology (bioactivity assays) led to the detection of promising vegetal species 12 to be used as ingredients for new consumer products. Field projects have established the conditions for biomass production under sustainable agricultural practices, as well as for their rural processing to afford essential oils, extracts, biocontrol solutions, composting material, and biofuel. Multiple challenges still need to be addressed, such as establishing commercial chains with reinvestment into the countryside, the achievement of origin recognition, road improvement, internet coverage, water supplies, and stable legislation. A successful example from Colombia’s Bio program, using the green growth model for the development of a real, sustainable value chain for natural ingredients in Colombia, is the case of naidí (Euterpe oleracea), which is presented in Chapter 9. From semi-structured interviews involving community-based organisations, private companies, NGOs, international organisations and universities, this case study assesses the full range of activities and relationships involved in creating value. The main results highlight that the factors which characterise the value chain consist of trusted relationships between small-scale producers, a low distribution of profits among local collectors, limited profitability supported by external subsidies, and low regional and national demand. The main results suggest that this value chain could be improved in the short- and mediumterm by increasing the capacity of cold chains and storage centres, fostering economic incentives for implementing agroforestry systems, developing strategies for promoting traditional diets by using its natural ingredients, and improving technology and business capabilities to enhance its commercialisation. Chapters 10 and 11 provide descriptive summaries of the taxonomic and geographical distribution of species, based on the uses in the database to produce the checklist of Useful Plants of Colombia (UPC) detailed in chapter 12. Chapter 10 provides notes on the taxonomic coverage of this checklist, highlighting the plant groups richest in useful plant species, categories of use, and phylogenetic spread. Potential taxonomic gaps are also identified. The checklist of useful plants comprises 7,472 species, 2,140 genera, and 258 families, representing all major plant groups. Most of the species are native (78.6%), and 6.4% are endemic to Colombia. At least 14% of the useful species are commercially cultivated, and 8.6% are introduced and naturalised. The families that are richest in useful plant species are Fabaceae, Asteraceae, and Poaceae. The genera that are richest in useful plant species are Solanum, Inga, and Passiflora. The categories of use with the most associated plant species are medicines (5,108 spp.), human food (3,806spp.), and materials (2,363spp.). Chapter 11 characterises the geographic distribution of the species listed in the checklist by describing distribution patterns at varying biological and geographical scales. In total, the plant species with documented uses are represented by 435,230 publicly available georeferenced records. Of the 13 bioregions assessed, the humid forests of the Andes made up the bioregion with by far the largest CATALOGUE OF USEFUL PLANTS OF COLOMBIA EXECUTIVE SUMMARY numbers of records, species, genera, and families. The humid forests of the Amazonia and Caribe followed in terms of species richness. At the regional level, Antioquia has the largest number of useful species, followed by Cundinamarca and Valle del Cauca, whilst at the municipality level, Solano (Caquetá), Manizales (Caldas), and Leticia (Amazonas) report the highest species richness. However, records are unevenly distributed across both taxonomic groups and geographic areas. Within all use categories, the species richness decreased with elevation. Potential drivers, implications and knowledge gaps are presented, and the current understanding of the distribution of useful plants can be used as a baseline to target future research efforts, supporting plant conservation planning and efforts across Colombia. Finally, a comprehensive checklist of the useful plants of Colombia is presented in Chapter 12. The chapter describes how the checklist was built and what kind of resources are presented for all useful species of Colombia. The list includes supraspecific taxa, accepted species and authors, common names, species origin, geographic information (habitats, regions, departments, and elevation range), conservation status (assessments of the accepted name), and level 1 of the category of use. Tracking species as their names change through time and maintaining the link to their uses, distributions, conservation status and common names, is an endless task. This is the reason for having the printed version of the catalogue linked to a dynamic online portal. For full details on the species, readers should scan the QR codes and visit the species profiles in https://www.colplanta.org, which include morphological descriptions, geographic distribution and maps, synonymy, notes on uses, links to herbarium specimens, images of live plants, illustrations, economic botany items and anatomical laminae, bibliography, and additional sources. The checklist will enable you to access its supplementary material, to explore open questions and opportunities to develop innovative ideas on alternative processes for consumption and production, novel technologies, and species conservation, to promote a social and environmental positive impact, and to inspire new generations. We hope that this Catalogue and ColPlantA (as its dynamic resource) will be used by the audience as a useful referential for decision-making, production, and demand building. We expect these resources to meet a broad audience (consumers, interested public, politicians, producers, regulators, scientists, stakeholders, and students), allowing Colombia as a Nation to achieve its Mission on Green Growth, and to use and conserve its natural resources in an integrated, equitable and sustainable way. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 13 Flower of Passiflora ambigua Hemsl. (Passifloraceae) used as human food in Colombia. 14 Mauricio Diazgranados CATALOGUE OF USEFUL PLANTS OF COLOMBIA Foreword Back in 2016, during the State Visit that President Juan Manuel Santos paid to the UK, there was an important event at the Natural History Museum, where HRH The Prince of Wales, President Santos and other high-level UK Government stakeholders discussed how to transform the scientific collaboration between the UK and Colombia. That episode marked the beginning of the UK Government’s Newton-Caldas Fund support to an inspirational programme called Colombia BIO, aimed at understanding Colombia’s vast biodiversity to ignite an unprecedented systematic international scientific collaboration, the epicentre of which was in rural areas of Colombia formerly affected by years of armed conflict. At that event, I met two members of the Kew Science team, Professor Monique Simmonds OBE and Dr Mauricio Diazgranados Cadelo, who have unintentionally contributed to both my professional and personal lives, transforming part of my own relationship with nature. In that meeting, the three of us drafted the first ideas to conceptualise what months later became ColPlantA, a project led by Royal Botanic Gardens, Kew, within the British Council’s Professional Development & Engagement strand of their Newton-Caldas Fund work. This project was carried out in partnership with what we know today as the Ministry of Science and Technology in Colombia, Minciencias. ColPlantA, an online tool to provide open information about Colombian plants and their uses, was developed to complement other work that the Royal Botanic Gardens, Kew was already doing with Colombian scientists and research institutions with the support of the UK Government. Once ColPlantA was established, the scientific team at Kew knew there was still work to do and worked hard to win a new Newton-Caldas Fund grant, administered by the British Council, which was announced during the visit of President Iván Duque to the UK as a guest of the Government, back in 2019. That, in my view, was the origin of the Useful Plants and Fungi of Colombia project. Part of my role at the British Embassy in Colombia is to connect the UK scientific ecosystem with its peer equivalent in Colombia. The Useful Plants and Fungi of Colombia project is a perfect example of how peers exchange knowledge and produce equal partnerships, within an epistemic exercise to put science at the front of a race to act against a changing climate that demands immediate solutions from all of us. Those solutions—we learn from the work of institutions such as RBG Kew, Instituto Alexander von Humboldt and a wide range of Colombia and UK academic institutions—have a lot to do with plants and the ways in which we use them. The more we learn about these uses, actual or prospective, the better equipped we are to play our individual role in solving a collective challenge such as climate change. I feel immensely proud to see research products such as the one you are reading now, and I am hugely grateful for the opportunity I have had to contribute to making this possible, almost literally from the inception stage. In a year when life as we used to know it has been radically changing, I truly believe in science not only as the way to find the answers to complex global development challenges but also as a superpower to transform lives in entire communities. This is perhaps one of the most important benefits of a research product like this. Offering Colombian communities the opportunity to learn about—and sustainably live from—their surrounding nature is, in itself, a paramount achievement that history will put into perspective in the years to come. I celebrate this unique piece of scientific collaboration and I have no doubt about the potential of this work to contribute not only to Colombia but also to the rest of global society. This book is testament to a deep bilateral scientific relationship I have had the immense honour to energise through my work, so I really thank those who made it possible and do hope you enjoy it and find it useful. Luis Calzadilla Waldmann Head of Science and Innovation British Embassy Colombia CATALOGUE OF USEFUL PLANTS OF COLOMBIA 15 Inflorescence of Puya trianae Baker (Bromeliaceae), a species from the páramos with Environmental and Animal Food uses. 16 Mauricio Diazgranados CATALOGUE OF USEFUL PLANTS OF COLOMBIA Acknowledgements This work was supported by a Professional Development & Engagement grant under the Newton-Caldas Fund partnership. The grant is funded by the UK Department for Business, Energy and Industrial Strategy (BEIS) and the Colombian Ministry of Science, Technology and Innovation (MinCiencias) and delivered by the British Council. For further information, please visit www.newtonfund.ac.uk. The Catalogue of Useful Plants of Colombia could have not been realised without the remarkable efforts of many individuals and the support of several institutions. It represents one of the main products of the project Useful Plants and Fungi of Colombia (UPFC), led by the Royal Botanical Gardens, Kew (RBG Kew), with the collaboration of the Alexander von Humboldt Biological Resources Research Institute (Colombia). This book is substantially based on the data from the ColPlantA portal, which is developed and maintained by RBG Kew, with the collaboration of various Colombian partners. The editors and authors are very grateful for the support of the funders regarding the production costs of this publication, RBG Kew for providing the research structure and internal services, the Humboldt Institute for the contribution of data and literature, and Kew Publishing for the final copy-editing and layout. The editors and authors wish to acknowledge their appreciation of all staff, past and present, of the institutions involved in this catalogue, for their dedicated research that resulted in the contents of this work, and of the numerous specialists who contributed to the elaboration and revision of the updated lists, and with valuable information on local and regional uses of the species. Special acknowledgements are due to the team working on taxonomic reconciliation and data management, especially Bob Allkin, Tiziana Cossu, Julia Carretero, Laura Kor, and Benedetta Gori, and to Joaquim d’Souza and Malcolm Stone for providing the PURLs that are used to make the QR Codes. Many thanks to those who were working dedicatedly to provide invaluable support on the compilation of the preliminary lists, references, specimens, common names, and all information related to species and their respective uses and conservation status, in both the UK and Colombia: Laura Green, Ellie Graves, Priscila Reis, Germán Torres-Morales, Alejandra Aguilar-Giraldo, Daniel Jiménez-Pastrana, Lina Isabel Guevara-Ruiz, Henry Agudelo and Fabio Ávila. The authors also wish to thank those who contributed chapters and images, and those who acted as reviewers and made valuable suggestions: Amalia Díaz, Benedetta Gori, Cristina López-Gallego, Dairon Cárdenas, David Hammond, Federico Padilla, Felipe García, Henry Yesid Bernal, Jennyfer Andrea Aldana, Justin Moat, Mark Nesbitt, Nelson Salinas, Phillipa Ryan, Rafael F. Almeida, Steven Bachman, Tiziana Ulian, and William Milliken. We are very grateful to all those who contributed species and field images (Benedetta Gori, Carolina CastellanosCastro, Dairon Cárdenas-López, Elena Stashenko, Elizabeth Hodson, Laura Green, Laura Kor, Mateo Fernández Lucero, Mauricio Diazgranados, Mónica Andrea Flórez Pulido, Priscila dos Reis and Ellie Graves), as well as to the Spatial Analysis team from RBG Kew who produced the species distribution maps available in the ColPlantA species profiles, especially Carolina Tovar, Ian Ondo, and Justin Moat. We are also grateful to Daisy Lafarge for allowing us to use excerpts from a poem she presented at the ‘Observations on Being’ exhibition developed as part of the Coventry City of Culture 2021. Thank you to Kew’s IT team for helping with IT and computational issues, specially Reinis Rozkalns, Erkki Mellin and Chris Soh. We would like to express our deepest thanks to our Project Manager, David Hammond, the Project Officer Kaitalin White and the Communication Specialists Camila Gutiérrez and María del Pilar Mira Pontón, for their enormous support of the Useful Plants and Fungi of Colombia project, which was extended during the entire production process for the publication of the Catalogue. Sadly, one of our contributing authors and colleagues, Dairon Cárdenas-López, passed away during the production of this catalogue. We dedicate this volume to him, his family and his life as a field botanist, scientific researcher, and highly regarded Director of the herbarium at the Amazonian Scientific Research Institute (SINCHI). We celebrate his valuable contribution and dedication to advancing our understanding of the flora of the Colombian Amazon, including the collection of more than 45,000 botanical specimens. Finally, we would like to thank the successive directors and researchers of the RBG Kew, the Humboldt Institute and the Ministry of Environment and Sustainable Development of Colombia, as well as the British Embassy in Colombia and the Colombian Embassy in the UK, for their continuous support. Many other people and institutions gave their support in one way or another to the preparation of this catalogue. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 17 Schoenocephalium teretifolium (Flor de Inírida de Verano), in white sand savannas of Inírida. 18 Mateo Fernandez Lucero CATALOGUE OF USEFUL PLANTS OF COLOMBIA List of Contributors CONTRIBUTORS INSTITUTION Alba Nohemí Téllez A. Alejandra Aguilar-Giraldo Alex Monro Andrés A. Barona-Colmenares Benedetta Gori Carlos Alberto Cortés Carolina Castellanos-Castro Catalina Mesa Sánchez Cristina Lopez-Gallego Dairon Cárdenas-López† Daniel Jiménez-Pastrana David Hammond Elena Stashenko Pontificia Universidad Javeriana Instituto de Investigación de Recursos Biológicos Alexander von Humboldt Royal Botanic Gardens, Kew Instituto Amazónico de Investigaciones Científicas SINCHI Royal Botanic Gardens, Kew Instituto de Investigación de Recursos Biológicos Alexander von Humboldt Instituto de Investigación de Recursos Biológicos Alexander von Humboldt Pontificia Universidad Javeriana Universidad de Antioquia Instituto Amazónico de Investigaciones Científicas SINCHI Instituto de Investigación de Recursos Biológicos Alexander von Humboldt Royal Botanic Gardens, Kew Centre for Research on Tropical Aromatic and Medicinal Plant Species (CENIVAM) – Industrial University of Santander Pontificia Universidad Javeriana Jardín Botánico de Bogotá José Celestino Mutis Instituto de Investigación de Recursos Biológicos Alexander von Humboldt Instituto de Investigación de Recursos Biológicos Alexander von Humboldt Universidad Nacional de Colombia Pontificia Universidad Javeriana Royal Botanic Gardens, Kew; University College London Universidade de São Paulo Royal Botanic Gardens, Kew Royal Botanic Gardens, Kew Royal Botanic Gardens, Kew Royal Botanic Gardens, Kew Instituto de Investigación de Recursos Biológicos Alexander von Humboldt British Embassy Colombia Instituto de Investigación de Recursos Biológicos Alexander von Humboldt Royal Botanic Gardens, Kew Royal Botanic Gardens, Kew Royal Botanic Gardens, Kew Royal Botanic Gardens, Kew Instituto Amazónico de Investigaciones Científicas SINCHI Instituto Amazónico de Investigaciones Científicas SINCHI Royal Botanic Gardens, Kew Royal Botanic Gardens, Kew Royal Botanic Gardens, Kew Elizabeth Hodson de Jaramillo Fabio Ávila Felipe García Germán Eduardo Torres-Morales Henry Agudelo Henry Yesid Bernal M Hernando Echeverri-Sanchez Jennyfer Andrea Aldana Mejía Joaquim de Souza Julia Carretero Kaitalin White Laura Kor Lina Isabel Guevara-Ruiz Luis Calzadilla Waldmann Mabel Tatiana Rojas María del Pilar Mira Mark Nesbitt Mauricio Diazgranados Natalí Sánchez-Garzón Nicolás Castaño-Arboleda Nórida Marín-Canchala Raquel Negrão Tiziana Cossu Tiziana Ulian CATALOGUE OF USEFUL PLANTS OF COLOMBIA 19 Brosimum alicastrum Sw. seeds. Néstor Plata Chapter 1 The Useful Plants and Fungi of Colombia (UPFC) project: delivering botanical knowledge to support conservation and sustainable development Mauricio Diazgranados1*, David Hammond 1, Tatiana Rojas2, Kaitalin White1, María del Pilar Mira1, Carolina Castellanos-Castro2, Camila Gutiérrez1 & Tiziana Ulian1 1 2 Royal Botanic Gardens, Kew Instituto de Investigación de Recursos Biológicos Alexander von Humboldt *Corresponding author: m.diazgranados@kew.org Keywords: bioeconomy, fungi uses, green growth, plant uses, sustainable development ABSTRACT Colombia ranks second in the world in biodiversity and is recognised as one of the most ethnically diverse countries. Despite its biocultural richness, the country is marked with vast social inequality and rural poverty. Following decades of internal armed conflict, the country’s 2016 Peace Agreement has provided new opportunities for its socio-economic growth, representing either a threat to Colombian biodiversity or an opportunity for sustainable development based on its treasured natural resources. The Useful Plants and Fungi of Colombia (UPFC) project aimed to develop pathways to enhance nature’s contribution to people in Colombia by increasing, consolidating and making accessible the knowledge on its useful plants and fungi for the benefit of local communities. The project has provided a framework to develop and promote a market for useful indigenous species and their high-value products whilst protecting the surrounding natural resources. It has produced over 140 dissemination outputs, including books, booklets, scientific journal publications, technical reports, websites, online portals, and educational tools. Also, it has delivered capacity-building events, reaching a broad audience. With the participation of a multinational team, the project compiled and generated knowledge on over 36,000 plants and fungi and developed pathways for tackling socioenvironmental challenges and contributing to Colombia’s green transformation. RESUMEN Colombia ocupa el segundo lugar en el mundo en biodiversidad y es reconocido como uno de los países con mayor diversidad étnica. A pesar de su riqueza biocultural, el país está marcado por una gran desigualdad social y pobreza rural. Tras décadas de conflicto armado interno, el Acuerdo de Paz de 2016 del país ha brindado nuevas oportunidades para su crecimiento socioeconómico que pueden representar tanto una amenaza para la biodiversidad colombiana como una oportunidad para un desarrollo sostenible basado en sus preciados recursos naturales. El proyecto Plantas y Hongos Útiles de Colombia (UPFC) tuvo como objetivo desarrollar vías para mejorar la contribución de la naturaleza a las personas en Colombia, mediante el aumento, la consolidación y la accesibilidad del conocimiento sobre sus plantas y hongos útiles para el beneficio de las comunidades locales. El proyecto ha proporcionado un marco para desarrollar y promover un mercado para especies autóctonas útiles y sus productos de alto valor, al mismo tiempo que se protegen los recursos naturales circundantes. Así, este proyecto ha producido más de 140 productos de difusión, incluidos libros, folletos, publicaciones en revistas científicas, informes técnicos, sitios web, portales en línea y herramientas educativas. También se han realizado eventos de creación de capacidad, llegando a una amplia audiencia. Con la participación de un equipo multinacional, se ha recopilado y generado conocimiento sobre más de 36.000 plantas y hongos, y se desarrollaron vías para abordar los desafíos socioambientales y contribuir con la transformación verde de Colombia. COLOMBIA’S RICH BIODIVERSITY IN DANGER Our life and well-being rely on the environmental goods and services provided by plants and fungi. Not only do they provide us with products, such as food, medicines, natural fibres, fuel, building materials, and cosmetics, but they also act to purify the air, enrich the soil, protect against erosion, regulate water flow and quality, and provide habitats for animals, while acting as a major store of global carbon among other functions (Millennium Ecosystem Assessment, 2005). However, conversion of natural habitats—primarily for farming—is fuelling land degradation, undermining the well-being of two-fifths of humanity, and raising the risk of migration and conflict (IPBES, 2018). Therefore, the preservation and sustainable use of Colombia’s biodiversity are essential to human wellbeing and future economic prosperity. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 21 CHAPTER 1 TABLE 1. Plant and fungal diversity of useful vs non-useful taxa in Colombia, based on data collected by this project. Numbers Plants Useful plants Fungi Useful fungi Plants & fungi Useful plants & fungi Species 28,947 7,472 7,140 374 36,087 7,790 Genera 3,583 2,140 1,763 192 5,346 2,318 Families 395 258 448 99 843 356 TABLE 2. Origin and conservation status of plant and fungal diversity in Colombia, based on data collected in this project. Trait Plants Useful plants Fungi Useful fungi Plants & fungi Useful plants & fungi Cultivated 1,077 1,077 N/A N/A N/A N/A Naturalised 677 638 N/A N/A N/A N/A Native 27,134 5,830 1,264 25 28,398 5,855 Endemic 9,473 478 206 8 9,679 486 Threatened 1,061 249 10 3 1,071 252 Colombia is one of the most biodiverse countries on Earth, comprising at least 28,947 plant species (24% endemic; >1,000 threatened), belonging to 3,583 genera and 395 families (Table 1). The country ranks second in terms of diversity of plants worldwide and is a centre of origin of tomatoes, peppers, potatoes, chillies, and many other crops and their wild relatives (Diazgranados et al., 2020, Khoury et al., 2016). Its fungal diversity is also impressive, with at least 7,140 species belonging to 1,763 genera and 448 families (Table 2). Given that the observed ratio between fungi and plant species in well-documented areas is 9.8:1, Colombia could harbour up to 300,000 fungi species, representing 9% of the global diversity (Gaya et al., 2021). Colombia is also described as one of the most ethnically diverse countries globally, consisting of at least 85 ethnic groups, with 68 recognised native languages. The knowledge associated with useful plants and fungi can be as rich as the cultural variety. The country has been considered a “cradle for modern ethnobotany”, with a plethora of recent studies on this topic (Bernal et al., 2011). However, ethnobotanical knowledge is still vastly under-documented in this region (Cámara-Leret et al., 2014). Regrettably, the country lost just under 6 million hectares of forest between 1990 and 2015, at an average rate of 237,000 hectares per year (FAO, 2015). This deforestation is mainly due to agricultural expansion, urban development and illegal mining, driven by the race for economic growth following the country’s peace process, and inadequate political decision-making that is often not supported by science (Eufemia et al., 2019, Sabater et al., 2017, Salazar et al., 2021). Because this impressive cultural and biological biodiversity is under threat, researchers and conservationists are in a race to protect it (Andrade, 2011, WWF-Colombia, 2017). As deforestation proceeds, the 22 disappearance of traditional knowledge on conservation, use, and management of plants and fungi accelerates with biodiversity loss. A recent review of studies across four Andean countries (Colombia, Ecuador, Peru, and Bolivia) evaluated the sustainability of wild plant use in the region and found that unsustainable harvesting and the loss of useful wild plant species repeatedly arose (Kor et al., 2021). Colombia has the highest percentage of studies reporting unsustainable outcomes (86%). The authors identified five main factors contributing to the unsustainable use of plants: 1) biology; 2) land tenure and access rights; 3) loss of knowledge, resources, and capacity; 4) economic and market pressures due to overexploitation; and 5) institutional structures, policy, and legislation. With its unique and extraordinary biocultural diversity and richness, Colombia can become an example for achieving the Sustainable Development Goals (SDG) (United Nations, 2015) by reducing inequality and poverty through the sustainable use of its natural resources, reducing the degradation of its plants and fungi. Although native non-crop plants and fungi have great potential to improve livelihoods and support economic development, knowledge on the most useful native species of Colombia remains highly dispersed, largely inaccessible, and susceptible to disappearing over time. CAN THE SUSTAINABLE USE OF BIODIVERSITY BECOME PART OF THE SOLUTION? Despite the unique biological and cultural richness of Colombia, vast social inequality and marked poverty in rural areas have triggered more than six decades of internal conflict. Plant and fungal diversity and its associated cultural knowledge could tackle socio-environmental challenges by boosting rural employment and incomes, improving CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 1 people’s livelihoods, reducing inequalities, and supporting peace. Improving access to critical biological information and developing markets for natural ingredients, while meeting sustainability benchmarks, can boost the stake of marginalised communities in the national economy by adding value to their traditional knowledge. Simultaneously, this process would empower meaningful participation, ultimately leading to more stable and equitable socioeconomic outcomes. This approach tackles several of the SDGs, most notably those related to the alleviation of poverty, improvement of health, empowerment of the disenfranchised, sustainability of economic growth and promotion of peace (United Nations, 2015). In recent years, Colombia has established several policies promoting the sustainable use of biodiversity to reduce the social gap and to consolidate peace whilst following a green growth approach to economic development (Baptiste et al., 2017). The country has stated its intention to develop as a bioeconomy, which is “an economy that efficiently and sustainably manages biodiversity and biomass to generate new value-added products, processes, and services based on knowledge and innovation” (CONPES, 2018). The commitment of the Colombian government to green growth has been demonstrated through the Colombia Bio programme and its principal aim to foster the development of the bioeconomy through science, technology, and innovation. As an offspring of this over-arching national effort, the Useful Plants and Fungi of Colombia project (UPFC) was supported by a Professional Development & Engagement grant under the Newton-Caldas Fund partnership. The £2.5 million grant was funded by the UK Department for Business, Energy, and Industrial Strategy (BEIS) and the Colombian Ministry of Science, Technology, and Education (MinCiencias) and delivered by the British Council over 28 months from November 2019 to February 2021. THE USEFUL PLANTS AND FUNGI OF COLOMBIA The overall goal of the UPFC was to develop pathways to enhance nature’s contribution to people in Colombia by increasing, consolidating, and making accessible the knowledge on its useful plants and fungi for the benefit of local communities. In addition, the project promoted a market for useful indigenous species and their high-value products to motivate the sustainable use of biodiversity whilst protecting the surrounding natural resources. This project documented and broadly disseminated knowledge on the useful plants and fungi of Colombia across a wide range of audiences. Outputs targeted policy and decision-makers in government, local NGOs, scientists, educators, and private and public companies, aiming to boost bioeconomic development. The project’s long-term vision was to support Colombia’s economic future by promoting the sustainable expansion of its bioeconomy, based on its unique plant and fungal diversity, whilst conserving its natural resources and associated traditional knowledge. The UPFC project was structured as four work packages (WP), each with a specific objective: 1. document knowledge on useful plants and fungi of Colombia (WP1) 2. synthesise and disseminate knowledge on useful plants and fungi of Colombia using various platforms for different audiences (WP2) 3. develop a framework for creating a sustainable value chain network (VCN) from plant and fungal diversity (WP3) 4. apply a VCN framework to improve local communities’ livelihoods in three pilot areas (WP4). DOCUMENT KNOWLEDGE ON USEFUL PLANTS AND FUNGI OF COLOMBIA (WP1) Native non-crop plants and fungi have great potential to improve livelihoods and economic development in the country. Still, high-quality information on their uses and properties remains scarce or inaccessible. This WP aimed to gather and compile this information to boost Colombia’s bioeconomy and benefit local communities. Significant effort to document the diversity of Colombian flora and fungi has been carried out in recent years (Andrade, 2011, Bernal et al., 2012, Bernal et al., 2019, Diazgranados et al., 2020, Gaya et al., 2021). Still, hundreds of sources ofpublished information had not been compiled and made readily available. The project brought together a binational team of researchers who collected and compiled data from available datasets and publications; digitised data that were not readily available; scanned specimens, illustrations, and records from scientific collections (from herbaria, fungaria, and libraries); generated an image bank of useful plants and fungi of Colombia; developed species distribution maps and models; and created species profiles available online through the project’s portals. The gathered information was analysed to identify taxonomic and geographic gaps and assessed for quality and potential contribution to conservation practices, among other aspects. In all, six research questions on useful plants and fungi were addressed, each with a scientific output (https://in-colombia.org/): • How can we assess the comprehensiveness (in quantity and quality) of the information available for each species? • Are there taxonomic gaps in the available information? • Are there geographic gaps? • What are the most likely factors influencing observed taxonomic and geographic patterns? • Which geographic areas are important for conservation? • Can traditional knowledge erosion be mitigated through Citizen Science? CATALOGUE OF USEFUL PLANTS OF COLOMBIA 23 CHAPTER 1 A B Percentage of well-surveyed cells FIGURE 1. Survey completeness for all vascular A and useful B plants by bioregion (Bystriakova et al., 2021). Percentage of 10×10 km wellsurveyed grid cells, defined as those with ≥ 25 observations, calculated by bioregion. Results from initial activities helped us to identify areas of the country that were poorly surveyed for useful plants and plants in general (Bystriakova et al., 2021), informing subsequent fieldwork and ethnobotanical priorities for WP1 (Figure 1). In addition to the research questions, three tools were designed to manage and collect data on useful plants and fungi, specifically: 1) the shinyCCleaner tool, to clean georeferenced records in the R environment (Ondo et al., pers. comm.); 2) an online survey built-in ESRI Survey123, to collect ethnobotanical information from mobile devices; and 3) a Data Management tool, with modules for data entry and exportation, taxonomic reconciliation, classification of uses, etc. (available online at https://upaf-uat.kew.org/). SYNTHESISING AND DISSEMINATING KNOWLEDGE ON USEFUL PLANTS AND FUNGI OF COLOMBIA (WP2) Access to information about plants and fungi and their uses underpins our collective effort to further understand Colombia’s diversity, allowing us to better conserve species, habitats, and ecosystems and use them sustainably to the benefit of society. Knowledge must be available for different audiences to empower Colombia to protect and sustainably utilise its valuable plants and fungi. This WP aimed to build a comprehensive synthesis and disseminate this through various media, including online portals, scientific journal publications, books, booklets, reports, magazine and news articles, social media streams, and educational tools. This wide range of dissemination outputs and formats aimed strategically to reach a broad audience of policymakers, researchers, and practitioners, including those in governmental, monitoring, and developmental aid 24 agencies, regulatory bodies, conservation organisations, biobusinesses, schools, and universities. Potential users included: innovators, developers, bioeconomists, herbal and food entrepreneurs (including plant and fungi importers, suppliers and retailers), biotechnologists, bioenergy experts, agronomists, forestry engineers, crop specialists, horticulturalists, arborists, weed scientists, forest managers, wildlife managers, health professionals, biopharmaceutical scientists, medicine regulators, conservationists, naturalists, photographers, environmental consultants, library managers, students, professors, researchers (biologists, botanists, ecologists, chemists, agronomists, biomedical researchers, ethnobiologists, etc.), scientific editors, people interested in culture and gardening, and the general public. The project built two websites (https://www.kew.org/ upfc, in English; and https://in-colombia.org/; in Spanish) to inform the public and to disseminate project outputs. In addition, it built on the previously created ColPlantA portal (http://colplanta.org/), further synthesising knowledge of Colombian plants and improving the user experience, content, and infrastructure (Diazgranados et al., 2020). It also created a companion portal for Colombian fungi, ColFungi (http://colfungi.org/) (Figure 2). Each portal was accompanied by an explanatory booklet (Diazgranados et al., 2020, Gaya et al., 2021) designed for a general audience. Comprehensive annotated checklists for both plants and fungi, along with a range of thematic chapters, are presented here and in the Catalogue of Fungi of Colombia (de Almeida et al., 2022) as a reference for scientists and long-term preservation of knowledge. A survey analysis was conducted to reveal how much Colombian society knows about their native diversity CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 1 of plants and fungi and their uses as part of this WP. An online national survey was developed to achieve this, which collected information on how Colombians generally use plants and fungi. Semi-structured interviews of 20 experts from different sectors of society, including academia, civil society, NGOs, and the private sector, were also undertaken. These experts provided information regarding the state of knowledge of useful plants and fungi. They advised on the limitations and opportunities of various studies and the inclusion of plants and fungi in value chains. In consideration of socio-cultural components, a Spanish-language storytelling series, “Somos Historias”, was developed to identify and promote practices and relationships that underlie key the value chains of key plants. The co-creation of short videos that portrayed the practices, knowledge, and relationships of local producers with these plants and their environment promoted value chains and increased project visibility to a wider audience, while also gathering qualitative information on the local use of the species. Finally, the UPFC project published over 50 blogs and notes in the press to raise awareness on the importance of useful plants and fungi for the well-being and prosperity of Colombians and humanity. BUILDING ON COLPLANTA A significant output under WP2 was work carried out to improve “ColPlantA – Colombian resources for Plants made Accessible” (http://colplanta.org/), an authoritative, expert-driven, open access, an online portal for botanical FIGURE 2. ColPlantA (https://colplanta.org/) and ColFungi (https://colfungi.org/) site homepages. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 25 CHAPTER 1 information on Colombian plants that was officially launched in August 2019 during the 10th Colombian Botany Meetings as part of a previous project (Diazgranados et al., 2019, Diazgranados et al., 2020). ColPlantA is a content aggregator site – it collects information from Kew’s databases and various external sources and displays it in one place while preserving the appropriate links and citations. Therefore, it is a comprehensive resource that for the first time, brings together high-quality taxonomic information with a wide range of data and information sources. In doing so, ColPlantA unlocks valuable information, enriches the data and its potential uses, and maximises the visibility of local resources. Only publicly available content is used and, as part of the content-gathering process, steps are taken to ensure that publishers have respected the IBCD Nagoya Protocol (to ensure the fair and equitable sharing of benefits arising from the utilisation of genetic resources) as well as Colombian laws relating to traditional knowledge. ColPlantA holds an online profile for each plant species known from Colombia, aggregating information from other databases and portals, and displaying previously inaccessible information. It links to type specimens from RBG Kew, species descriptions, maps on geographic distribution, published plant uses, sustainable practices, conservation status, photographs, common names, and information on ecology and natural history. Currently, ColPlantA holds more than 27,400 accepted species profiles (including 19,644 with images), 145,000 scientific names, and circa 20,000 distribution maps for Colombian plants (ColPlantA (http://colplanta.org/), consulted on 18 February 2022). Since its launch, the portal has hosted approximately 31,500 unique users in 97 countries, averaging 1,400 sessions by 1,260 users each month. The site has been crawled and indexed by Google, with roughly 86,900 web pages indexed (SEOQuake, consulted on 18 February 2022). The portal is linked to the Plants of the World Online (POWO) database – an RBG Kew Strategic Output – and any change to POWO is updated automatically in ColPlantA. ColPlantA supports the identification and understanding of the benefits provided by plant diversity, utilising new technologies, concepts, and social frameworks. Also, it helps to underpin the wider goals of Colombia’s green growth by providing evidence of links between plants, uses and sustainable development goals. DIGITAL OUTREACH OPPORTUNITIES The project reached a range of audiences and created new alliances and opportunities for collaboration through an array of varying virtual events and social media activities. Despite the recent challenges presented by the coronavirus pandemic, the project found new methods to engage with its target audiences through virtual events, such as Hay Festival Cartagena Virtual, Hay Festival Digital Queretaro 2020, Hay Festival Cartagena 2022 FIGURE 3. The VCN website (“Red de Ingredientes Naturales – Colombia”; https://redin-colombia.org/), displaying the registration form for connecting to the market chain. The site also includes a blog, a serving BOT, and a practical guide to understanding the legal processes in Colombia for developing and commercialising natural ingredients or products of any kind from non-crop plants and fungi. 26 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 1 the Colombian Chef Masterclass series, and virtual workshops, reaching over one million viewers. In addition, it delivered more than 23 virtual conferences, five seminar presentations, three teaching modules, 30 socialisation workshops, and fifteen radio interviews. It had more than 90,000 engagements in its social accounts (Facebook and Twitter), and the project published ten blogs and more than 90 popular articles and media notes, all of which helped to broaden the project’s recognition. BUILDING A VALUE CHANGE NETWORK TO SUPPORT COLOMBIA’S BIOECONOMY (WP3) Innovative projects to deliver natural ingredients or other bioproducts to the market often fail because a viable market has not been established. The project developed a value chain network (VCN) to provide pathways to develop such markets. This VCN was centred on a digital platform that facilitates interaction between local, national, and international actors in the production, refinement, distribution and exportation of plant- and fungi-based products. The VCN aims to enable the connections between plant and fungal diversity, knowledge of its uses and local livelihoods which depend on it. It is the first VCN focused on useful plants and fungi for the country and the continent. The VCN connects scientists, technologists, innovators, producers, and consumers. It enables the flow of materials downstream (e.g., the supply of natural ingredients produced by local communities to consumers) and information upstream (e.g., consumer demand to producers) whilst linking products to species names on the basis of a taxonomic backbone. This functionality allows two-way linking between the VCN platform and the scientific portals (ColPlantA and ColFungi), enabling users to browse specific uses, read interesting plants and fungi species profiles, and contact product providers directly. A website for the VCN (https://www.redin-colombia.org/) was created in Spanish to maximise outreach among the Colombian audience (primary users) (Figure 3). In addition, the project delivered the first comprehensive practical guide (174 pages) to understanding the Colombian legal processes for developing and commercialising natural ingredient products from non-crop plants and fungi (Rojas et al., 2021, available at https://redin-colombia.org/guide/). This freely available guide is in Spanish and is organised around four central roles within the value chain: producer, refiner, national wholesaler/retailer, and exporter. Content focuses on permissions, regulatory compliance, and certification concerning three types of natural ingredients: harvested non-timber forest products, nursery-grown products, and genetic material products. It articulates processes related to intellectual property rights and benefitsharing agreements, a threatened species, among others related to national and international regulations, compliance, and market access. Eleven semi-structured interviews were conducted with staff from the Ministry of Environment and Sustainable Development, ProColombia, natural product companies, and NGOs. Two online tools were created to accompany the guide: 1. a serving BOT, which is an automated tool designed to guide users to obtain information from within the guide by answering a series of targeted questions regarding their needs and the type of commercial role they play within the value chain (e.g., producer, refiner, distributor, retailer, exporter). The serving BOT is accessible via the VCN website (https://www.redin-colombia.org/) 2. an online quiz in Spanish, drawing upon the information contained within the guide, that tests users’ knowledge regarding documentation, certification and responsible agencies related to the production, distribution, and exportation of bioproducts from Colombia. Access to the live quiz platform in Kahoot can be found through the link: https://kahoot.it/challenge/00115304 or in the practical guide. To establish a baseline for the current status of the Colombian bioeconomy, researchers produced a 124-page report (Rojas et al., 2020), also freely accessible at https:// in-colombia.org/. Highlights included: 1. a prioritisation exercise for potential species in Colombia’s bioeconomy 2. a broad directory of companies engaged in the use of prioritised species of native plants and fungi (25 in total) 3. a Science, Technology, and Innovation capability analysis around the Bioeconomy and Natural Ingredients sector 4. a multi-criteria value chain infrastructure for biodiversitybased products 5. a case study of the value chain analysis for the first prioritised species, naidí (Euterpe oleracea Mart.), in the Pacific Coast region of Colombia 6. a search and analysis of patents and publications related to naidí in Colombia and the rest of the world Finally, to inform the VCN, a standard for controlled vocabulary on natural ingredients and bioproducts was developed, which included a harmonised table and accompanying explanatory report. The standard will allow their classification, indexing, cataloguing, description, and analysis. It was based on the harmonisation of three classification structures linking natural ingredient products: Kew’s Economic Botany Data Collection Standard (EBDCS), the UN Central Product Classification scheme (CPC) and the Departamento Administrativo Nacional de Estadistica’s Clasificacion Central de Productos (DANE CPC). These three structures were reconciled on the basis of each hierarchy’s concept, definition, and rules to identify a common, core standard linking botanical (e.g., species), national and international bioproduct classification structures. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 27 CHAPTER 1 FIGURE 4. Map of urban centres in which the main collaborators are based, and pilot areas selected for the implementation of the VCN framework: Bahía solano (Chocó), Becerril (Cesar), and Otanche (Boyacá). Photograph credits: Mateo Hernández (Becerril), Mauricio Diazgranados (Otanche), and Daniel Uribe (Bahía Solano). 28 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 1 IMPLEMENTING A VALUE CHANGE NETWORK IN THREE PILOT AREAS OF COLOMBIA (WP4) Three biodiverse pilot areas of Colombia that had been affected recently by the armed conflict were selected to test the potential of the VCN framework to improve impoverished communities’ livelihoods (food security, health, and prosperity): Bahía Solano (Chocó), Becerril (Cesar) and Otanche (Boyacá) (Figure 4). The selection of the areas considered several criteria within five categories: environmental, social, governance, business, and economic factors (Diazgranados et al., 2021). The implementation included: 1. assessment of the useful local plants and fungi resources 2. evaluation of the market status based on local native diversity 3. assessment and modelling of current and potential outcomes of the VCN implementation 4. identification of the most promising species and types of natural ingredients 5. the gleaning of lessons from the implementation and community feedback The project expected that implementing the VCN would improve family livelihoods over the middle- to long-term through its role in facilitating diversified, market-based approaches to income generation. The social impact assessment carried out during the implementation of the VCN will allow the measurement of middle- and long-term changes in cultural, economic, and social outcomes. The outcome monitoring activities implemented at the time of writing include participant surveys (collecting data on gender, age, locality, ethnic group, family composition, etc.) and analysis of statistics relating to the potential implementation. The project published a booklet to connect experiences in local communities within the three pilot areas with a broader, more general audience. It included highlights, notes on the prioritisation of species, stories from local initiatives and experiences (QuiñonesHoyos et al., 2021). THE UPFC TEAM The UPFC project was led by RBG Kew in collaboration with the Instituto de Investigación de Recursos Biológicos Alexander von Humboldt in Colombia. It directly involved 85 people (50 from Kew and 35 from the Humboldt Institute) and received contributions from 34 other researchers representing 26 institutions across Colombia and Europe. Diverse and gender-balanced participation was accomplished in all activities to ensure a more inclusive environment and in line with SDG5 (Empower women). We have been collaborating to include minorities such as indigenous people, Afro-Colombians and the LGBT+ community, always following RBG Kew’s and Humboldt Institute’s policies on diversity and inclusion. Of 115 project participants, 64 (56%) were female and 51 (44%) were male. The project deemed this gender balance to have been appropriate given the traditional tendency of Botany to be a male-dominated discipline. REACHING COMMUNITIES ACROSS SOCIETY The UPFC project built and maintained connections with other government and non-government initiatives. Through various fora and media, objectives and outcomes were communicated to the Presidency of the Republic, the National Committee of Sustainability, the Green Business Bureau of the Ministry of Environment, the Programa de Transformación Productiva (PTP) of the Ministry of Commerce, ProColombia, the Instituto de Investigaciones Ambientales del Pacífico John von Neumann (IIAP) and the Green Growth Mission (GGM) created by the Colombian National Planning Department in 2017. The GGM, together with the Biointropic Corporation and Suricata Consulting, identified seven factors to boost the country’s bioeconomy: 1) research and technological development; 2) regulatory framework; 3) market dynamics; 4) human capital; 5) financing and investment; 6) environmental or ecosystem services; and 7) infrastructure. By contributing directly to factors 1–4, the project supported GGM’s mission and the Green Growth Policy, i.e., “Promote by 2030 the increase in productivity and economic competitiveness of the country, while ensuring the sustainable use of natural capital and social inclusion, in a compatible way with the climate”. It engaged the Colombian National Board of Natural Ingredients (Programa de Interés estratégico-PINE), an initiative led by the Program Colombia + Competitiva of SwissContact as part of a 5-year plan, to boost natural ingredients for cosmetics. The Humboldt Institute leader of the board’s six sub-activities regarding the governance and competitiveness of native plant species’ value chains were key in maintaining this interaction. The project also connected with the private sector, such as Crepes & Waffles, the Chamber of Food Industry, the National Association of Industrials of Colombia (ANDI), Corpocampo, Naidí Pacífico SAS, Fondo Acción, Planeta CHB SA, Partnerships for Forest, MUCHOCOL, UNIANDES (Faculty of Economics), Apsacesar, Envol-vert, Selvacéutica S.A.S., E3 Asesorías, Corporación Boyapaz, Alianza Quinchas, among many others. The botanical work, particularly around the initial consolidation of the ColPlantA portal, involved botanists from several Colombian organisations, including the Asociación Colombiana de Herbarios (ACH), the Grupo Etnobotánico Latinoamericano (GELA) and several universities such as Universidad de Antioquia, Universidad de la Amazonia, Universidad de los Llanos, Universidad de Nariño, Universidad de Pamplona, Universidad del Cauca, Universidad Distrital “Francisco José de Caldas”, Universidad Nacional de Colombia, Universidad Pedagógica y Tecnológica de Colombia, and Universidad Tecnológica del Magdalena. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 29 CHAPTER 1 FIGURE 5. UPFC fieldwork activities in the pilot areas. Top left: collecting useful plants and fungi with the help of local “sabedores” (wisdom people). Top right and bottom: species prioritisation workshops with local communities. Photograph credits: Mateo Fernández (top left), Sabina Bernal (top right and bottom right), and Edgar Padilla (bottom left). Fieldwork was carried out through a participatory approach involving local organisations, social leaders, and communities from the veredas of Caño Rodrigo and Río Maracas in Becerril (Cesar), and Altazor, Betania, Camilo, Cunchalita, Curubita, El Carmen and Nazareth in Otanche (Boyacá) (Figure 5). In conclusion, the UPFC project played a pivotal role by reaching communities across society, promoting the conservation and sustainable use of the country’s plants and fungi, combining traditional and scientific knowledge to boost the national bioeconomy, and raising awareness about the importance of these resources for the country. On a broader scale, it represents a sustainable development model of green growth in megadiverse nations worldwide. UNEXPECTED IMPACTS AND CHALLENGES DUE TO COVID-19 The pandemic of 2020–21 had significant impacts on where and how the UPFC project team carried out their work due to the restriction of travel and facility access, recruitment delays, and the shortage of human resources. 30 Mitigation plans proved effective in buffering some of the effects, especially regarding WP1, WP2 and WP3 workflows and timelines, which used increased teleconferencing and online stakeholder consultations with several crosscutting contingency actions. Nevertheless, capacity building (e.g., the internship and research visitor programmes) and fieldwork activities had to be scaled down considerably, and future funds could help to strengthen the implementation of the VCN at the national scale. PATHWAY TO IMPACT ON ECONOMIC DEVELOPMENT AND SOCIAL WELFARE This project has developed novel pathways to enhance nature’s contribution to people in Colombia. It has made available ‘state-of-the-art’ knowledge on the nation’s useful plants and fungi, while unveiling how this information can be used to promote and develop markets for these species and their high-value products. CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 1 Beyond the delivery of specific outputs, a broader set of project outcomes were achieved. These include: The project pathway to impact on economic development and social welfare was based on a stepwise approach: • a significant increase in online, open-access knowledge on useful plants and fungi of Colombia • Short-term (1–3 years): the project documented and broadly disseminated knowledge on useful plants and fungi of Colombia, reaching both general and academic audiences, as well as policy and decision-makers. It also contributed to capacity building. Last, by implementing the VCN in three pilot areas, the project aimed to support improvements to communities’ livelihoods by establishing a baseline that can be monitored and provide initial training to communities. • increased capacity of Colombian scientists to research plant and fungal diversity and uses • increased public perception and awareness about the importance of plant and fungal diversity and their uses to support green growth in Colombia • improved opportunities for alternative local income streams based on value-added bioproducts through VCN development, at least in the three pilot areas • supported expansion of a natural ingredients market at the national level • opportunities opened for further improvement to people’s livelihoods in the three pilot areas • informed development of a bioeconomy, bringing together science, policy, and industry. The predicted spillover effects included: • increased interest by Colombian institutions in expanding plant and especially fungal research • enhanced recognition of market possibilities for natural ingredients based on native species of plants and fungi at the Colombian national level • incentive to develop specific areas of commercial businesses based on natural ingredients from native Colombian plants and fungi, e.g., by the British and Colombian Chamber of Commerce, ProColombia, etc. • expansion of Colombian bioeconomy by strengthening industrial endeavours to add value to useful plants and fungi products. In addition to the contributions to the United Nations SDGs, the project outputs and activities also contributed to several objectives of the Convention of Biological Diversity (CBD) relating to the targets defined from the Global Strategy for Plant Conservation (GSPC) 2010–2020, particularly those linked to ecosystem goods and services (GSPC1), recognition, respect, and maintenance of indigenous and local community knowledge (GSPC 5), and improved awareness of the urgency of plant conservation and public participation (GSPC 6). As most communities in rural Colombia are maledominated, with men focused on farming, mining and construction, the project emphasised pathways for women to play lead roles in the obtention and commercialisation of natural ingredients, potentially bringing additional income to the household. Thus, the implementation of the VCN could improve gender equality and opportunities, supporting SDG5 (Empower women). • Mid-term (3–10 years): the project outputs will increase awareness in Colombian society about the importance of the country’s diversity of useful plants and fungi, and will inform policy and decision-makers in government and non-government organisations, thereby boosting sustainable biobusinesses and supporting national and international-scale connections. The VCN may improve the livelihoods of communities at the national scale, providing new opportunities to reduce poverty and gender inequality, and to improve health and well-being. • Long-term (10–15 years): Colombia will rise as a successful example of a developing country that is expanding its bioeconomy based on its unique plant and fungal diversity while preserving its natural environment. FUTURE PERSPECTIVES AND FINAL REMARKS The project has contributed significantly to the Colombian Green Growth Policy’s scope to “Promote by 2030 an increase in productivity and economic competitiveness of the country, while ensuring the sustainable use of natural capital and social inclusion, in a compatible way with the climate” (CONPES, 2018). While the transformation of a country is a very complex, long-term process, usually involving all levels of society, the UPFC project is confident that its outputs and outcomes have supported Colombia’s transformation towards a greener, more sustainable economy. This has been achieved by enhancing critical baseline knowledge on plants and fungi and re-imagining how access and practical use of this information could be improved through open-access digital platforms and tools. The impact of the UPFC project will continue, as the information generated here has been made broadly available to Colombian companies that have an interest in the sustainable development of biodiversity-based products (bio-based), typically the agricultural, pharmaceutical, food, human health, and personal care sectors. Colombia’s green transformation has begun! CATALOGUE OF USEFUL PLANTS OF COLOMBIA 31 CHAPTER 1 Acknowledgements The UPFC project thanks all the researchers and contributors who made this project possible: Adriana Corrales, Aida Marcella VascoPalacios, Alba Nohemi Téllez A., Alejandra Aguilar, Alejandra Osejo, Alex Monro, Aline Dufat, Amalia Díaz, Ana Cristina Bolaños-Rojas, Ana Esperanza Franco-Molano, Ana Garrido, Andrea Baquero, Angélica Herrera, Angie Karina Rengifo Fernandez, Anna Haigh, Benedetta Gori, Bibiana Moncado, Bob Allkin, Camilo Garzón, Carlos Cortés, Carolina Quiñones, Carolina Soto, Carolina Tovar, Catalina Mesa Sánchez, Clara Morales, Cristian Lasso-Benavides, Dairon López, Daniel Jiménez, Daniel López, David Bishop, David Díaz-Escandón, Diana Marcella Moreno, Diego Simijaca-Salcedo, Edier Soto-Medina, Eduardo Toledo Romero, Edwin Rios, Ehidy Rocio Peña-Cañón, Elizabeth Hodson, Ellie Graves, Emmerson Pastás, Ester Gaya, Fabiola Eugenia González-Cueller, Felipe García, Germán Torres, Guilana Furci, Harrie J.M. Sipman, Henry Agudelo, Henry Yesid Bernal M., Hugh Pritchard, Ian Ondo, James Morley, Jenny Williams, Jennyfer Andrea Aldana Mejía, Joaquim d’Souza, Joseph Ruff, Julia Carretero, Justin Moat, Laura Green, Laura Kor, Lee Davies, Lina Guevara, Louise Colville, Luis Fernando Coca, Lynn Parker, Malcolm Stone, Marcela Cely, Margarita María Jaramilo-Ciro, Mariana Pizano Noguera, Mario Murcia, Mark Nesbitt, Martha Lucía Ortiz-Moreno, Mateo Fernández, Mauricio Ramírez-Castillón, Mauricio Salazar-Yepes, Meike Piepenbring, Melanie-Jayne Howes, Mimi Tanimoto, Nadia Bystriakova, Natalia Vargas-Estupiñán, Nataly Gómez-Montoya, Nestor García, Nestor R. Salinas, Nicolas Black, Pablo Hendigo, Paola Acosta, Paola Andrea Torres-Andrade, Paul Kersey, Paul Wilkin, Priscilla Reis, Rafael Felipe de Almeida, Raquel Negrao Baldoni, Rene Camacho, Robert Lücking, Rose Taylor, Sabina Bernal, Santi Gómez, Sarah Phillips, Silvia Restrepo, Tatiana Sanjuan, Tim Utteridge, Tiziana Cossu, Vicky Murphy, Viviana Motato-Vásquez, Wiebke Hillebrecht, William Baker, William Milliken, and Yeina Milena Niño-Fernández. The project also thanks the local communities and authorities in the pilot areas, as well as all the stakeholders involved. References Andrade CMG (2011) Estado del conocimiento de la biodiversidad en Colombia y sus amenazas: consideraciones para fortalecer la interacción Ciencia-Política. Rev. Acad. Colomb. Cienc. Exactas Fis. Nat., 35 (137): 491-507. Baptiste B, Pinedo-Vasquez M, Gutierrez-Velez VH, Andrade GI, Vieira P, Estupiñán-Suárez LM, Londoño MC, Laurance W, Lee TM (2017) Greening peace in Colombia. Nat. Ecol. 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Available at: https://www.ipbes.dk/ udgivelser-og-viden/ipbes-udgivelser/ Khoury CK, Achicanoy HA, Bjorkman AD, Navarro-Racines C, Guarino L, Flores-Palacios X, Engels JMM, Wiersema JH, Dempewolf H, Sotelo S, Ramírez-Villegas J, Castañeda-Álvarez NP, Fowler C, Jarvis A, Rieseberg LH, Struik PC (2016) Origins of food crops connect countries worldwide. P Roy Soc B-Biol Sci, 283: 20160792. Kor L, Homewood K, Dawson TP, Diazgranados M (2021) Sustainability of wild plant use in the Andean Community of South America. Ambio, 50: 1681-1697. doi: 10.1007/s13280-021-01529-7 Millennium Ecosystem Assessment (2005) Ecosystems and human wellbeing. Washington, DC. 156p. https://www.millenniumassessment. org/documents/document.356.aspx.pdf CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 1 Quiñones-Hoyos C, Rengifo-Fernández A, Bernal-Galeano S, Peña R, Fernández M, Rojas T, Diazgranados M (2021) Una mirada a las plantas y los hongos útiles en tres áreas biodiversas de Colombia. London, Royal Botanic Gardens, Kew & Bogotá, Instituto de Investigaciones en Recursos Biológicos Alexander von Humboldt. Rojas MT, Cortés C, Noguera M, Acosta P, Diazgranados M (2021) Guía práctica para potencializar el uso sostenible de los ingredientes naturales en Colombia. London, Royal Botanic Gardens, Kew & Bogotá, Instituto de Investigaciones en Recursos Biológicos Alexander von Humboldt. 170p. Rojas T, Cortés C, Noguera M, Ulian T, Diazgranados M (2020) Evaluación del estado de los desarrollos bioeconomicos colombianos en plantas y hongos. London, Royal Botanic Gardens, Kew & Bogotá, Instituto de Investigación de Recursos Biológicos Alexander von Humboldt.123p. Sabater S, González-Trujillo JD, Elosegi A, Rondón JCD (2017) Colombian ecosystems at the crossroad after the new peace deal. Biodivers. Conserv., 26: 3505–3507 doi: 10.1007/s10531-017-1415-8 Salazar A, Salazar JF, Sánchez-Pacheco SJ, Sanchez A, Lasso E, Villegas JC, Arias PA, Poveda G, Rendón ÁM, Uribe MR, Pérez JC, Dukes JS (2021) Undermining Colombia’s peace and environment. Science, 373: 289-290. United Nations (2015). Transforming our World: the 2030 Agenda for Sustainable Development, Document A/70/L.1. UNGA, New York. WWF-Colombia (2017) Colombia Viva: un país megadiverso de cara al futuro. Informe 2017. Cali, WWF-Colombia. 166p. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 33 CHAPTER 1 Field expedition with community participants in the Municipality of Otanche. Laura Kor 34 CATALOGUE OF USEFUL PLANTS OF COLOMBIA Chapter 2 Assessing extinction risk coverage and identifying geographical hotspots to guide conservation action for Colombia’s useful plants Laura Kor 1,2, Cristina Lopez-Gallego 3, Carolina Castellanos-Castro 4 & Mauricio Diazgranados 1* 1 2 3 4 Research Department, Royal Botanic Gardens, Kew, Richmond, TW9 3AE, UK Department of Geography, Bush House NE, King’s College London, London, WC2B 4BG, UK Instituto de Biología, Universidad de Antioquia, Medellín, Colombia Instituto de Investigación en Recursos Biológicos Alexander von Humboldt, Bogotá, Colombia *Corresponding author: m.diazgranados@kew.org Keywords: Conservation planning; Conservation prioritisation; Important Plant Areas; Red List; in-situ conservation ABSTRACT Colombia is one of the most biologically and culturally diverse countries in the world, supporting over 7,000 plant species with documented human uses. However, large gaps remain in understanding the country’s biodiversity and its role in human welfare. Simultaneously, global conservation strategies increasingly highlight the importance of the protection and sustainable use of socially, economically, and culturally valuable species. Drawing on the Checklist of Useful Plants of Colombia (CUPC), this chapter proposes conservation actions based on: (1) taxonomic gaps in extinction risk assessments and (2) geographic hotspots for native useful plants of Colombia (UPC). We identified 544 species of conservation concern – threatened with extinction and/or endemic to Colombia. However, 45% of native UPC lack extinction risk assessments. The proportion of unassessed species was significantly higher in 13 of the 256 useful plant families. We propose that these families should be prioritised for extinction risk assessments. There were also variations between use categories, with 73% of fuel species assessed compared to 42% of species with environmental uses. Existing records of native UPC were distributed across all major ecosystems in Colombia. Applying three definitions of hotspots and using a regionalised approach that combines habitat and municipality boundaries, we identified 78 units of analysis (89,901 km2) as hotspots for native UPC. In-situ conservation gaps exist, with 48 hotspots having less than 25% of their land within Colombia’s current protected area or Indigenous and Community Conserved Area systems. We discuss how these hotspots can contribute to the identification of Important Plant Areas in Colombia. While prioritisation is required to target limited conservation planning and resources efficiently, stakeholder engagement will be crucial to develop and implement conservation action and sustainable use for the benefit of people and nature. RESUMEN Colombia es uno de los países del mundo con mayor diversidad biológica y cultural, al albergar más de 7.000 especies de plantas con usos documentados. Sin embargo, siguen existiendo grandes vacíos en la comprensión de la diversidad biológica del país y su papel en el bienestar humano. Al mismo tiempo, las estrategias de conservación global de la biodiversidad destacan cada vez más la importancia de la protección y el uso sostenible de especies de valor social, económico y cultural. A partir de la Lista de Verificación de Plantas Útiles de Colombia (CUPC), este capítulo propone acciones de conservación basadas en (1) vacíos taxonómicos en las evaluaciones de riesgo de extinción y (2) puntos geográficos críticos para las plantas nativas útiles de Colombia (UPC, por su sigla en inglés). Identificamos 544 especies de interés para la conservación – amenazadas de extinción o endémicas de Colombia. Sin embargo, el 45% de las UPC nativas carecen de evaluaciones de riesgo de extinción, con una proporción de especies no evaluadas significativamente mayor en 13 de las 256 familias de plantas útiles. De esta manera se propone priorizar estas familias para futuras evaluaciones de riesgo de extinción. También se encontraron variaciones en este aspecto entre las categorías de uso, con el 73% de las especies usadas para combustible evaluadas comparado con el 42% de las especies con usos ambientales. Los registros existentes de las UPC se distribuyeron a largo de los principales ecosistemas de Colombia. Aplicando tres definiciones de puntos críticos y utilizando un enfoque regionalizado que combina hábitats y límites de municipios, identificamos 78 unidades de análisis (89.901 km2) como puntos críticos para las UPC nativas. Se identificaron vacíos en términos de áreas gestionadas para la conservación in-situ, con 48 regiones críticas que tienen menos del 25% de su área terrestre dentro de un área protegida actual de Colombia o dentro de territorios colectivos indígenas o de afrodescendientes. Discutimos cómo estos puntos críticos pueden contribuir a la identificación de Áreas de Importancia para la Conservación de las Plantas (IPAs por sus siglas en inglés) en Colombia. Si bien se requiere priorización para orientar de manera eficiente la ejecución de los limitados recursos para conservación, la participación de las partes interesadas será crucial para desarrollar e implementar acciones de conservación y uso sostenible para las personas y la naturaleza. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 35 CHAPTER 2 INTRODUCTION Plants are the primary producers in almost all ecosystems, providing the structure and resources needed for life on earth. Additionally, more than 40,000 plant species have a documented human use, from food and fuels to plants with ritual or religious significance (Diazgranados et al., 2020). Despite their importance, two in five plant species globally are threatened with extinction, driven by habitat conversion, overexploitation and climate change (Nic Lughadha et al., 2020). Effective plant conservation is therefore urgently needed for the wellbeing of both people and biodiversity. Colombia is one of the most biodiverse countries on earth (Corzo et al., 2010). The region is also rich in cultural diversity, with thousands of plant uses documented across Colombia and the wider Andean region (Bernal et al., 2011, Paniagua-Zambrana & Bussmann., 2020, Kor et al., 2021). Following decades of internal conflict, the 2016 Peace Agreement in Colombia has provided an opportunity to improve biodiversity knowledge and simultaneously has generated socio-economic changes, with impacts on natural resources (Bridge Colombia, 2017). This combination of factors makes Colombia a particularly pertinent country for socio-ecological approaches to plant conservation. The identification of conservation priorities is a crucial step in this process to enable targeted conservation planning and resource allocation. This chapter summarises current plant conservation efforts before discussing and evaluating useful plant conservation in Colombia. We aim to propose conservation priorities based on useful plant species by highlighting gaps in extinction risk assessments and identifying geographic hotspots. We draw on the Checklist of Useful Plants of Colombia (CUPC; Chapter 12) to ask: (1) which useful plant species are known to be at risk of extinction? (2) what are the gaps in extinction risk assessments? (3) where are the hotspots for useful plant species? and (4) how well are they represented in national protected areas and Indigenous and Community Conserved Areas (ICCA)? International framework for plant conservation The Convention on Biological Diversity (CBD) is a major international agreement that guides plant conservation efforts. Plants have their own taxonomically focused strategy that was adopted in 2002, the Global Strategy for Plant Conservation (GSPC), which aims to “halt the continuing loss of plant diversity” (CBD, 2012). This strategy was updated in 2010 in line with the Aichi Biodiversity Targets, and the Post-2020 Global Biodiversity Framework is currently being developed (CBD, 2021). While all plant conservation efforts can benefit useful plants, more targets in the Draft Post-2020 GSPC directly highlight socio-economically important species and sustainable use than ever before TABLE 1. Preliminary 2030 targets of the Global Strategy for Plant Conservation draft that have direct relevance to useful plant species (values denoted as “a significant percentage” are still to be confirmed at the time of writing). 36 Target Detailed aims 4 By 2030, there has been at least a [50%] reduction in the number of plant species threatened by international trade and by unsustainable levels of harvesting. 9 9a: By 2030, a significant percentage (to be defined) of areas under agriculture, aquaculture and forestry are managed sustainably, ensuring the conservation of associated wild and crop plant diversity 9b: By 2030, a significant percentage of crop varieties, landraces, forest genetic resources, crop wild relatives (CWR) and other domesticated socio-economically and culturally valuable plant species are conserved ex-situ, and viable populations are effectively managed in-situ to prevent genetic erosion and safeguard their genetic diversity. 12 By 2030, a significant percentage of countries are benefitting from the exchange of plant materials and associated traditional knowledge to support plant conservation, ecological restoration, and sustainable use. 13 By 2030, at the latest, plant diversity values have been integrated into rural and urban development and poverty reduction strategies and planning processes and have been implemented in natural capital and other national accounting mechanisms and reporting systems worldwide. 19 19d: By 2030, the value of plant diversity and responsibility for its protection is universally recognised by the world’s people, including the ecosystem services they provide and the steps that can be taken to conserve and use plants sustainably. 20 By 2030, with the full and effective participation of indigenous and local communities, at all relevant levels, the traditional knowledge, innovations, and practices of indigenous and local communities relevant for the conservation and sustainable use of plant diversity are respected, safeguarded, and preserved to support customary and cultural use of these resources. CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 2 (Table 1). These conventions and strategies demonstrate international recognition of the importance of the direct use of plants and the need to recognise and conserve local and indigenous knowledge for biodiversity conservation and sustainable development. An additional international treaty that contributes to the conservation of plants is the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). CITES was established to regulate the international trade in specimens of wild animals and plants and currently includes over 32,800 plant species, most of which are of socio-economic importance. Important Plant Areas With global conservation efforts commonly focused on area-based protection, Targets 2 and 19b of the Post-2020 GSPC are also relevant to useful plant conservation. They aim for the identification of “the most important areas for plant diversity [to be] identified” and “adequately protected for plant conservation”. Traditional approaches to protected areas often involved fortress-conservation strategies, which restricted access to ecosystem services and ignored the role of local and indigenous communities (Palomo et al., 2014). The Important Plant Area (IPA) programme, established to support GSPC targets, does not confer national legal protection and can instead be used as a tool to support national and regional strategic planning and to drive community-led sustainable management (Darbyshire et al., 2017). This tool is crucial as we increasingly recognise that for scientific findings to translate into sustainable use, there needs to be an understanding and integration of the social, ethical, economic, political, and religious factors which influence conservation outcomes (Cunningham, 2001, Kor et al., 2021). IPA identification is based on three globally consistent criteria, first published by Anderson (2002). The criteria were updated in 2017 to include socially, economically, or culturally valuable species – or useful plant species (Darbyshire et al., 2017). However, the application of IPA criteria to useful plants is limited, with a notable exception being IPAs for medicinal plants in the Himalayas (Hamilton et al., 2007). PLANT CONSERVATION IN COLOMBIA As a party to the CBD, Colombia has developed policies and strategies for biodiversity conservation, including a National Biodiversity Strategy and a NSPC – National Strategy for Plant Conservation (or ENCP, Estrategia Nacional para la Conservación de Plantas in Spanish). The latter has driven the country’s plant conservation policy and action, including the publication of a Catalogue of Plants and Lichens (Bernal et al., 2020) and a series of Red Books of Plants in Colombia (Garcia et al., 2007). Risk assessments for species are ongoing and follow the methodologies of the International Union for Conservation of Nature’s (IUCN) Red List of Threatened species (hereafter, IUCN Red List). The implementation of Colombia’s NSPC was reviewed by the Humboldt Institute and the National Network of Botanical Gardens in 2010 (García Martínez et al., 2010), with the publication of an Action Plan in 2017 (CastellanosCastro et al., 2017). Aligned to the NSPC, a diagnosis and guidelines for the conservation of medicinal plants have been developed (Bernal et al., 2011), and conservation action plans for taxonomic groups of charismatic and socioeconomic importance, such as orchids, palms, cycads, and some timber trees, have been published. Other conservation measures in Colombia include a National System of Protected Areas (SINAP), with public national protected areas consisting of 122 sites within nine categories (UNEP-WCMC, 2020). Colombia also has a large national system of indigenous and afro-descendant reserves with legally recognised collective land tenure and co-management strategies (MADS, 2017), plus much smaller public and private protected areas within regional and local systems. Some international conservation areas have been designated, including Important Bird Areas (IBAs) and RAMSAR sites. However, IPAs (or AIPs, Áreas de Importancia para la Conservación de Plantas in Spanish) are yet to be identified at the national level. A methodology for IPA identification in Colombia has been proposed (Diazgranados & Castellanos-Castro 2021). This methodology draws on global IPA criteria, with adjustments to account for the country’s rich diversity and the lack of reliable species distribution data. Identifying IPAs in Colombia may help promote the use of plant and fungi information in biodiversity conservation planning, which has traditionally focused on vertebrate species (Corlett, 2016). Additionally, useful plants can drive IPA identification based on the presence of threatened and/or range-restricted useful plant species under criterion A or sites holding concentrations of useful species under criterion B (Darbyshire et al., 2017). Such an approach would support the NSPC Action Plan, which recognises that continued efforts are required to meet targets on sustainable use. Conservation priority setting The global and national level structures described above can help to direct overarching conservation approaches. However, prioritisation is also important to ensure that resources and funds for conservation action are targeted effectively to prevent biodiversity loss (Wilson et al. 2009, Arponen, 2012). This prioritisation can be undertaken at different biological and geographical scales and can apply a range of methods. At the species level, conservation prioritisation has considered a range of factors such as evolutionary uniqueness and economic and cultural importance (Brehm et al., 2010, Arponen, 2012). Extinction risk assessments have also become one of the most often applied criteria for prioritisation, based on the urgency of action to avoid the loss of a species. The IUCN Red List of Threatened Species (IUCN, 2021) is the most comprehensive and recognised source on extinction risk, with replicated protocols to CATALOGUE OF USEFUL PLANTS OF COLOMBIA 37 CHAPTER 2 produce red lists at the national level. However, its limitations include taxonomic and geographic bias in the species assessed, with many gaps persisting in plant assessments (Nic Lughadha et al., 2020). Spatial approaches are also widely used in conservation priority setting to identify protected or other conservation areas, as well as biodiversity hotspots (Wilson et al., 2009). The principles of Systematic Conservation Planning (SCP) are often applied (Margules & Sarkar 2007). However, despite the gaps in extinction risk assessments, spatial prioritisation is often based on the presence of threatened species listed as Vulnerable (VU), Endangered (EN) or Critically Endangered (CR) on the IUCN and other red lists. MATERIALS AND METHODS Useful plants of conservation concern We compiled a preliminary list of useful plant species of conservation concern in Colombia based on their current extinction risk and geographical distribution. Uses were based on the Checklist of Useful Plants of Colombia (CUPC), produced as part of the Useful Plants and Fungi of Colombia project (UPFC, 2020), with nomenclatural reconciliation of plant names undertaken to match extinction risk assessments with the CUPC (Chapter 12). The CUPC was compiled from relevant databases, reports, and scientific literature, identifying 7,472 species within 2,140 genera and 258 families (Figure 1). Uses were grouped within the ten categories applied in the World Checklist of Useful Plant Species (Diazgranados et al., 2020), based on the ‘Level 1 States’ of the Economic Botany Data Collection Standard (EBDCS) (Cook, 1995). A stepwise process was used to identify species of conservation concern (Figure 1). First, only species that are native to Colombia were considered. Second, native species were only included if known to be: (i) internationally threatened; (ii) nationally threatened; or (iii) endemic. Native and endemic species were identified using the Colombian Catalogue of Plants and Lichens (CCPL) (Bernal et al., 2020) and Plants of the World Online (POWO, 2019). International and national conservation status was based on the IUCN Red List (IUCN, 2021) and on a list of national extinction risk assessments carried out by the IUCN Colombian Plant Specialist Group, respectively. We considered CR, EN or VU species as threatened. Additionally, as this study focuses on socially, economically, or culturally valuable species, IUCN Extinct (EX) or Extinct in the wild (EW) species were also included. Extinction risk assessment gaps Extinction risk assessments have not been undertaken for many plant species, representing gaps in our knowledge of their risk of extinction (Nic Lughadha et al., 2020). We calculated the proportion of Colombian useful plant species assessed at the international (IUCN Red List) or national level within each plant family and use category. We then used the Chi-square test and post-hoc analyses applying the standardised residual method (Beasley and Schumacker, 1995) to identify statistical variations in the proportion of species assessed in each family, thereby enabling families 38 with the largest species gaps in extinction risk assessments to be identified. Only families with 15 or more reported useful plant species were included in the analyses. Equivalent statistical analyses between use categories were not undertaken as species can fall within multiple categories. Distribution data for useful plants Existing public distribution data were used to identify hotspots for all known native useful plants of Colombia. All 5,830 native useful plants were queried against the Global Biodiversity Information Facility (GBIF) names backbone, and occurrence records linked to matched names and synonyms were downloaded from GBIF on 15 July 2021 (GBIF Occurrence Download, 2021). Data were cleaned to remove records with coordinates which were: outside Colombia; duplicated; assigned to country centroids or institutions; non-terrestrial; rounded to less than one decimal place; with equal or zero longitude and latitude; and within 1 km of the capital. This data cleaning resulted in 366,936 records for 5,241 native useful species (90.1% of total species). Identifying hotspots for useful plants To identify hotspots for useful plants, we defined our units of analysis (UA) based on ecological and political criteria. These were determined by splitting the 30 synthesised ecosystems defined by the Mapa de Ecosistemas Continentales Marinos y Costeros de Colombia (MADS et al., 2017) at the borders of Colombia’s 1,122 municipalities, resulting in 8,930 polygons. Occurrence records for native useful plants of Colombia (UPC) were joined to this multipolygon layer to enable analyses. This a priori regionalisation approach follows the proposed IPA methodology for Colombia (Diazgranados & CastellanosCastro 2021). While the alternative grid cell approach has been applied to IPA identification in other countries and has certain advantages (Hamidah et al., 2020), it makes it difficult to ensure comprehensiveness of ecosystem coverage – a key concept in Systematic Conservation Planning (Margules & Sarkar, 2007). We applied three methods for hotspot identification, with UAs falling under one or more of these definitions highlighted as hotspots for UPC. First, ‘national 5%’ hotspots are the UAs with the highest species richness and account for 5% of the total land area of Colombia. This method involved calculating the number of native UPCs recorded in each UA, sorting the UAs in decreasing order of richness, then selecting the UAs with the highest species richness until 5% of Colombia’s terrestrial area (c. 56,825 km2) was reached. This threshold is based on consistent findings that the richest 1–5% of the land area can represent a substantial proportion of species (Orme et al., 2005) and reflects the approach taken by Chi et al. (2017) to identify conservation priorities for medicinal plants in China. The next two methods took an ecosystem-level approach, further enabling a more comprehensive representation of the diversity of habitats in Colombia. ‘Ecosystem richest’ hotspots are defined as the UAs that support the highest species richness of each ecosystem type. This method is based on CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 2 criterion B1 for areas of exceptional botanical richness in the methodology for IPA identification in Colombia, with one UA per ecosystem type identified as a hotspot (Diazgranados & Castellanos-Castro 2021). Finally, ‘ecosystem 5%’ hotspots are the UAs with the highest species richness and account for 5% of the total land area of each ecosystem type. This method applied the ‘national 5%’ methodology at the ecosystem level, thereby combining the ecosystem-specific approach recommended by the IPA methodology with a threshold previously applied for hotspot identification at the national level in other countries (Chi et al., 2017). For the ecosystem-dependent analyses, all aquatic ecosystem types were combined, and ecosystems classified as ‘other areas’ and ‘no information’ were not included, resulting in a total of 19 ecosystem types (Figure 3). Spatial conservation gaps for useful plants To detect in-situ conservation gaps, all identified hotspots for UPC were overlaid with the distribution of protected areas and ICCA in Colombia. Hotspots were categorised depending on percentage overlap with these area-based designations: (1) high conservation gap (≤25% overlap); medium conservation gap (26–75% overlap); and low conservation gap (>75% overlap). Protected area data were downloaded from the World Database on Protected Areas (UNEP-WCMC, 2020) and include areas designated at the national, regional, and local levels. ICCA areas were downloaded from Agencia Nacional de Tierras (2017). Data reconciliation, preparation, and analyses were primarily conducted in R v.3.4.3 (R Core Team, 2020), using the packages dplyr v.1.0.6 (Wickham et al., 2021), rgbif v3.5.2 (Chamberlain et al., 2021), rgdal v1.5–23 (Bivand et al., 2021), and CoordinateCleaner (Zizka et al., 2019). Geographic analyses were conducted using a combination of R and ArcGIS Pro 2.8.0 (Esri Inc., 2021). RESULTS Useful plants of conservation concern and risk assessment gaps Native useful plant species in Colombia occurred across 232 families, with the most species-rich families being Fabaceae (530), Asteraceae (300), and Rubiaceae (257). We identified 544 useful plant species of conservation concern—as defined in Section 3.2.1—within 111 families. Of these, 294 species were identified as internationally or nationally threatened, while 250 unassessed species were included as endemics (Figure 1). As only 55% of UPC have FIGURE 1. Breakdown of orders of useful plants of Colombia and the stepwise process used to determine the species of conservation concern. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 39 CHAPTER 2 A B FIGURE 2. Proportion of useful plant species threatened at national and/or global levels or lacking extinction risk assessments across A families and B use categories. Numbers show total species per group, with some species in multiple-use groups. In A, only families that significantly contributed to the chi-square statistic (χ2), which tested differences in the proportion of species assessed, are shown, with remaining families grouped in ‘other’; * indicates families that are significantly under represented in Red Lists. 40 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 2 FIGURE 3. Ecosystem map of Colombia (MADS et al., 2017) and distribution of native useful plants of Colombia, based on occurrence points downloaded from GBIF (GBIF Occurrence Download, 2021). CATALOGUE OF USEFUL PLANTS OF COLOMBIA 41 CHAPTER 2 TABLE 2. Hotspots for useful plant species of Colombia by designation type and percentage overlap with existing protected areas (PA) or Indigenous and Community Conserved Area (ICCA). Hotspot type Number of UAs Total area (km2) Native UPC recorded PA overlap (%) ICCA overlap (%) PA or IR overlap (%) National 5% 3 52,384 1,644 44.3 26.3 70.5 Ecosystem 5% 76 77,593 4,039 44.4 16.5 56.6 Ecosystem richest 19 60,074 2,894 49.1 20.1 69.1 All hotspots 78 89,901 4,134 36.5 24.2 60.4 A 0 150km B 0 150km been assessed, the list of taxa under conservation concern is preliminary and could be extended in the future. A total of 151 families supported fewer than 15 native UPC and were not included in the following analyses. The proportion of useful plant species with extinction risk assessments at the national or international level was unevenly distributed across families (χ 2 = 1213, df = 94, p < 0.01). On average, 48.7% of species were assessed per family, with 28 families significantly contributing to the unevenness of coverage in risk assessments (χ 2 post hoc analysis based on standardised residuals p < 0.01). Thirteen of those had a significantly smaller proportion of species with extinction risk assessments than other families, while the rest had a significantly higher proportion of species assessed (Figure 2A). The 13 families highlighted in Figure 2 with a lower proportion of assessments can be used as a basis to prioritise UPC for further extinction risk analysis. Variation was also seen between the Level 1 use categories. Species with environmental and medicinal uses were under represented in extinction risk assessments, with 42% and 47% assessed, respectively. This scenario was in contrast to species in the fuels, invertebrate food, and materials categories, which had levels of assessment that were higher than the average (73%, 62% and 61% assessed, respectively). As many species occur in multipleuse categories, variables are not mutually exclusive, with the Chi-square test of significance not applied. Geographic hotspots and conservation gaps Records of native useful plants of Colombia were distributed across 967 municipalities (Figure 3), with Solano (Caquetá department) found to support the greatest diversity. Hotspots were distributed across all ecosystem types, with agroecosystems (4,523 species) and forest habitats supporting the most species (4,405). In total, 78 UAs were identified as hotspots for UPC (Figure 4a), classified under one or more of the three definitions applied, and covering a total area of 89,901 km2 across 57 municipalities. Hotspots had a mean size of FIGURE 4. A Units of analysis identified as hotspots for useful plant species in Colombia, and the hotspot definitions they fall under. B Percentage overlap of hotspots with protected areas or Indigenous and Community Conserved Areas (ICCAs). 42 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 2 1,152.6 km2, ranging from 2.5 km2 (swamp habitat in Santa Fé de Bogotá municipality, Cundinamarca) to 40,076.5 km2 (forest habitat in Solano municipality). The identified hotspots are known to support 4,134 native UPC (70.9% of total) while covering 7.9% of the country’s terrestrial land area. Sixty-eight of the hotspots include records for UPC that are of conservation concern (threatened and/or endemic; Figure 1), including a total of 353 species (64.9% of all UPC of conservation concern; Figure 1). This includes 67 species listed as VU on the IUCN Red List, 41 listed as EN, 11 species assessed as CR, four species categorised as EW and EX (two species in each category), and 226 endemic species to Colombia. However, only 36.5% of the total hotspot area is currently included in the protected areas system of Colombia, with 24.2% represented in ICCAs (Table 2). Hotspots were ranked for their conservation priority based on the percentage gap in coverage by existing protected areas or ICCAs (Figure 4B). Forty-seven of the hotspots (16,775 km2) overlap with existing designated areas by less than 25% and are therefore considered as high conservation priority. These hotspots include ten different ecosystem types distributed across 37 municipalities. Thirteen hotspots (17,870 km2) overlap with designated areas by more than 75%, while the remaining 18 hotspots (55,256 km2) are considered as medium priority. DISCUSSION Despite plant conservation efforts, studies indicate that targets for useful wild plant species have not been met, and overexploitation continues to be a major driver of extinction (Khoury et al., 2019). Based on the Checklist of Useful Plants of Colombia, we present priorities to guide two key actions to contribute to conservation in the country: extinction risk assessments at the species level and area-based management at the ecosystem level. To inform this process, we compiled a list of useful plant species of known conservation concern in Colombia, highlighted families and use groups that are underrepresented in extinction risk assessments and identified geographic hotspots and area-based conservation gaps for useful plant species in Colombia. These are important first steps in the long-term conservation and sustainable use of plants in one of the most biodiverse countries in the world (UN-WCMC, 2014). Useful plants of conservation concern and their risk assessments We identified 544 UPC that are known to be endemic and/ or threatened at national or global levels. However, with 45% of native useful plant species not represented in either the Colombian or IUCN Red Lists (Figure 1), many more UPC may be at risk of extinction. This level of extinction risk assessment is greater than the average coverage of global assessments across plant species (roughly 30%), reflecting findings that useful plants are over represented in red listing (Nic Lughadha et al., 2020). However, it is important to fill the remaining gaps, as extinction risk assessments often form the basis of conservation policy, planning, and action. Gaps in assessments may therefore lead to key species being ignored in conservation efforts, including spatial conservation prioritisation. Undertaking risk assessments is resource-intensive, requiring a high level of specialist knowledge and robust data. To target limited conservation resources efficiently, our analyses highlighted 13 families in which significantly fewer useful plant species have been assessed (Figure 2). This count includes some of the most species-rich groups both in Colombia and globally, such as Asteraceae and Poaceae. Meanwhile, at the level of use categories, species with medicinal and environmental uses were found to be the most under represented in extinction risk assessments – two of the top three most species-rich categories in the country. We propose that these families and use groups should be considered as priorities for ongoing extinction risk assessments in Colombia with a view to informing subsequent conservation prioritisation. While red listing is an important first step for prioritisation, extensive further work across disciplines is required to achieve sustainable use and management at species- and location-specific levels. This includes developing knowledge of plant biology and harvesting practices; understanding relevant economics and market pressures; recognising and supporting land rights holders and enabling management which facilitates responsible use; building capacity and providing resources for sustainable harvesting practices; and developing institutional structures and policies that support such practices (Kor et al., 2021). Geographic hotspots and conservation gaps for useful plants In total, 78 hotspot UAs for useful plants of Colombia were highlighted, falling under any one of three hotspot definitions applied. These areas should be prioritised for conservation measures that incorporate sustainable plant use and provide a first step towards identifying a national IPA network for socially, economically, and culturally valuable species. By focusing on useful species, this study supports national priorities to integrate biodiversity conservation into Colombia’s post-conflict development and the NSPC Action Plan, whose main objectives include sustainable use and protecting useful plant species (Castellanos-Castro et al., 2017). Additionally, it supports global conservation efforts (CBD, 2021). A recent assessment of the GSPC’s progress highlighted that while there have been many successes in global plant conservation, the targets where the least progress have been made include in-situ conservation and sustainable use (Sharrock, 2020). CATALOGUE OF USEFUL PLANTS OF COLOMBIA 43 CHAPTER 2 In-situ conservation gaps Our results showed that UPC are not well represented in Colombia’s current protected area system, with just 36.5% of the total hotspot area covered by local, regional, or national PAs in the country (UNEP-WCMC, 2020). Adding ICCAs to this analysis lessoned the conservation gap, but 48 of the 78 hotspot UAs remained as high priorities, with less than 25% of their land area occurring within any designated areas. While indigenous reserves and afro-descendant territories are primarily designated for recognising and protecting ethnic and biocultural diversity, they present great relevance to biodiversity conservation, with many occurring in ecosystems of rich biodiversity and 62% of land use associated with forest protection (Arango, 2017). Conservation areas focusing on useful plants remain scarce globally. However, their potential has been shown in Colombia, with the Santuario de Flora Plantas Medicinales Orito Ingi-Ande national park designated as the first and only protected area for medicinal plants in the country (IUCN, 2008). It is located in territories used by the Kofanes community in southwestern Colombia, who first proposed the concept. Species characterisation was undertaken by them in collaboration with a wide range of stakeholders over several years. The hotspots presented in this chapter are therefore only the first step in the process of conservation designation, and designation is by no means a guarantee of sustainable plant use, with a multitude of social, political, and economic, as well as further ecological considerations, required. Additionally, data challenges arose which reflect those commonly met in conservation planning. Data and methodological limitations Our analyses were based on existing species occurrence records available through large, global datasets (GBIF Occurrence Download, 2021). However, records for many of the priority UPC were scant or missing altogether. Additionally, the geographical distribution of occurrence records was unequal, favouring ecosystems such as agroecosystems and forests, which are more easily accessible or of greatest economic and research interest. The high occurrence of useful plant species in agroecosystems (Section 3.3.2) may also result from historic records that are no longer present following land-use conversion. As a result, while many UPC are not represented in the hotspots identified, other useful species reported as present may no longer be in the locations highlighted, with targeted ground-truthing required in some areas before designating conservation protection. This scenario is exacerbated by the uncertainty associated with occurrence information, which tends to lack taxonomic verification. Therefore, the data used may also include misidentified species, with case studies finding that rates of species misidentification in plant occurrence information can range from <1% to 17% (Meyer et al., 2016). These biases, gaps and uncertainties are common challenges that have affected conservation planning globally, including IPA networks (Blasi et al., 2011, Hamidah et al., 2020). 44 Further conservation action and prioritisation based on the high-level gaps identified in this chapter should include wider sources of data, such as herbarium records and geographic data from published literature. In addition, to improve data availability more generally, it is important to continue supporting national and regional herbaria in increasing the digitisation and public accessibility of their records, which can include important specimens and provide local data for conservation planning and a wide range of other research (Sweeney et al., 2018). These actions are a resource-intensive procedure and require constant curation to ensure that high-quality data are provided digitally. Supporting participatory research can also help document the vast amount of traditional knowledge held by local experts that should form the basis for conservation decision-making. In addition to the data challenges, the methodologies applied for hotspot identification represent just some of many potential approaches, each with associated benefits and limitations. While we applied three methods, all were based on measures of overall richness of native useful plant species in Colombia. These methods did not account for species’ severity of extinction risk within the UAs, with 32% of UPC of conservation concern (Figure 1) not represented within the hotspots. This approach was partly chosen due to current gaps in extinction risk assessments. However, future analyses could instead focus on hotspots for species of conservation concern, in line with Criterion A of the IPA methodology, which focuses on the presence of threatened and/or range-restricted plant species (Darbyshire et al., 2017). In addition, the UAs used were of varying size, which can create a bias towards larger areas due to the fundamental species-area relationship. Alternative approaches applying grid cells as UAs (e.g., Hamidah et al., 2020) or accounting for ratios such as species density per area and species numbers per record could inform further analyses and provide useful comparisons to the approach taken in this chapter. From conservation priorities to action This chapter presents the first steps in identifying gaps and suggesting priorities for the conservation of useful plant species in Colombia. However, undertaking extinction risk assessments and identifying geographic hotspots for useful plants are not sufficient to achieve their conservation. Other important steps must be taken to transform the hotspots identified into locally relevant, sustainably managed areas and to use information from extinction risk assessments to inform effective conservation prioritisation and action. Crucially, a wide range of socio-economic considerations must be determined in collaboration with a range of stakeholders, including levels of government, land right-holders, and community members, to ensure that overexploitation of useful species does not occur (Margules and Sarkar, 2007). Social-Ecological Systems (SES) science is increasingly used as a framework to support transdisciplinary approaches in environmental management and biodiversity CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 2 conservation (Virapongse et al., 2016). Such an approach is particularly important when considering useful plants, whose sustainable use and conservation require the involvement of resource users and is dependent on market forces and national policies (Kor et al., 2021). Simultaneously, linking ethnobotany and conservation enables the recognition and incorporation of local, traditional, and indigenous knowledge in biodiversity management and can support the conservation of ‘knowledge networks’ and cultural heritage. In-depth botanical and ecological knowledge data are also required to strengthen the definition of IPA boundaries and to enable the development of effective management plans for the sustainable use of species. Despite including both ecosystem and municipality boundaries in our definition of UAs, some identified hotspots were extremely large (Section 3.3.2), with further geographic refinement required to determine feasible units for conservation management. These hotspots should be supported by targeted field data collection and consultation with experts. Meanwhile, ex-situ conservation is also vital to the long- term survival of threatened species, providing insurance against extinction and the potential for species restoration and reintroduction. However, global gaps currently exist in the representation of useful plants in such repositories (Khoury et al., 2019). Interdisciplinary research projects such as the UPFC, which unite a diverse range of knowledge bases through collaboration with local communities, alongside the involvement of researchers from multiple disciplines, are significant contributors to this process (UPFC, 2020). We believe that through sustained collaborative approaches, the conservation priorities presented in this study can act as a crucial starting point for useful plant conservation action in Colombia, and can enable participatory approaches to identify IPAs. While published conservation criteria and available conservation tools provide invaluable guidance for planning and prioritisation, stakeholder engagement is ultimately crucial to develop a realistic, practical, and sustainable network of sites that will result in effective conservation on the ground. Participatory mapping to identify local conservation priorities in the Municipality of Becerril. Mauricio Diazgranados CATALOGUE OF USEFUL PLANTS OF COLOMBIA 45 CHAPTER 2 Acknowledgments This work directly contributes to the ‘Useful Plants and Fungi of Colombia’ project, supported by a Professional Development & Engagement grant under the Newton-Caldas Fund partnership. The grant is funded by the UK Department for Business, Energy and Industrial Strategy (BEIS) and Minciencias, and delivered by the British Council. LK is supported by a studentship awarded by the Natural Environment Research Council [Grant Number NE/ S007229/1]. References Agencia Nacional de Tierras (2017). 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CATALOGUE OF USEFUL PLANTS OF COLOMBIA 47 CHAPTER 2 Espeletia killipii var. chisacana – medicinal endemic species from Colombia, occuring in Páramos. 48 Mauricio Diazgranados CATALOGUE OF USEFUL PLANTS OF COLOMBIA Chapter 3 Endemic Medicinal Plants of Colombia Henry Yesid Bernal M.1* & Catalina Mesa Sánchez2 1 2 Ex-Profesor Asociado del Departamento de Biología, Facultad de Ciencias, Universidad Javeriana, Herbario de la Pontificia Universidad Javeriana (HPUJ), Bogotá, D.C., Colombia. Bióloga, M.Sc. en Gestión Ambiental, Facultad de Ciencias, Departamento de Biología, Pontificia Universidad Javeriana, Bogotá, D.C., Colombia. *Corresponding author: henryesid@gmail.com Keywords: useful plants, ethnobotany, phytopharmaceuticals, phytomedicines, phytotherapeutic products ABSTRACT The medicinal species are highlighted from the Colombian biodiversity and its useful plants, including many native and a significant number of species endemic to the country. However, the medicinal species of Colombia are still underused or poorly known, with several of them already at risk of extinction. Medicinal species constitute a true natural heritage that requires concrete and innovative actions for their knowledge, conservation, and sustainable use. This study was carried out between 2014 and 2021, with the main goals to: a) review and update the inventory of medicinal plants used in Colombia (exotic and native species); b) specify which are the medicinal species endemic to Colombia; and c) evaluate the current state of traditional, scientific, and technological knowledge of the endemic species to Colombia used as medicinal and their potential contribution to the treatment of prevalent pathologies in the country. A total of 3,005 medicinal plants used in Colombia were identified and documented, of which 1,719 are native to this country (including 204 endemics), 558 exotic, and 728 species with an uncertain geographical distribution. The lack of knowledge on conservation and sustainable use of most of the Colombian endemic medicinal species was verified, and their traditional, scientific, and technological knowledge was characterised. Only three species have the traditional and scientific knowledge classified as ‘acceptable’: Achyrocline bogotensis (Asteraceae), Espeletia killipii (Asteraceae) and Pentacalia corymbose (Asteraceae). Another ten species have evidence of traditional knowledge classified as ‘acceptable’ and evidence of scientific knowledge classified as ‘regular’: Berberis goudotii, Berberis rigidifolia (Berberidaceae), Cecropia telealba, Cecropia mutisiana (Urticaceae), Chromolaena barranquillensis (Asteraceae), Espeletia grandiflora, Espeletiopsis muiska (Asteraceae), Ocotea caparrapi (Lauraceae), Passiflora antioquiensis (Passifloraceae) and Senecio niveoaureus (Asteraceae). Those 13 species present one or several mentions of uses for different pathologies with higher impact in Colombia. RESUMEN Dentro de la biodiversidad de Colombia y sus plantas útiles se destacan las especies medicinales, siendo muchas nativas y dentro de ellas un buen número son endémicas del país, pero son subutilizadas o poco conocidas y varias de ellas están en peligro de extinción. Se considera que estas especies medicinales constituyen un verdadero patrimonio natural que requiere de acciones concretas e innovadoras para su conocimiento, conservación y uso sostenible. Entre los años 2014 y 2021 se desarrolló esta investigación que tuvo como objetivos: a) revisar y actualizar el inventario de las plantas medicinales de uso en Colombia (exóticas y nativas); b) precisar cuáles son las plantas medicinales endémicas del país y c) evaluar el estado actual de los conocimientos tradicional, científico y tecnológico de las plantas medicinales endémicas de Colombia y su posible contribución para la atención de las patologías prevalentes en el país. En esta investigación se identificaron y documentaron 3.005 plantas medicinales de uso en Colombia de las cuales 1.719 son nativas de Colombia (204 endémicas), 558 son foráneas y 728 tienen distribución geográfica incierta. Se comprobó la falta de conocimientos para la conservación y uso sostenible de la mayoría de las especies medicinales endémicas de Colombia, y se logró la caracterización de sus conocimientos tradicional, científico y tecnológico. Solo 3 especies cuentan con conocimiento tradicional y científico clasificados como ‘aceptable’ Achyrocline bogotensis (Asteraceae), Espeletia killipii (Asteraceae) y Pentacalia corymbosa (Asteraceae), y 10 especies tienen evidencia de conocimiento tradicional clasificada como ‘aceptable’ y evidencia de conocimiento científico clasificada como ‘regular’ Berberis goudotii (Berberidaceae), Berberis rigidifolia (Berberidaceae), Cecropia telealba (Urticaceae), Cecropia mutisiana (Urticaceae), Chromolaena barranquillensis (Asteraceae), Espeletia grandiflora (Asteraceae), Espeletiopsis muiska (Asteraceae), Ocotea caparrapi (Lauraceae), Passiflora antioquiensis (Passifloraceae) y Senecio niveoaureus (Asteraceae). Estas 13 especies presentan uno o varias menciones de uso para las diferentes patologías de mayor impacto en Colombia. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 49 CHAPTER 3 INTRODUCTION Since ancient times, all societies have resorted to plants as a source of medicines. According to the World Health Organization (WHO, 2021), 80% of the world’s population use traditional medicine to meet their needs for primary health care. It is recognised that traditional medicine and medicinal plants are the main important ways to improve health for the planet’s population. Several guidelines are found in a significant number of publications (WHO, 2021). The conservation of biodiversity at the global level is a crucial issue for future possibilities for adapting living beings in a world of continuous change. The undoubted medicinal resource that plants constitute is under threat and is a treasure that cannot go to waste (Bernal, 2012). Many initiatives on biodiversity continue to register and count species, but there is a big gap in the associated traditional, scientific, and technological knowledge needed to conserve and use biodiversity sustainably. The humid tropics in Latin American countries, especially Colombia, are a stripe of intense biological activity but low economic production of natural products. Several conditions can explain this contradiction, such as poor soils, excessive rainfall, plant and animal diseases, inadequate agricultural systems, physical structural deficiency, and lack of medium- and long-term investments. Paradoxically, Colombian biodiversity is considered megadiverse, and it is estimated that in terms of plant species, the country comprises between 30,000 and 41,000 species (Andrade, 2011). These figures place Colombia as one of the richest floras in the world in species number, which are mainly underutilised, underexploited, and poorly known in terms of their conservation and economic potentialities. The great importance of environmental goods and services derived from biodiversity must be highlighted. The survival of humanity relies on biodiversity as a great source of food, fossil fuels, natural fibres, phytomedicines, biocides, natural colourants, soil productive capacity, water and air cycling, and many other environmental services. Likewise, biodiversity is the world’s natural capital. In particular, the native and endemic species of megadiverse countries such as Colombia constitute its greatest biological heritage and represent fundamental options for equitable, democratic, and sustainable development (Bernal, 2012). The main mortality-causing pathologies in Colombia are ischemic heart diseases, cerebrovascular diseases, chronic diseases of the lower respiratory tract, diabetes mellitus, acute respiratory infections, hypertensive diseases, malignant tumours of the stomach, trachea, bronchi, and lungs, and heart failure (Bernal et al., 2011). Similarly, in descending order, the main morbidity-causing pathologies in Colombia are periodontal disease, dental caries, iron deficiency anaemia, asthma, trichuriasis by Trichuris trichiura, hypertensive heart disease, proteincalorie malnutrition, hookworm, abortion, unipolar major depression, and diarrheal diseases (Bernal et al., 2011). 50 The main pathologies on epidemiological surveillance in Colombia are classical dengue fever, haemorrhagic dengue fever, leishmaniasis, Plasmodium vivax and P. falciparum malaria, and tuberculosis (Bernal et al., 2011). In Colombia, important ethnobotanical, phytochemical, and pharmacological investigations have been carried out on the use of medicinal plants as a support treatment for several tropical pathologies. Regarding the inventory and the state of traditional, scientific, and technological knowledge of medicinal plants used in the country, it is worth mentioning the studies produced by: a) Enrique Pérez Arbeláez (1953–1966, 1978a,b); b) Victor Manuel Patiño (1967); c) Hernando García Barriga (1974, 1975 and 1992); d) José Joaquín Montes Giraldo (1981); e) Daniel J. González Patiño (1984); f) Mauro Hernández Mesa (1992); g) Ramiro Fonnegra et al. (Fonnegra & Jiménez, 1992, Fonnegra et al., 2006); h) the Ibero-American Program of Science and Technology for Development (CYTED), Subprogram of Fine Pharmaceutical Chemistry and the International Organization of the Andrés Bello Agreement from the publication “Promising plant species from the Andrés Bello Agreement countries” (Bernal & Grupta 1995, ; Correa & Bernal, 1989–1995); and i) other Ibero-American medicinal plants studies (Bernal & Correa, 1995; Martínez et al., 2000; Bernal & Jiménez, 2008). In 2008, the Ministry of Social Protection of Colombia published the first edition of the Colombian Vademecum of Medicinal Plants and updated guidelines, from which 127 plant species used as medicinal in Colombia were regulated (including 87 Neotropical species that are not native to Colombia and only two endemic species – Achyrocline bogotensis and Cecropia mutisiana) (Ministerio de Protección Social, 2008). More recently, Bernal et al. (2011) published a book entitled ‘Guidelines for Knowledge, conservation, and sustainable use of native medicinal plants of Colombia’. The inventory from this book was carried out by Vásquez & Bernal (2011), recording 2,404 medicinal plant species used in Colombia, including 1,870 native, 214 endemic, and 534 exotic species. In 2021, Bernal & Gómez prepared the second edition of the project “National collection of medicinal plants”, which is a list of the endemic medicinal plants of Colombia and the native medicinal species of Colombia that have been prioritised for cultivation at the Quindío Botanical Garden (Calarca, Colombia). The data and results presented here are part of a project developed between 2014 and 2021 from extensive previous studies on the medicinal plants of Colombia (Bernal & Mesa 2014). This chapter has as main goals: a) to review and update the inventory of medicinal plants used in Colombia (exotic and native); b) to specify which endemic species are used as medicines in Colombia; and c) to evaluate the current state of traditional, scientific, and technological knowledge on endemic species used as medicines and their possible contribution to the treatment of the prevalent pathologies in Colombia, according to data on mortality, morbidity, and diseases from epidemiological surveillance. CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 3 TABLE 1. National and international databases consulted. Database name BHL Biodiversity Heritage Library BIOVIRTUAL (common names). Instituto de Ciencias Naturales-Universidad Nacional de Colombia BIOVIRTUAL. Ejemplares de herbario Instituto de Ciencias Naturales-Universidad Nacional de Colombia Catálogo de plantas y líquenes de Colombia URL https://www.biodiversitylibrary.org/ http://www.biovirtual.unal.edu.co/ nombrescomunes http://www.biovirtual.unal.edu.co/ICN/ http://catalogoplantasdecolombia.unal. edu.co/es https://www.cbif.gc.ca/acp CBIF. Canadian Biodiversity Information Facility CEIBA. Catalogador de información biológica. Instituto de Investigación de http://www.i2d.humboldt.org.co/ceiba Recursos Biológicos Alexander von Humboldt-Colombia http://www.ciencias.bogota.unal.co/menuColombia diversidad biótica. Instituto de Ciencias Naturales de la Universidad principal/publicaciones/biblioteca-digital/ Nacional de Colombia Colombia-diversidad-biotica CoL. Catalogue of Life https://www.catalogueoflife.org ColPlantA Colombian Resources for Plants made Accessible https://colplanta.org DR. DUCKE’s. Phytochemical and Ethnobotanical Databases http://www.ars-grin.gov/duke EOL. Encyclopaedia of Life. National Museum of Natural History. Smithsonian http://www.eol.org/ http://www.ciencias.bogota.unal.edu.co/ Flora de Colombia. Monografías del Instituto de Ciencias Naturales de la menu-principal/publicaciones/bibliotecaUniversidad Nacional de Colombia digital/flora-de-colombia Flora de la Real Expedición Botánica. Real Jardín Botánico de España https://www.bibdigital.rjb.csic.es/medias GBIF. Global Biodiversity Information Facility http://www.gbif.org/species Global Compositae database. International Compositae Alliance https://compositae.org/ GRIN. Germplasm Resources Information Network http://www.ars-grin.gov Ildis database. International Legume Database and Information Service http://www.ildis.org/ IPNI. International Plant Name Index https://www.ipni.org/ IUCN-Red List. The IUCN red list of the threatened species https://www.iucnredlist.org/species ITIS. Integrated Taxonomic Information System http://www.itis.gov JSTOR Global Plants database https://www.plants.jstor.org/ Kew herbarium catalogue. Royal Botanic Gardens, Kew http://www.apps.kew.org/herbcat/ Kew Plant Information Portal. Royal Botanic Gardens, Kew https://www.epic.kew.org/ Napralert Database. University of Illinois at Chicago http://www.napralert.org/ NCBI. National Centre for Biotechnology Information. National Library of https://www.ncbi.nlm.nih.gov/ Medicine. National Institutes of Health. USA NMNH. Smithsonian National Museum of Natural History https://www.collections.nmnh.si.edu/ PL@NTUSE. Usos de las plantas https://use.plantnet-project.org RACCEFYN. Revista de la Academia Colombiana de Ciencias Exactas, https://raccefyn.co Físicas y Naturales REFLORA. Virtual Herbarium from the Flora of Brazil. Rio de Janeiro Botanic Garden. http://reflora.jbrj.gov.br/ Science Direct https://www.sciendedirect.com/ Scopus-Elsevier. Bibliographic database http://www.scopues.com SIAC. Sistema de información ambiental de Colombia http://www.siac.gov.co/biodiversidad SIB. Sistema de Información sobre la Biodiversidad de Colombia http://www.dta.sibcolombia.net/inicio.htm SPECIES 2000 and ITIS. Catalogue of life. Annual checklist. Integrated Taxonomic http://www.catalogueflife.org/ Information System The Plant List http://www.theplantlist.org TROPICOS. Vascular Plants of the Americas http://www.tropicos.org/project/VPA USDA-Plants Database. United States Department of Agriculture http://www.plants.usda.gov Useful Tropical Plants Database https://www.tropical.the ferns.info WCSP. World Checklist of Select Plant Families. Royal Botanic Gardens, Kew https://www.wcsp.science.kew.org/ WCVP. The World Checklist of Vascular Plants. Royal Botanic Gardens, Kew https://www.wcvp.science.kew.org/ WFO. World Flora Online Consortium http://www.floraonline.org CATALOGUE OF USEFUL PLANTS OF COLOMBIA 51 CHAPTER 3 MATERIALS AND METHODS This study used a revised and updated version of the list of medicinal plants used in the country published by Vásquez & Bernal (2011, pages 165 to 231) as a starting point to identify the endemic medicinal plants of Colombia. Specialised bibliographic searches were carried out at the national and international levels covering the period from 2014 to 2021, and the corresponding databases were built. A total of 13 Colombian herbaria were consulted: Herbarium of the Technological University of Chocó (CHOCO); Herbarium of the Eloy Valenzuela Botanical Garden (CDMB); Colombian Amazon Herbarium; Amazon Institute for Scientific Research, Sinchi (COAH); Colombian National Herbarium (COL); Herbarium of the University of Antioquia (HUA); Enrique Forero Herbarium of the University of the Amazon (HUAZ); Herbarium of the University of Córdoba (HUC); Herbarium of the Universidad Católica de Oriente (HUCO); Herbarium of the University of Quindío (HUQ); Herbarium of the Botanical Garden of Bogotá “José Celestino Mutis” (JBB); Herbarium of the Guillermo Piñeres Botanical Garden (JBGP); Herbarium of the Juan María Céspedes Garden. (TULV); and Herbario Forestal Gilberto Emilio Mahecha Vega of the District University (UDBC). Colombian and international databases were also consulted (Table 1), aiming to define the geographical distribution of medicinal species to Colombia, from which the species were grouped into three categories: a) restricted geographical distribution in one or two departments of Colombia, b) wide geographical distribution in three or more departments of Colombia, and c) uncertain geographic distribution in Colombia. The current states of traditional, scientific, and technological knowledge of the endemic species of Colombia used as medicinal were based on the studies carried out by Mesa (2013). Theoretical variables were defined for each species to analyse the current state of knowledge (based on ethnobotanical, scientific, and technological evidence). Indicators were established for each of the variables according to the following variation scale: 5–7 (acceptable), 3–4 (regular), and 0–2 (scarce) with scores that allow the definition of the current state of knowledge for species (Tables 2, 3, and 4). The indicators of the evidence of traditional knowledge were: a) historically supported use based on reliable documentary references; b) pathologies prevented or treated by the used species; c) mentions of therapeutic uses in humans or animals; d) safety (no history of toxicity according to the ethnobotanical literature review); e) part(s) of the plant that are used for therapeutic purposes in humans or animals; f) traditional way of preparation; g) posology; h) dosage; i) topical administration route (external use); and j) systemic administration route (internal use) (Table 2). The indicators of evidence of scientific knowledge were a) description and taxonomic diagnosis; b) morphology; c) geographical and altitudinal distribution; d) ecology; e) phytochemical profile; f) molecular characterisation; g) extraction and separation of active ingredients; h) identification of compounds and determination of their molecular structures; i) acute toxicity; j) subacute toxicity; k) chronic toxicity; l) subchronic toxicity; m) in vitro biological activities; n) biological activities in animal models; o) phase I clinical studies; p) phase II clinical studies; q) phase III clinical studies; and r) phase IV clinical studies (Table 3). The indicators of evidence of technological knowledge were a) micropropagation; b) sexual propagation; c) asexual spread; d) soil preparation; e) planting and spacing; f) fertilisation TABLE 2. Operationalisation for the theoretical variable ‘ethnobotanical evidence’ related to endemic species of Colombia used as medicinals. Factor Traditional knowledge 52 Theoretical variable Concept Indicators Scale Ethnobotanical evidence Ethnobotanical evidence that an endemic plant species is used as medicinal in Colombia (in this study) is based on its cultural therapeutic use to treat or prevent diseases in humans or animals that mainly indigenous and local communities give to these species. It is based on the reliable bibliographic documentation available in the national and international literature. • Use historically supported by reliable bibliographic references • Pathologies that are prevented or treated • Mentions of therapeutic uses in humans or animals • Safety (no history of toxicity according to the available botanical bibliography) • Part(s) of the plant that are used for therapeutic purposes in humans or animals • Traditional preparation method • Posology/dosage • Dose • Topical route of administration (external use) • Systemic route of administration (internal use) • Acceptable • Regular • Scarce CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 3 TABLE 3. Operationalisation for the theoretical variable ‘scientific evidence’ related to endemic species of Colombia used as medicinals. Factor Scientific knowledge Theoretical variable Scientific evidence Concept Indicators The scientific evidence related to endemic species of Colombia used as medicinals (in this study) is based on the current state of taxonomic, geographical, ecological, and phytochemical knowledge, toxicity, biological and pharmacological activities, and health studies (preclinical and clinical). It is based on the reliable bibliographic documentation available in the national and international literature. Scale Taxonomic characterisation: • Valid scientific name • Synonyms • Infraspecific categories • Cultivars • Taxonomic description. • Taxonomic diagnosis • Habit (life form) • Annual-perenniality data • Morphological studies • Molecular studies • Methods for the identification and taxonomic determination of species and infraspecific categories • Methods for the identification and taxonomic determination of cultivars Geographical and ecological characterisation: • Geographical distribution • Altitudinal distribution • Ecological studies (habitat, phenology, natural regeneration, populations, communities, plant-animal relationship, conservation status, conservation measures, etc.) Phytochemical characterisation: • Phytochemical workflow [extraction, fractionation, or separation, qualitative characterisation (preliminary • Acceptable or qualitative phytochemical workflow), isolation and • Regular purification (preparative or quantitative phytochemical • Scarce workflow)] • Active compounds recognition • Identification of compounds and determination of molecular structures (structural interpretation) Characterisation of innocuousness/safety: • Acute toxicity • Subacute toxicity • Chronic toxicity • Subchronic toxicity • Contraindications and precautions • Interactions with other drugs Preclinical characterisation: • Biological and pharmacological activities in vitro • In vivo biological and pharmacological activities in animal models • Existence of studies on pharmacodynamics • Existence of studies on pharmacokinetics • Existence of studies on the kinetics Clinical characterisation: • Existence of Phase I sanitation/health studies • Existence of Phase II sanitation/health studies • Existence of Phase III sanitation/health studies • Existence of Phase IV sanitation/health studies CATALOGUE OF USEFUL PLANTS OF COLOMBIA 53 CHAPTER 3 TABLE 4. Operationalisation for the theoretical variable ‘technological evidence’ related to endemic species of Colombia used as medicinals. Factor Technological knowledge Theoretical variable Technological evidence Concept The technological evidence related to endemic species used as medicinal in Colombia (in this study) is based on the current state of agrotechnological knowledge and the technologies for preparing medicinal products made from them. It is based on the reliable bibliographic documentation available in the national and international literature. Indicators Agrotechnologies for organic and sustainable cultivation: • Micropropagation • Sexual propagation • Asexual propagation • Soil preparation • Planting and spacing • Fertilisation and supply of nutrients • Organic farming and agroforestry (monoculture or polyculture) • Cultural work and plant care • Diseases and treatment • Harvest and collection of the part(s) of the plant used as medicine • Acceptable • Postharvest • Regular • Performance • Scarce • Costs of organic and sustainable production Technologies used in the elaboration of medicinal products: • Technologies used in the extraction of plant raw material • Technologies used in the pharmaceutical formulation of medicinal products • Technologies for the use of plant-based excipients and auxiliary formulation to produce medicinal products • Quality control of medicinal products (thin-layer chromatography and high-performance liquid chromatography) and nutrient supply; g) organic cultivation and agroforestry (monoculture or polyculture); h) cultural work and plant care; i) diseases and treatments; j) harvest and collection of parts for medicinal use; k) postharvest; l) performance; m) costs of organic and sustainable production; n) technologies for the extraction of vegetable raw materials; o) technologies for pharmaceutical formulation; p) technologies for the use of excipients and formulation aids; and q) quality control of medicinal products (Table 4). The current state of traditional, scientific, and technological knowledge of the endemic species used as medicinals was based on the variables, indicators, scale, and scores indicated above, and defined according to five steps: 1) the endemic species used as medicinals were listed under ‘acceptable’, ‘regular’, or ‘scarce’ evidence for each of the three types of knowledge (traditional, scientific, and technological); 2) the endemic species used as medicinals that were associated with acceptable evidence of scientific and traditional knowledge were selected, and their geographical distributions in Colombian departments were presented; 3) species associated with acceptable evidence 54 Scale of traditional knowledge, but also with regular evidence of scientific knowledge, were selected and the same procedure was carried out; 4) species with acceptable evidence of scientific and traditional knowledge and acceptable or regular evidence of technological knowledge were analysed; and 5) a matrix was elaborated to compare pathologies of greater impact in the country in terms of mortality, morbidity and pathologies under epidemiological surveillance versus mentions of therapeutic uses considering the list of endemic medicinal species of Colombia. RESULTS In this study, we identified 3,005 plant species that have medicinal uses (2,847 spermatophytes and 158 species of mosses, hornworts, liverworts, or lycophytes) in Colombia, with 1,719 being native to this country (including 204 endemics), 558 exotics, and 728 with uncertain geographical distribution. Almost the totality of Colombian endemic species used as medicinals is represented by flowering plants (133 species) distributed into 50 families, with a single species of Lycophytes reported (Selaginella rosea – Selaginellaceae). CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 3 The endemic species to Colombia used as medicinals are represented mainly by 195 species (95.6%) of angiosperms distributed as follows: Compositae (Asteraceae, 52 species); Araliaceae (12 species); Gesneriaceae (nine species); Lamiaceae and Piperaceae (eight species each); Campanulaceae, Malvaceae, and Rubiaceae (seven species, each); Symplocaceae (six species); Caprifoliaceae and Fabaceae (five species each); Acanthaceae, Rosaceae, and Styracaceae (four species each); and Berberidaceae, Brassicaceae, Clusiaceae, Erythroxylaceae, Gentianaceae, Melastomataceae, and Urticaceae (three species each). The remaining families are represented by only two or a single species. Among the 204 endemic species of Colombia used as medicinal, 123 species (60.3%) are widely distributed TRADITIONAL KNOWLEDGE acceptable 13% Aa regular 2% scarce 85% (in three or more departments), 64 species (31.4%) have restricted distributions (in one or two departments), and 17 species (8.3%) have an uncertain distribution in the country. From the operationalisation and factors score for each theoretical variable, most species were considered as having “scarce” evidence of traditional or scientific knowledge (Figure 1A, B, respectively) Only 26 (13%) endemic species used as medicinals in Colombia had their traditional knowledge defined as ‘acceptable’ (Achyrocline bogotensis, Berberis goudotii, Berberis rigidifolia, Brachyotum strigosum, Cecropia mutisiana, Cecropia telealba, Chromolaena barranquillensis, Copaifera canime, Draba litamo, Espeletia grandiflora, Espeletia killipii, Espeletiopsis muiska, Justicia phytolaccoides, Ocotea caparrapi, Pachira speciosa, Passiflora antioquiensis, Pentacalia corymbosa, Peperomia garcia-barrigana, Peperomia putumayoensis, Psychotria boqueronensis, Rubus choachiensis, Senecio niveoaureus, Symplocos theiformis, Valeriana arborea, Vasconcellea goudotiana, and Verbena valerianoides) (Figure 1A). Only three species had the scientific knowledge factor qualified as ‘acceptable’ (Achyrocline bogotensis, Espeletia killipii, and Pentacalia corymbosa) (Figure 1B). The same three species were defined as presenting ‘acceptable’ evidence on both factors (scientific and traditional knowledge) (Table 5, Figure 2). An important result is that no endemic species used as a medicinal in Colombia, presents technological knowledge assessed as ‘acceptable’. 8 7 6 SCIENTIFIC KNOWLEDGE Bb acceptable 1% 5 4 regular 9% 3 2 1 0 scarce 90% Achyrocline bogotensis (Kunth) DC. Espeletia killipii Cuatrec. Scientific Knowledge FIGURE 1. A Proportion of species endemic to Colombia used as medicinals, from which the traditional knowledge was defined as ‘acceptable’, ‘regular’ or ‘scarce’ according to ethnobotanical evidence. B Proportion of species endemic to Colombia used as medicinals, from which the scientific knowledge was defined as ‘acceptable’, ‘regular’ or ‘scarce’ according to scientific evidence. Pentacalia corymbosa (Benth.) Cuatrec. Traditional Knowledge FIGURE 2. Species endemic to Colombia used as medicinals, presenting ‘acceptable’ evidence of traditional and scientific knowledge. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 55 CHAPTER 3 TABLE 5. Medicinal species endemic to Colombia with ‘acceptable’ traditional and scientific knowledge (*) and species with ‘acceptable’ traditional and ‘regular’ scientific knowledge (**). (***) m a.s.l. = metres above sea level; (****) Habitat Classification based on Cuatrecasas (1958). Altitude (***) (m a.s.l.) and habitat (****) Geographic distribution (Colombian Departments) Achyrocline bogotensis (Kunth) DC. (Compositae-Asteraceae) 1,600 – 4,000 Sub-Andean Forest, Andean Forest, Subpáramo, Páramo, and Superpáramo Boyacá, Cundinamarca, Magdalena, Nariño, Norte de Santander, and Santander Espeletia killipii Cuatrec. (Compositae-Asteraceae) 2,120 – 4,010 Páramo, Subpáramo, and Andean Forest (occasional) Cundinamarca and Caldas (?) Pentacalia corymbosa (Benth.) Cuatrec. (Compositae-Asteraceae) 1,800 – 3,960 Andean Forest, Sub-Andean Forest, Subpáramo, and Páramo Antioquia, Boyacá, Cauca, Cundinamarca, Meta, Norte de Santander, and Santander Berberis goudotii Triana & Planch. (Berberidaceae) 1,900 – 3,960 Sub-Andean Forest, Andean Forest, Subpáramo, Páramo, and Superpáramo Antioquia, Boyacá, Cundinamarca, and Meta Berberis rigidifolia Kunth (Berberidaceae) 1,750 – 4,200 Sub-Andean Forest, Andean Forest, Subpáramo, Páramo, and Superpáramo Boyacá, Cundinamarca, Norte de Santander, and Santander Cecropia mutisiana Mildbr. (Urticaceae) 500 – 1,800 Lower Neotropical Forest and SubAndean Forest Cauca, Caldas, Cundinamarca, Huila, Quindío, Risaralda, and Tolima Cecropia telealba Cuatrec. (Urticaceae) 1,650 – 2,600 Sub-Andean Forest, Andean Forest Caldas, Cauca, Risaralda, Valle del Cauca, and Quindío 8 Chromolaena barranquillensis1 (Hieron.) R.M. King & H. Rob. (Compositae-Asteraceae) 0 – 600 Lower Neotropical Forest Atlántico, Bolívar, Chocó, Córdoba, La Guajira, Magdalena and San Andrés Isla 9 Espeletia grandiflora Humb. & Bonpl. (Compositae-Asteraceae) 2,120 – 4,300 Páramo and sometimes in the upper part of the Andean Forest Boyacá, Cundinamarca, Chocó (?), Meta and Santander (**) Espeletiopsis muiska (Cuatrec.) 10 Cuatrec. (Compositae-Asteraceae) 2,170 – 3,800 Subpáramo, Páramo and sometimes in the upper part of the Andean Forest Boyacá, Cundinamarca, Norte de Santander and Santander Ocotea caparrapi1 (Sand.-Groot ex 11 Nates) Dugand (Lauraceae) 1,000 – 2,380 Sub-Andean Forest Antioquia, Cundinamarca and Santander Passiflora antioquiensis H. Karst. 12 (Passifloraceae) 1,800 – 3,000 Sub-Andean Forest, Andean Forest Antioquia, Boyacá, Caldas, Cauca, Cundinamarca, Putumayo, Quindío, Risaralda, Tolima and Valle del Cauca 2,220 – 4,600 Páramo and sometimes in Andean Forest Antioquia, Arauca, Boyacá, Caldas, Cauca, Cundinamarca, Huila, Meta, Nariño, Norte de Santander, Quindío, Santander, Tolima and Valle del Cauca No. Scientific name/Family 1 2 (*) (*) (*) 3 4 5 6 7 (**) (**) (**) (**) (**) (**) (**) 13 56 Senecio niveoaureus Cuatrec. (Compositae-Asteraceae) (** CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 3 0 1 2 3 4 5 6 7 Berberis goudotii Berberis rigidifolia Cecropia telealba Espeletiopsis muiska Cecropia mutisiana Espeletia grandiflora Ocotea caparrapi Chromolaena barranquillensis Passiflora antioquiensis Senecio niveoaureus Scientific Knowledge Traditional Knowledge FIGURE 3. Colombian endemic species used as medicinals, presenting ‘acceptable’ of traditional knowledge evidence and ‘regular’ scientific knowledge evidence. In terms of ‘acceptable’ traditional and ‘regular’ scientific knowledge, 10 species were recorded: (4) Berberis goudotii, (5) Berberis rigidifolia, (6) Cecropia mutisiana, (7) Cecropia telealba, (8) Chromolaena barranquillensis, (9) Espeletia grandiflora, (10) Espeletiopsis muiska, (11) Ocotea caparrapi 1, (12) Passiflora antioquiensis, and (13) Senecio niveoaureus (Table 5, Figure 3). Whilst only five species presented regular traditional knowledge (Clusia ellipticifolia, Clusia inesiana, Hesperomeles goudotiana, Salvia bogotensis, and Viburnum cornifolium), 18 species presented regular scientific knowledge (Baccharis bogotensis, Baccharis boyacensis, Berberis goudotii, Berberis rigidifolia, Calea peruviana, Cecropia goudotiana, Cecropia mutisiana, Cecropia telealba, Chromolaena barranquillensis1, Chromolaena perglabra, Clusia ellipticifolia, Espeleptiosis muiska, Espeletia grandiflora, Ocotea caparrapi1, Passiflora antioquiensis, Pentacalia ledifolia, Salvia bogotensis, and Senecio niveoaureus), and only three species presented regular technological knowledge (Achyrocline bogotensis, Passiflora antioquiensis, and Vasconcellea goudotiana). A matrix representing the relation between 28 prevalent pathologies in Colombia and those 13 species used as medicinals with acceptable traditional knowledge and 1 acceptable or regular scientific knowledge is presented (Table 6). The results show, for example, that chronic lower respiratory diseases and acute respiratory infections, ranked, respectively, in third and the fifth place on the list of causes of mortality in Colombia, could be treated by at least by seven different species (1) Achyrocline bogotensis, (4) Berberis goudotii, (7) Cecropia telealba, (8) Chromolaena barranquillensis, (9) Espeletia grandiflora, (10) Espeletiopsis muiska, and (12) Passiflora antioquiensis. Also, species such as Pentacalia corymbosa could treat various known diseases such as dental caries, diarrheal diseases, hypertensive diseases, leishmaniasis, Plasmodium vivax malaria, and periodontal disease (Table 6). DISCUSSION It is important to note that no other country is known for such a high number of native species with potential medicinal uses from the literature available on medicinal plants. The species represent just one of the hidden natural treasures of Colombia that should be broadly known, conserved, managed, and sustainably used by present and future generations. Several native plant species present in Colombia are recognised as having some type of medicinal activity. However, there Ocotea caparrapi and Chromolaena barranquillensis have been synonymised; however, for this chapter, these taxa do not follow the taxonomic criteria which proposed the synonymy of Ocotea caparrapi as Mespilodaphne cymbarum (Ocotea cymbarum) and Chromolaena barranquillensis as Chromolaena odorata. A different point of view has been prepared by the authors as a publication on the taxonomy, nomenclature, geographical distribution, and ecological status of these two species, which are recognized as exclusive to Colombia. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 57 CHAPTER 3 TABLE 6. Matrix of prevalent pathologies in Colombia and endemic medicinal species of Colombia with acceptable traditional knowledge and acceptable or regular scientific knowledge. Codes (present = 1 and absent = 0) represents mentions of any therapeutic use of each species for a prevalent disease in Colombia and shows the total of species that have uses mentioned for a certain pathology and the number of mentions of use for each species for these pathologies. Prevalent pathologies/species 1 2 3 4 5 6 7 8 9 10 11 12 13 Total Ischemic heart disease 0 0 0 1 1 0 0 0 0 0 0 0 0 2 Cerebrovascular diseases 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Chronic lower respiratory diseases 1 0 0 1 0 0 1 1 1 1 0 1 0 7 Mellitus diabetes 0 0 0 1 0 0 0 0 0 0 1 0 0 2 Acute respiratory infections 1 0 0 1 0 0 1 1 1 1 0 1 0 7 Hypertensive diseases 0 1 1 0 0 0 0 0 0 0 0 0 0 2 Malignant stomach tumour 1 0 0 0 0 1 0 1 0 1 0 0 0 4 Malignant tumour of the trachea, bronchi, and lung 1 0 0 0 0 1 0 1 0 1 0 0 0 4 Kidney disease pain 1 0 0 0 0 0 0 0 0 0 0 0 0 1 Heart failure 0 0 0 1 1 0 0 0 0 0 0 0 0 2 Periodontal disease 0 0 1 1 0 0 0 0 0 0 0 0 0 1 Dental caries 0 0 1 1 0 0 0 0 0 0 0 0 0 1 Iron deficiency anaemia 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Asthma 0 0 0 0 1 0 1 1 1 0 0 0 0 4 Trichuriasis by Trichuris trichiura 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Hypertensive heart disease 0 0 1 0 0 0 0 0 0 0 0 0 0 1 Protein-calorie malnutrition 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Hookworm 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Abortion 0 0 0 0 0 0 1 0 0 0 0 0 0 1 Unipolar major depression 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Diarrheal diseases 0 0 1 0 0 0 0 0 0 1 0 0 0 2 Malaria by Plasmodium vivax 0 1 1 0 0 0 0 0 0 0 0 0 0 2 Leishmaniasis 0 1 1 0 0 0 0 0 0 0 0 0 0 2 Classic Dengue 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Malaria by Plasmodium falciparum 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Tuberculosis 0 0 0 0 0 0 0 0 0 0 0 0 0 0 AIDS HIV 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Dengue haemorrhagic fever 0 0 0 0 0 0 0 0 0 0 0 0 0 0 TOTAL 5 3 7 7 3 2 4 5 3 5 1 2 0 ------ MORTALITY MORBIDITY EPIDEMIOLOGICAL SURVEILLANCE 58 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 3 are no detailed publications for each species on the current state of traditional, scientific, and technological knowledge. Also, there is no published compilation to facilitate the identification and characterisation of species and with the potential to promote their use in primary health care. Likewise, there is also a lack of research, development, and innovation projects (I+D+i) (especially for endemic species) to allow the chemical-pharmaceutical synthesis or the sustainable use of raw materials to elaborate pharmaceutical preparations and phytotherapeutic products. Achyrocline bogotensis, Espeletia killipii, and Pentacalia corymbosa are endemic plant species of Colombia used as medicinals that present the most extensive information on traditional and scientific knowledge, and used to treat and prevent some of the most prevalent pathologies within the country and in other tropical countries (see Table 6). However, it is still necessary to carry out studies focusing on these species regarding safety policies (toxicity) and indepth investigations on efficacy policies for medicinal use, including phytochemical investigations, investigations of in vivo and in vitro biological activities and clinical studies. In the same way, it is still necessary to carry out new studies to expand the knowledge on agrotechnologies for organic and sustainable cultivation and on technologies for the elaboration of pharmaceutical preparations and phytotherapeutic products made from those three species. Mentions of the traditional use of Achyrocline bogotensis are very varied, with most of the available information relating to conservation status and sustainable use. Regarding the evidence of traditional and scientific knowledge, the information on anti-inflammatory activity demonstrated from studies is remarkable. Several scientific investigations show that this species is also antioxidant, anticancer and has biocidal activities (García 1992; Pombo, 2003; Bernal et al., 2011). This species also has trypanocidal activity against Trypanosoma epimastigotes and proven properties against Staphylococcus aureus, Staphylococcus epidermidis, Staphylococcus pyogenes, Mycobacterium avium, Mycobacterium abscessus, Mycobacterium fortuitum, Mycobacterium chelonae, Microsporum gypseum, and Trichophyton mentagrophytes (Bernal et al., 2011; Baldisserra et al., 2014). Evaluation of the extracts and fractions of leaves, stems and flowers demonstrated in vitro antiviral activity against a rotavirus (RRV strain) and an astrovirus (Yuc8 strain) (Téllez et al., 2015). In the Colombian phytotherapeutic market, a product of Achyrocline bogotensis registered by the National Institute of Food and Drug Surveillance (Invima) (No. PFT2018-0000487-R1) presents sale conditions with medical formula as adjunct support in the treatment of mild inflammation of the lower urinary tract. Also, Téllez et al. (2015) developed a phytotherapeutic product to treat acute diarrheal disease on an industrial pilot scale obtained from the aerial parts of this species. The bioactive fraction and the formulation of the phytotherapeutic product were patented in 2017 (Patent No. 16-017515 Superintendency of Industry and Commerce Colombia). It is important to record the scientific findings on Achyrocline bogotensis because no other specific antiviral therapy reduces acute diarrheal illness. The World Health Organization (WHO, 2021) estimates that each year there are 1,700 million episodes of diarrhoea in children under five years in developing countries and 760,000 deaths due to acute diarrhoea related to 50–70% of cases with dehydration. For example, rotavirus alone produces about 500,000 deaths of children under five years old, and the protocols described by the WHO to treat this pathology are solely aimed at the rehydration of patients. For Espeletia killipii, the small farmers of Colombia agree on their traditional medicinal uses, mainly for asthma and lung diseases, which seem to be endorsed by the scientific research that has been carried out (Torrenegra et al., 1994; Bernal et al., 2011). According to Torrenegra & Téllez (1996), the kaurenoic derivatives of this plant are toxic to the potato (Solanum tuberosum) white worm (Premnotrypes vorax). This species also presents marked antibacterial activity against Bacillus subutilis, Salmonella tiphimurium, Staphylococcus aureus, and Streptococcus pneuminiae ( Bernal, in prep.). In general, the traditional uses of Pentacalia corymbosa are not aligned with the results of the scientific investigations. In Colombian popular medicine, this plant is used as a vulnerary alexipharmaceutical and antisyphilitic. Several chemical compounds identified from this species are antioxidants (Uribe-Holguín, 2010), and they act against dermatophyte (Trichophyton mentagrophytes), have cytotoxic and antitumor action and are antifungal against Botrytis cinerea, Fusarium oxysporum, and Trichoderma viride (Torrenegra et al., 2000). From the ten endemic medicinal species in Colombia that have acceptable traditional knowledge and regular scientific knowledge (see Figure 3), the species from the genera Berberis, Cecropia, Chromolaena, and Ocotea stand out. Berberis goudotii and Berberis rigidifolia are used in popular medicine as a febrifuge, cutaneous leishmaniasis treatment, diaphoretic, haemostatic, treatment for pyorrhoea, purgative, remedy for throat infections, and tonic (García, 1974; Bernal et al., 2011; Sequeda-Castañeda et al., 2019). These species can play a significant role in the treatment or prevention of diseases that are important causes of morbidity in Colombia, since it has been proven that they contain chemical compounds that can act in a curative way in certain forms of cutaneous leishmaniasis which is a fairly frequent epidemiological surveillance disease in the country. More recently, a preliminary phytochemical analysis of Berberis goudotii determined that this species presents antimicrobial activity against bacteria associated with caries and periodontal disease (Sequeda-Castañeda et al., 2019). Cecropia mutisiana and Cecropia telealba are myrmecophilic species that are important for feeding birds and mammals in the Southern Neotropical rainforest. The Andean and sub-Andean forests of Colombia have many uses in traditional medicine (Bernal 2012). However, there are only a few scientific investigations to verify their biological activities (Sequeda-Castañeda et al., 2015). The main uses in traditional medicine are as antiasthmatics, antibilious medicines, arthritis treatments, cardiotonics, chorea medicines, diuretics, hypoglycaemics, agents to improve nervous mobility of the body, and rheumatism remedies. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 59 CHAPTER 3 Cecropia mutisiana extracts have cardiotonic activity (Pinzón & Eraso, 1975). Something particular demonstrated by Zambrano-Ospina (2000) is that although the extract of the leaves is not effective against chronic seizures induced by pentylenetetrazol (PTZ), nor does it increase the latency time of seizures when used at normal doses and under test conditions; doses of 1,000 mg/kg protect 40% of the test animals from death. From this group of 10 endemic medicinal species, Chromolaena barranquillensis has traditional uses mentioned as an antioxidant and cytotoxic plant, and as a treatment for boils, inflammations, and skin ulcers. There have been phytochemical studies on this species (MuñozAcevedo et al., 2011). The antioxidant activity of the flowers and leaves of this species was also verified (Muñoz-Acevedo et al., 2011). Ocotea caparrapi is a tree up to 30 m high that grows between 1,000 and 2,380 m a.s.l. in the subAndean forests of Colombia. Its oil, called “caparrapi oil”, has a very broad historical record of traditional uses: as an alexipharmic, alexiteric, anticancer agent, antidote against wasp and scorpion stings, antivenom, antirheumatic, antiviral, mosquito repellent, cosmetic, anti-inflammatory, vulnerary, and veterinary medicine, as well as as a treatment for gonorrhoea, hair loss, bruises, diarrhoea, epigastric pain, eczema, skin diseases, erysipelas, intermittent fevers, malarial fevers, herpes, hydrophobia, ant (diseases of the mule and horse hoof), leprosy, lepromas, epithelial lesions, leucorrhoea, snake and rabid dog bites, insect bites, stingray stings, scabies, ringworm, tumours, ulcers, atonic ulcers, warts, and bronchial, skin and lung infections (extensive literature reviewed by Bernal, in prep.). This species has many phytochemical studies carried out by various authors between 1950 and 2010, and there are some basic studies on skin cancers (epitheliomas) (Bernal, in prep.). This research corroborated the lack of knowledge on the conservation and sustainable use of medicinal species in Colombia, especially endemic species. All the endemic species to Colombia used as medicinals lack clinical studies or, at the very least, preclinical studies (including in vitro and in vivo biological and pharmacological activities, pharmacodynamics, pharmacokinetics, and kinetics). Most of these species have few scientific and technological studies. Only Achryrocline bogotensis, Espeletiopsis muiska, and Vasconcellea goudotiana have advances in this type of knowledge. The species Achyrocline bogotensis, Ocotea caparrapi and Salvia bogotensis present preclinical studies of biological and pharmacological activity in vitro and in vivo. The species Berberis rigidifolia and Clusia ellipticifolia have preclinical studies of biological and pharmacological activity in vivo; whilst the species Cecropia goudotiana, Cecropia telealba, Chromolaena perglabra, Espeletia grandiflora, Espeletia killipii, Espeletiopsis muiska, Passiflora antioquiensis, Pentacalia corymbosa, Pentacalia ledifolia, Senecio niveoaureus, and Vasconcellea goudotiana have preclinical biological and pharmacological studies of activity. In general, all endemic medicinal species of Colombia show insufficient knowledge about agrotechnologies for their organic and sustainable cultivation and studies 60 on agroforestry in polyculture. Likewise, there are no technological studies on raw materials and elaboration of medicinal products from these species. The native medicinal species of Colombia, especially the endemic species, constitute a true natural heritage that requires concrete and innovative actions to develop knowledge, conservation, and sustainable use. Several actions are currently taking place at the Alexander von Humboldt Biological Resources Research Institute, whose next publication will document the updated inventory and monographs on the current state of traditional, scientific, and technological knowledge on the main endemic medicinal plants in Colombia (Bernal, in prep.). Also, there are great efforts and initiatives from the Quindío Botanical Garden to establish on its premises the “National Collection of Medicinal Plants”, supported by the National Botanical Gardens Network of Colombia, The Quindío Botanical Garden also and which seeks to establish a living reference collection for the country’s native medicinal plants with an emphasis on endemic species (Bernal & Gómez, 2021). In general, the native medicinal species of Colombia (especially the South American and Andean species present in the country and several of its endemic species) have great potential to prevent or treat the prevalent pathologies in Colombia and surely in other mainly tropical countries. However, scientific, and technological studies must be continuous to allow the conservation and sustainable use of medicinal species. We strongly recommend the enrichment of programs and projects on research, development and innovation (R + D + I) which aim: i) to conserve and take advantage of species sustainably and profitably, either through the route of chemical-pharmaceutical synthesis for the generation of international exchange for patenting, or through the manufacture of medicinal products that serve to improve social security systems in health; ii) to reduce imports of phytotherapeutic products, and iii) to generate international exchange for exports of medicinal products made from native medicinal species to Colombia, in particular from those species that are to this country, which by their very nature present differential competitive advantages in the market of medicinal products at national and international levels. Acknowledgements The authors would like to thank Hernando García Barriga (q.e.p.d), Professor-Investigator at the Institute of Natural Sciences of the National University of Colombia; Elizabeth Hodson de Jaramillo, Former Director of the Department of Biology of the Pontificia Universidad Javeriana, and Hernando García Martínez, Director of the “Alexander von Humboldt” Institute for Research on Biological Resources, who have studied, and have always supported the scientific research, on the flora of Colombia and in particular the native medicinal plants of the country, and have been able to collect Colombian plants and put into practice a phrase by Enrique Pérez Arbeláez, who in 1956 pointed out that “a plant or animal cannot be considered as a natural resource, until when, as result of the investigation, be put at the service of humans”. 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(Berberidaceae) with anticariogenic and antiperiodontal activities. Scientia Pharmaceutica, 87(1): 2. doi: 10.3390/scipharm87010002 Sequeda-Castañeda LG, Célis C, Ortíz-Ardila AE (2015) Phytochemical and therapeutic use of Cecropia mutisiana Mildbr. (Urticaceae) an endemic plantfrom Colombia. Harmacolog Onl., 3:62–64. Téllez MA, Téllez AN, Vélez F, Ulloa JC (2015) In vitro antiviral activity against rotavirus and astrovirus infection exerted by substances obtained from Achyrocline bogotensis (Kunth) DC. (Compositae). BMC Complement Altern Med, 15(1):1–11 doi: 10.1186/s12906-015-0949-0 Torrenegra RD, Pedrozo J, Téllez AN, Cabeza G, Granados A, Méndez D (2000) Chemistry and antifungal activity of Pentacalia corymbosa (Asteraceae - Senecioneae). Rev Latinoam de Química, 28: 31–34. Torrenegra RD, Téllez AN (1996). Phytochemistry of Espeletia killipii Cuatr. and gibberellic activity of some of the isolated compounds. Rev Latinoamer Quím, 24, 2–6. Torrenegra RD, Téllez AAN, García G (1994) Quimica de especies del género Espeletia: Espeletia killipii - Espeletia tunjana. Rev Colomb de Química, 23(2): 29–35 Uribe-Holguín C (2010) Evaluación de la actividad antioxidante de las hojas de Pentacalia corymbosa y Pentacalia nitida (Asterales: Asteraceae). Pontificia Universidad Javeriana. Vásquez C, Bernal H (2011) Plantas medicinales de Colombia: origen, uso tradicional, eficacia, seguridad y aplicabilidad desde el sistema de salud. In: Pautas para el conocimiento, conservación y uso sostenible de las plantas medicinales en Colombia. 45-130pp. Bogotá, Alianza Ediprint Ltda., Instituto de Investigaciones de Recursos Biológicos Alexander von Humboldt. WHO (World Health Organization) (2021) Temas de salud. Mortalidad y morbilidad. Available at: https://www.who.int/home/search. Accessed in June 2021. Zambrano-Ospina AM (2000) Evaluación de la actividad ansiolítica y anticonvulsivante de las hojas de Cecropia mutisiana. Bogotá, Universidad Nacional de Colombia. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 61 CHAPTER 4 Boronía (confit aubergine with mashed plantain, crispy chickpeas and edible flowers and leaves); recipe from the Celele restaurant, in Cartagena, Colombia). . 62 Mauricio Diazgranados CATALOGUE OF USEFUL PLANTS OF COLOMBIA Chapter 4 The hidden food basket of Latin America: an overview of Colombian edible plant diversity and its distribution Benedetta Gori 1*, Tiziana Ulian1, Henry Yesid Bernal2 & Mauricio Diazgranados1* 1 2 Royal Botanic Gardens, Kew Ex-Profesor Asociado del Departamento de Biología, Facultad de Ciencias, Universidad Javeriana, Herbario de la Pontificia Universidad Javeriana (HPUJ), Bogotá, D.C., Colombia. henryesid@gmail.com *Corresponding authors: b.gori@kew.org, m.diazgranados@kew.org Keywords: Edible plants, Colombia, biogeography, conservation. ABSTRACT Colombia is the second most biodiverse country globally, hosting more than 30,000 plant species and an extraordinary diversity of natural ecosystems. More than 7,000 of these species have reported human uses, and 3,806 of them are food plants. However, despite the high availability of natural resources, Colombia is characterised by widespread poverty and food insecurity. With growing attention being paid to the economic and nutritional benefits of neglected and underutilised edible plants, the present chapter explores the unrivalled diversity of edible plants in Colombia, providing a basis for future research on priority edible species for conservation and sustainable use in the country. It is focused on four fundamental questions: What does Colombian edible plant diversity look like? What is the distribution of Colombian edible species diversity across different bioregions? What is the conservation state of Colombian edible plant diversity in-situ and ex-situ? And, finally, what are the underutilised edible species that have the potential to improve the food security of the local communities and to be commercialised in national and international markets? Results show that the incredible biocultural richness of Colombian edible plants currently receives inadequate conservation actions, both in-situ and ex-situ. There are significant gaps in knowledge regarding the conservation status of some of the most important edible plant genera. The biogeographic analysis highlights the “Tropical Andes” as a diversity hotspot for edible plants, emphasising the urgent need for the preservation of this habitat. Further research is critical to understanding the complex interactions between people, food and the natural environment. At the same time, conservation efforts need to consider the intrinsic value of edible plant diversity and to identify its full potential for socio-economic development at the local and national levels. RESUMEN Colombia es el segundo país con mayor biodiversidad del mundo. Alberga más de 30.000 especies de plantas, así como una extraordinaria diversidad de ecosistemas naturales. Más de 7.400 especies presentan registros de usos humanos, y 3.806 de ellas son plantas alimenticias. Sin embargo, a pesar de la alta disponibilidad de recursos naturales, Colombia se caracteriza por una pobreza e inseguridad alimentaria generalizadas. Siguiendo la creciente atención a los beneficios económicos y nutricionales de las plantas comestibles desatendidas y subutilizadas, el presente capítulo explora la diversidad inigualable de plantas comestibles en Colombia, con el objetivo de proporcionar una base para futuras investigaciones sobre especies comestibles prioritarias para la conservación y el uso sostenible en el país. Este estudo se centra en cuatro preguntas fundamentales: ¿cómo es la diversidad de plantas comestibles colombianas? ¿cuál es la distribución de la diversidad de especies comestibles colombianas en diferentes bioregiones? ¿cuál es el estado de conservación insitu y ex-situ de la diversidad de plantas comestibles colombianas? y ¿cuáles son las principales especies alimenticias subutilizadas de Colombia, que pueden ser usadas para mejorar la seguridad alimentaria de las comunidades locales y que tienen potencialidades económicas en los mercados nacionales e internacionales? Los resultados muestran que la increíble riqueza biocultural de las plantas comestibles colombianas recibe en la actualidad acciones de conservación inadecuadas, tanto in-situ como ex-situ, y existen importantes lacunas en el conocimiento sobre el estado de conservación de algunos de los géneros de plantas comestibles más importantes. El análisis biogeográfico destaca las bioregiones “Andes” y “Páramo” como puntos críticos de diversidad para las plantas comestibles, enfatizando la urgente necesidad de su preservación. Es fundamental realizar más investigaciones para comprender las complejas interacciones entre los pueblos, los alimentos y el medio ambiente natural, mientras que los esfuerzos de conservación deben considerar el valor intrínseco de la diversidad de plantas comestibles e identificar todo su potencial para el desarrollo socioeconómico a nivel local y nacional. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 63 CHAPTER 4 INTRODUCTION Food represents one of the deepest forms of interaction between humans and the environment. It lies at the foundation of human experience, shaping our relationship with non-human living beings and embedding forms of intangible cultural legacy. According to the World Checklist of Useful Plant Species (Diazgranados et al., 2020), more than 7,000 plants are currently known to be edible. Many of them are part of the traditional gastronomic heritage of human populations and have the potential to support food security and develop sustainable agriculture around the world (Ulian et al., 2020). However, only ten species contribute to almost all human caloric intake nowadays (FAO, 2015). This incongruous trend was triggered by the green revolution (Wolff, 2004), which started to replace traditional landraces and wild species with a restricted assortment of modern commercial hybrids by favouring yield production (Paludosi et al., 2011). The downsides of such direction were manifold: on the one hand, people started to diminish their interest in local edible plants while progressively decreasing their attention on the wellbeing of the ecosystems hosting them, which led to their degradation (Kalamandeen et al., 2018). On the other hand, increasing pressure on a narrow portion of natural resources, together with unsustainable cultivation and harvesting practices, resulted in a rapid depletion of natural plant populations (Pilgrim et al., 2008; Nogueira et al., 2011). In response to this trend, international policy frameworks combining biodiversity protection and sustainable development have gained attention in recent decades (FAO, 2019). Efforts have increased focus on the revitalisation and greater use of what Borelli et al. (2020) defined as “orphan crops” and wild edible plants (Borelli et al., 2020). Over the past few decades – more than 50 years after the green revolution – “Neglected and Underutilised Species” (NUS) have been shown to hold crucial importance for building sustainable livelihoods and mitigating environmental deterioration (Hunter et al., 2019; Borelli et al., 2020; Ulian et al., 2020). They are also known to hold critical biocultural values, as they are linked to local agricultural and culinary traditions and practices, symbolising the organic relationship between nature and culture. Growing evidence has demonstrated that the value that peoples give to local plant resources can play a crucial role in their engagement in conservation and sustainable management (Brehm et al., 2010; N’Danikou et al., 2011). “Conservation-through-use” approaches – aimed at encouraging nature conservation through the sustainable use of its resources – are increasingly being applied in conservation programs worldwide (Kor et al., 2021). Colombia is one of the world’s “megadiverse” countries (Renjifo et al., 2020). More specifically, it is the second most biodiverse country on the planet, hosting 10% of the world’s total biodiversity (Clerici et al., 2019) and bringing together an unequalled number of distinct natural ecosystems and human cultures. However, despite its unrivalled biocultural richness, Colombia is nowadays characterised by 64 widespread poverty and food insecurity (Hurtado-Bermudez et al., 2020). While local communities had historically taken extensive advantage of local plant diversity, such diversity has been increasingly neglected over the past decades and replaced by high yielding commercial varieties (Grau & Aide, 2008). In addition, new land uses are now causing landscape conversion, driven by export-oriented industrial and agricultural policies and market conditions (Grau & Aide 2008; Boron et al., 2016). Considering these circumstances, investigating, and understanding the complex interactions between people, food, and their environment (Pieroni et al., 2016) acquires crucial importance for formulating targeted and effective conservation and development activities at the local level. Within Colombian edible plants, numerous NUS hold the potential to address environmental degradation while creating sustainable livelihoods and boosting a new green growth, although being unknown outside of the country (Padulosi et al. 2011; Ulian et al. 2020). By combining biodiversity informatics and taxonomic approaches, this chapter reviews Colombian edible plant diversity, estimating its biogeographical distribution, highlighting biodiversity hotspots, and providing the basis for future efforts towards the revitalisation and conservation of edible plants in Colombia. MATERIALS AND METHODS Methods used for this study followed the approach employed by Diazgranados et al. (2020) for the World Checklist of Useful Plant Species. The Checklist added information from three large international datasets to the first State of the World’s Plants report published in 2016 (RBG Kew, 2016). To date, the World Checklist of Useful Plant Species is the most comprehensive piece of work on useful plants at a global scale, containing key taxonomic and ethnobotanical information on 40,292 species. Among all of the plants classified as “useful”, 7,039 are classified under the category of “Human Food”, according to the Economic Botany Data Collection Standard categorisation (Cook, 1995). The same taxonomic backbone of the World Checklist of Useful Plant Species was adopted to reconcile taxonomically species from a combination of ten new Colombian and international datasets and publications, in addition to data from the Checklist (cf. Annotated Checklist of Useful Plants of Colombia). The new datasets were cleaned using RStudio4.1.0 v. (R Core Team, 2021). Taxon names were reconciled to Plants of the World Online (POWO, 2021) and, occasionally, to Tropicos (Missouri Botanical Gardens, 2021) using the “Plyr” and “Dplyr” packages (Wickham, 2011; Wickham & Francois, 2021). Higher taxonomy information was obtained from the Global Biodiversity Information Facility (GBIF, 2021). Data on species edibility was retrieved from the cleaned datasets. Data on the in-situ conservation actions for edible species were obtained from the IUCN Red List of Threatened Species (IUCN, 2021). PlantSearch (BGCI, 2021), a platform developed by Botanic Gardens Conservation International that contains taxon-level data from worldwide gene and seed banks, was used to gather data on ex-situ conservation actions. CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 4 Occurrences of edible species were downloaded from GBIF (2021) through the “RGbif” package (Chamberlain et al., 2021) and cleaned using the “ShinyCCleaner” package (Ondo, in prep.) in RStudio. Filters for removing occurrences recorded within urban centres and institutions, areas (e.g., Botanic gardens and ex-situ repositories) were also applied to remove points in the sea and the centre of Colombia (i.e., centroid points). Moreover, latitude and longitude points with fewer than three decimals were removed, as were occurrences with equal latitude and longitude and occurrences with either latitude or longitude equal to zero. Mapping of edible species across Colombian departments and ecoregions was carried out in R 4.1.0 (R Core Team, 2021). Heatmaps showing species richness per department and grid were developed using the “ggplot2” package (Wickham, 2016). Grid analysis to measure species richness by ecoregions was carried out using the “rgdal” package (Bivand et al., 2021), raster (Hijmans & van Etten, 2012), and sp (Bivand et al., 2013). ArcGIS pro 2.8.1 was also employed to carry out biodiversity quantification. SDMtoolbox (Brown et al. 2017) – a python-based toolbox for spatial analysis – was used to obtain several biodiversity metrics such as species richness (i.e., the sum of species per cell), weighted endemism (i.e., the sum of the proportion of species’ ranges found in a given grid cell – emphasising areas that are rich in species with restricted distributional ranges) and corrected weighted endemism (i.e., weighted endemism divided by the total number of species in a cell – emphasising areas that are rich in species that have restricted ranges, but that are not necessarily speciesrich), employing a geographic resolution of 0.1 degrees. FIGURE 1. Richness per department. FIGURE 2. Richness of observations of Colombian edible plants by department. of Colombian edible plant species RESULTS AND DISCUSSION Overview of Colombian edible plants The current work resulted in the cataloguing of 7,205 useful species found in Colombia, among which 3,805 were categorised as edible (i.e., characterised by a history of consumption by human populations). If compared to the data showcased by the World Checklist of Useful Plants (Diazgranados et al., 2020), this number acquires crucial significance: in fact, the proportions of useful species and edible species at the global level (respectively, 40,292 and 7,039 in number) are remarkably lower than those specific to Colombia. According to the Economic Botany Data Collection Standard (Cook, 1995), at the global level, only 17.4% of useful plants have been recorded as “Human food” (Diazgranados et al., 2020). In Colombia, this percentage rises to 53.6%, making this country a global reservoir for edible plant diversity. The distribution of Colombian edible species across different departments is showcased in Figures 1 and 2. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 65 CHAPTER 4 Based on the clean georeferenced records for Colombian edible plants, the departments of Antioquia and Meta stand out for species richness. However, these are also the departments where most occurrences were recorded, and linear regressions revealed a significant correlation (p<0.001) between species richness and the number of georeferenced records (Figure 2). Therefore, it is possible to argue that other departments may not stand out simply because of a lack of adequate on-site investigation and the scarcity of georeferenced records collected. Colombian edible plants are represented by 219 families and 1,389 genera. The most important families (i.e., those containing the highest number of species), taking into account both native and non-native species, include Fabaceae (119 genera, 351 species), Asteraceae (86 genera, 136 species), Poaceae (72 genera, 140 species), Arecaceae (55 genera, 180 species), and Rubiaceae (52 genera, 137 species). This scenario partly reflects the global trend highlighted by the World Checklist. However, if only native species are considered, families such as Melastomataceae (14 genera, 119 species) and Malvaceae (35 genera, 91 species) obtain greater relevance, followed by Moraceae (18 genera, 74 species), Annonaceae (14 genera, 66 species), and Myrtaceae (11 genera, 61 species). Important genera, including both native and non-native species, comprise Inga (84 spp.), Passiflora (73 spp.), Miconia (63 spp.) Solanum (61 spp.), Pouteria (54 spp.), Protium (33 spp.), Annona (32 spp.), and Bactris (28 spp.). Genera such as Ficus, Diospyros, and Garcinia, known to be among the most relevant edible plants at the global level (Diazgranados et al., 2020), are not significantly rich in edible species in Colombia. On the other hand, genera such as Passiflora, Inga, Bactris, and Pouteria are characterised by a high number of edible species and may represent a new frontier for ethnobotanical and bromatological studies in the country. Colombian edible plants comprehend a great variety of growth forms, from trees to herbs, climbers, and epiphytes. Trees constitute the most dominant habit, with more than 1,500 species, followed by herbs, shrubs, and climbers. Growth habit highly reflects the biogeographic distribution of edible plants across various ecoregions (Figure 3). A B 1000 35 30 800 25 600 20 15 400 10 200 5 C D 1000 15 200 800 10 150 600 100 50 400 50 200 FIGURE 3. Distribution of Colombian edible plants by growing habit: A tree species, B shrub species, C herb species and D climbing species. The four distributions show relatively congruous trends, with high species concentrations across the Andean region. 66 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 4 Some of the most important tree genera include Inga (64 tree spp.), Pouteria (49 spp.), Miconia (39 spp.), Protium (32 spp.), Annona (27 spp.), Ficus (21 spp.), Casearia (21 spp.), and Matisia (20 spp.). On the other hand, important herbs include Solanum (30 herb spp.), Cyperus (16 spp.), Miconia (14 spp.), Oxalis (11 spp.), and Eragrostis (11 spp.). Predominant genera for shrub species include Miconia (54 shrub spp.), Solanum (33 spp.), Bactris (25 spp.), Casearia (20 spp.), Piper (15 spp.), and Senna (15 spp.). Finally, the most important genera for edible climbers comprise Passiflora (64 climbing spp.), Ipomoea (15 spp.), Dioscorea (15 spp.), Paullinia (14 spp.), and Solanum (12 spp.). Examples of important genera Passiflora (Passifloraceae) Several species are characterised by great economic potential, but only eight of them are currently cultivated (P. antioquiensis, P. caerulea, P. edulis, P. ligularis, P. mixta, P. quadrangularis, P. tarminiana, and P. tripartite). Other species, including P. vitifolia, are wildly harvested for local consumption or sale in local markets. There is substantial morphological variation within the genus (e.g., in the colour and shapeof the fruits and flowers). The fruits can be eaten raw, cooked, or even used to make drinks, such as P. antioquiensis. The pulp is very aromatic, and flowers and leaves are also edible sometimes. In the case of P. foetida, the leaves are cooked and used as an ingredient in soups. The pulp of fruits is very variable in terms of taste: it can be sweet (e.g., P. ligularis), juicy and acidflavoured (e.g., P. coccinea), or aromatic and mildly biting (e.g., P. cumbalensis). Finally, it is important to mention P. mollissima, which is traditionally eaten in various departments of Colombia and shows a great versatility of use. In fact, it can be consumed and marketed in the form of juice, sorbet, ice cream, or jam. Solanum (Solanaceae) With 61 known edible species, Solanum is the third most important genus in Colombia for food plants. In addition to some of the most economically valuable plants that the world has seen, such as S. lycopersicum (tomato), Nicotiana tabacum (tobacco), and S. tuberosum (potato), this family includes an outstanding number of edible plants, many of which hold lesser economic importance and are exclusively known, grown, and consumed at the local level. In Colombia, these are especially present in the Andean region and are still largely unexplored from a taxonomic, agronomic, and bromatological perspective. Examples include S. cajanumense, a fast-growing evergreen shrub whose goldenyellow fruits are usually eaten fresh when fully ripe, S. capsicoides, whose poisonous fruits can be eaten when roasted or cooked, and S. pectinatum, whose pale orange fruit, characterised by a sweet-acidic flavour, is delicious when cooked with sugar (Food Plants International, 2021). Only 14 out of 61 species are currently being cultivated. 51 species out of 61 are native to the country, and only one of them is endemic. Inga (Fabaceae) The edible fruit of this genus is very popular throughout South America, where it is commonly gathered from the wild and often cultivated (Lorenzi et al., 2000). Inga edulis is the best known and most consumed species. Both the seeds and the white and jelly pulp surrounding them within the fruit’s seedpod can be eaten. The pulp is characterised by a sweet and highly aromatic taste (Martin et al., 1987), and it is usually eaten raw. The seeds are eaten cooked, usually boiled, or roasted, like in I. ilta. When young, they can also be eaten raw, blanched and salted, and added to salads (Food Plants International, 2021). Inga trees are commonly planted in coffee or cacao plantations to provide shade to the surrounding environment, and fruits are often sold in local markets (Leon, 1966). Out of 84 species found in Colombia, 76 are native to the country, and none of them is endemic. Only six species are currently being cultivated (I. densiflora, I. edulis, I. feuillei, I. ornate, I. spectabilis, and I. vera). Bactris (Arecaceae) Bactris is a genus of multi-purpose spiny palm trees native to Latin America, the most popular of which is B. gasipaes, commonly known as peach palm. In total, 28 species of Bactris are known to be edible in Colombia, but only B. gasipaes is currently cultivated. All 28 species are native to Colombia, and only one species – B. chocoensis – is endemic. The fruit of most species is inedible when raw (Blombery & Rodd, 1992). They are usually boiled in salted water for 30 to 60 minutes and eaten as a vegetable (Food Plants International, 2021). The pulp is characterised by a nutty flavour and a floury texture, and by its remarkably high nutritious qualities resulting from its high protein and carbohydrate contents. Fruits can also be made into flour and baked into bread, cakes, and other processed foods (Food Plants International, 2021; Blombery & Rodd, 1992). Seeds, such as those of B. major and B. gasipes, can be consumed raw (Wickens, 1995) as nuts. The palm heart of some species (e.g., B. riparia and B. corossilla) is also eaten raw, in salads, or cooked. Finally, fruits of B. guineensis can be fermented and used to produce a drink, known in Colombia as Chicha de Corozo (Chízmar, 2009; Rodriguez, 2019). Vasconcellea (Caricaceae) This genus contains 21 species, which were for a long time classified within the genus Carica. They are of great importance as genetic resources (wild relatives) for the improvement of the papaya (Carica papaya) and for their edible fruits. Additionally, they can be of great importance for producing papain enzyme and latex, which, as raw materials, are promising resources for food and non-food agro-industrial processes. Colombia has eight species popularly known as “tall papayas”, of which three are endemic (Vasconcellea goudotiana, Vasconcellea longiflora and Vasconcellea sphaerocarpa). Several ethnobotanical CATALOGUE OF USEFUL PLANTS OF COLOMBIA 67 CHAPTER 4 studies carried out in the southern Colombian Andes show that V. cundinamarcensis and V. goudotiana are very often reported for their edible and medicinal uses, for both humans and animals (Fuertes, 2019). In the Andes, the “highland papayas” (Vasconcellea spp.) are mainly consumed fresh, toasted, juiced, as jams, sometimes preserved or cooked in sauces, or as fillings for cakes and pickles. These plants, therefore, possess great potential for use. They are usually grown on small Andean farms for home consumption for their tasty and aromatic fruits. Conservation state and current gaps Out of 3,806 Colombian edible plants, only 1,742 have been assessed by the International Union for Conservation of Nature. Out of these, 1,532 have been categorised as LC (Least Concern), 38 as NT (Near Threatened), 69 as VU (Vulnerable), 44 as EN (Endangered), 16 as CR (Critically Endangered), one as EW (Extinct In the Wild), and 40 as DD (Data Deficient). Additionally, two species have been categorised as EX (Extinct). Among the most threatened species, there are some endemic species, such as Melicoccus antioquensis (Sapindaceae) and Panopsis hernandezii (Proteaceae), as well introduced ones, such as Eugenia cotinifolia (Myrtaceae), Araucaria angustifolia (Araucariaceae), Hyophorbe verschaffeltii (Arecaceae), and Lathyrus odoratus (Fabaceae). The most common threats to these species include agricultural expansion and biological resource use. In fact, the degradation of habitats in which these species could be found in the past is mainly due to activities related to the expansion of the agricultural frontier (e.g., forest clearance for cattle ranching and crops cultivation) (IUCN, 2021). What is more, the usefulness of these plants makes them particularly subject to overexploitation and unsustainable harvesting. A significant portion of Colombian edible species boasts uses of other types, for example, having wood that is used for construction or other plant parts that are used as medicines. For instance, in the case of Araucaria angustifolia, in addition to more than 3,000 tons of fruits and seeds collected every year for human consumption, resource depletion can be attributed to its massive exploitation for timber (IUCN, 2021). The sustainable harvesting and use of natural resources remain key issues for determining their conservation: while overexploitation represents the second major driver of plant extinction, the sustainable use of local natural capital can be vital for the development of future livelihoods for indigenous peoples (Kor et al., 2021). As displayed by the significant portion of edible species whose conservation status has not yet been assessed, significant gaps in the knowledge required to support the protection of these species can be identified. For instance, out of the 73 Colombian edible species belonging to the genus Passiflora, none have been assessed by the IUCN. The same trend can be found for other genera, such as Bactris: only three out of 28 species present in Colombia have been assessed. Therefore, we can conclude that the conservation status of two of the richest genera for edible plants in Colombia is unknown. 68 Ex-situ conservation The comparison between the Colombian edible plant dataset and PlantSearch showed a total match of 3,042 species. However, 547 species (17.9% of the total) are not currently kept in any collection (Total gardens = 0). Therefore, the number of edible species included in the present study and currently maintained within the ex-situ collections of the BGCI global consortium is 2,495. In this case, 82.1% of all edible species found in Colombia are currently preserved in at least one ex-situ collection. Categorical analysis results are summarised, with 379 useful plant taxa conserved in only one BGCI botanic garden’s living collection, 780 taxa conserved within 2–10 collections, 290 taxa within 11–20 collections, 344 taxa within 2–40 collections, and 702 taxa within more than 40 collections each (Table 1). TABLE 1. Ex-situ conservation of Colombian edible plants. No. of ex-situ collections No. of edible species % of the total 0 547 17.9 1 379 12.4 2–10 780 25.6 11–20 290 9.5 21–40 344 11.3 >40 702 23 Results show that important genera for edible species, such as Bactris, Inga, and Pouteria, lack adequate ex-situ conservation. For instance, out of 28 edible Bactris species, one is currently not conserved in any ex-situ repository globally, and only two species are currently conserved in more than 15 repositories. Out of 84 edible species of Inga found in Colombia, 59 are not currently conserved in any ex-situ repository, and only one species – Inga edulis – is conserved in more than 15 repositories. Finally, out of 54 edible Pouteria species, 42 are not conserved in any ex-situ collection, and only two species – P. caimito and the introduced P. sapota – are maintained in more than 15 of them. Promising edible species and diversity hotspots The spatial analysis results carried out on the clean dataset, containing 221,838 georeferenced records for 3,220 Colombian edible species, revealed several hotspots for edible species diversity in the northern and north-western Andean region (Figure 4). The corrected weighted endemism distribution pattern for Colombian edible species emphasises areas that are characterised by a high proportion of species with restricted distributional ranges. Therefore, these areas are not only remarkably high in edible species richness but also stand out as diversity hotspots for rare and endemic species. Colombian Andes CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 4 FIGURE 4. Diversity patterns of Colombian edible plants. A Species richness map, showing the Tropical Andes as the richest area. B Weighted endemism map. C Corrected weighted endemism, with the Andes standing out, together with the tropical rain forest and SubAndean rainforest of the Nariño department, the Savanna between the Amazonas and Caqueta departments, and the Paramo of la Guajira. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 69 CHAPTER 4 form part of the Tropical Andes, extending across the north of Chile, Argentina, Bolivia, Peru, Ecuador, Colombia, and Venezuela for over 1.5 million km2, between the latitudinal range of 11° N to 30° S, and with an elevational range of approximately 500 to 6,000 m a.s.l. (Bax & Francesconi, 2019). According to Myers et al. (2020), the Tropical Andes support approximately 45,000 plant species, with nearly half of them endemic to the Andean ecoregion (Meyers et al., 2020). In particular, a significantly high density of narrowdistribution edible plants has been recorded in the northern part of the Huila department. Huila is considered one of the richest regions of Colombia in terms of plant biodiversity due to its great variety of thermal floors, from Páramo zones to extensive areas of tropical dry forest (Figure 5). At least 120,000 hectares of the department are localised within the Páramo area, with elevations ranging between 2,900 and 5,000 m a.s.l. Notably, the area has been described as one of the richest in the country in terms of the diversity of the genus Passiflora (Ocampo, 2013). Another interesting hotspot was identified within the high Guajira department, in the proximity of the Macuira National Park. The Serrania de Macuira, located in the extreme north of La Guajira’s department, is considered of great environmental importance. In fact, their evergreen cloud forest ecosystem, although localised outside of the Andean mountains, appears incredibly similar to the Andean forests located near the Páramo line (altitudes above 2,700 m a.s.l.). Despite this similarity, they are located at only 550 m a.s.l. This ecosystem functions as a water reserve for the surrounding desert region and supports a unique floral diversity. Furthermore, as showcased by the present study, it represents a national reservoir for unique edible plants. Therefore, the present biogeographic analysis highlights Andean ecosystems as important repositories for singular edible plants, which are highly adapted to specific environmental conditions and characterised by narrow distributions. The number of food species present in the Colombian ecosystems that are of low and regular altitudes is very high compared to that in high Andean forests and Páramo, which are characterised by more extreme weather patterns. Therefore, to guarantee their conservation and use in the long term by local populations, it is critical to register the most important species found within these ecosystems, their culinary uses, and their ecological value. Páramo exhibits an unparalleled flora originating from sophisticated evolutionary processes over the past 3 million years and is characterised by a great proportion of endemic species (Londono et al., 2014). However, due to increasing human-driven alterations of natural ecosystems and the progressive impact of climatic variations, ecosystems such as the Tropical Andes are nowadays recognised as one of the most critically threatened ecoregions in the tropics (Noh et al., 2020). This scenario constitutes a particularly serious hazard to the native endemic edible species that inhabit the region (Figure 6), characterised by extremely 70 specific habitat needs. In light of this, Colombian Andes are expected to lose a significant proportion of their native plant diversity due to environmental degradation (Brooks et al., 2002; Noh et al., 2020). This scenario would represent a significant loss not only for the biological heritage that is a feature of the region but also for its socio-cultural heritage, embedded in local traditional agricultural practices and gastronomy. High-altitude ecosystems and their edible flora must therefore be protected and yet further investigated from an ethnobotanical perspective to deepen the evolutionary and socio-cultural practices that underlie a remarkable gastronomic heritage. Finally, based on the obtained results, it is key to showcase the most promising underutilised and neglected species (native and non-native) that are present and easily adapted to different Colombian ecosystems. Such species present significant competitive advantages compared to other cultivated species, which could greatly improve local communities’ economies and, consequently, the country’s bioeconomy. For this purpose, Colombia’s most important food species for the warm, cold, and temperate thermal floors are summarised in Tables 2 and 3. CONCLUSIONS In conclusion, the showcased results highlight consistent discrepancies between the diversity and richness of Colombian edible plants and their current conservation. Both wild and domesticated edible plants have played a significant role in the subsistence of human communities across the country. Their preservation and valorisation, today, is more crucial than ever. After more than six decades of internal conflict, Colombia is currently going through an important time in the country’s development, which will determine the fate of its natural resources in the coming years. However, agricultural expansion and urban development lead to deforestation and habitat loss, causing unprecedented levels of biodiversity loss nationwide. Local underutilised edible plants hold great potential for developing new livelihoods and consolidating a readjusted bioeconomy based on the sustainable use of local plant diversity. The present chapter represents a first review of the Colombian edible diversity from a taxonomic, biogeographical and conservationist point of view. Taxonomic investigation revealed new plant groups worth deepening from an ethnobotanical and commercial perspective, laying the foundation for future efforts to revitalise underutilised edible species. Additionally, the spatial analysis highlighted the biodiversity hotspots in the country. Further analysis considering simultaneously the extinction risk assessments and the in-situ and ex-situ conservation actions in place for the most important edible plant groups could facilitate pinpointed conservation endeavours. CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 4 FIGURE 5. Corrected weighted endemism of Colombian edible plants with bioregions layer. FIGURE 6. Overview of Colombian bioregions. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 71 CHAPTER 4 TABLE 2. Native and non-native edible species suitable for cold thermal floors in Colombia. (*) Altitudinal range (m a.s.l.); (**) Habitat Classification based on Cuatrecasas (1958). Scientific name Family Altitudinal range (*) and habitat (**) Habit 2,400 – 3,500 Herb Andean Forest, 1.5 – 2.0 m Subpáramo and Páramo Main uses Human food (seeds) and animal forage (the entire plant) Amaranthus caudatus L. Amaranthaceae Amaranto, coime, achita Cavendishia bracteata (Ruiz & Pav. Ex J.St.Hil.) Hoerold Ericaceae 2,400 – 3,200 Uva de anís, uva Shrub Andean Forest and de monte, asnalulo 0.8 – 3.0 m Subpáramo Human food (fruits) Chalybea macrocarpa (Uribe) Morales P. & Penneys Melastomataceae Churumbelo Small tree 5 – 12 m Human food (fruits) Cañahua, kañihua Human food Herb 3,500 – 4,200 (seeds) and 0.2 – 0.8 m Páramo and Superpáramo animal forage (stems) 2,400 – 4,000 Herb Andean Forest, Páramo, 1.0 – 2.5 m Subpáramo and Superpáramo Chenopodium pallidicaule Aellen Amaranthaceae 2,400 – 2,900 Andean Forest Human food (seeds) Chenopodium quinoa Willd. Amaranthaceae Quinua, quinoa Gaylussacia buxifolia Kunth Ericaceae Agracillo, mosquito, uvo Shrub 2,500 – 3,500 0.5 – 1.5 m Andean Forest, Subpáramo and Páramo Human food (fruits) Hesperomeles goudotiana (Decne) Killip Rosaceae Mortiño, cerote, manzano, motemote Small tree or shrub 2–8m 2,500 – 3,500 Andean Forest, Subpáramo and Páramo Human and animal food (fruits) 2,400 – 3,100 Andean Forest and Subpáramo Human food (seeds) Juglans neotropica Diels Juglandaceae Nogal, cedro nogal, Tree cedro negro, cedro 15 – 45 m bogotano, tocte Fabaceae (Leguminosae) Lupino, tarwi, altramuz, chocho 2,400 – 3,800 Herb Andean Forest, 1.8 – 2.0 m Subpáramo and Páramo Human food (seeds) Macleania rupestris (Kunth) A.C. Sm. Ericaceae Uva camarona, uvita, uvilla, chaquilulo, otros 2,400 – 4,100 Shrub Andean Forest, Páramo, 0.6 – 2.5 m Subpáramo and Superpáramo Human food (fruits) Miconia squamulosa Triana Melastomataceae Tuno esmeraldo, tunacòn Small tree or shrub 2–8m Human food (fruits) Lupinus mutabilis Sweet 72 Common name(s) 2,400 – 3,200 Andean Forest and Subpáramo Mirabilis expansa (Ruiz & Pav.) Standley Nyctaginaceae Mauka, chago, 2,400 – 3,500 Herb arricòn, miso, taso, Andean Forest, 0.8 – 1.0 m pega pega Subpáramo and páramo Human food (leaves and tuberous roots) and animal food (leaves, stems, and roots) Myrcianthes leucoxyla (Ortega) McVaugh Myrtaceae Arrayán, guayabo silvestre mirto, champo Human food (fruits) Tree 10 – 15 m 2,400 – 3,300 Andean Forest and Subpáramo CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 4 Scientific name Oxalis tuberosa Molina Family Oxalidaceae Common name(s) Oca, oka, papa oca, ibia, cubio Habit Altitudinal range (*) and habitat (**) Main uses Herb 0.2 – 0.3 m 2,400 – 3,900 Andean Forest, Subpáramo and Páramo Human food (tubers) Passiflora antioquiensis Passifloraceae H. Karst. Curuba, curuba Vine antioqueña, curuba 8 – 10 m quiteña 1,800 – 2,700 Andean and Sub-Andean Forest Human food (fruits) Passiflora mollissima (Kunth) Bailey Passifloraceae Curuba, tacso, tumbo, poroporo Woody climber 1,800 – 3,000 Andean and Sub-Andean Forest Human food (fruits) Physalis peruviana L. Solanaceae Uchuva, guchuva, uvilla, aguaymato, vejigón Herb 0.3 – 1.0 m 2,400 – 3,300 Andean and Sub-Andean Forest Human food (fruits) Plutarchia guascensis (Cuatrec.) A.C. Sm. Ericaceae Azabache, coronilla, corales, Shrub coralitos, uvo, uvito 0.5 – 2.5 m de guasca 2,800 – 4,200 Andean Forest, Páramo, Subpáramo and Superpáramo Human food (fruits) Prestoea acuminata (Willd.) H.E. Moore Arecaceae (Palmae) Palmito de altura 2,400 – 2,650 Andean Forest Human food (buds, palm heart) Rubus nubigenus Kunth Rosaceae Morón de páramo, Climbing mora gigante, mora shrub de oso 1–4m 2,600 – 3,200 Andean Forest and Subpáramo Human food (fruits) Smallanthus sonchifolius (Poepp. & Endl.) H. Robinson Compositae (Asteraceae) Yacón, llacòn, aricona, jícama, jíquima Herb 1.5 – 2.5 m 2,400 – 3,000 Andean Forest Human food (tubers) Solanum vestissimum Dunal Solanaceae Lulo de tierra fría, toronjo, tumo, cuque Shrub 2–5m 2,400 – 3,000 Andean Forest and Subpáramo Human food (fruits) Tropaeolum tuberosum Ruiz & Pav. Tropaeolaceae Mashua, añu, isaño, cubio, majua, otros 2,400 – 3,900 Crawler 0.8 Andean Forest, – 1.0 m Subpáramo and Páramo Human and animal food (tubers) Ugni myricoides (Kunth) Myrtaceae O. Berg Arrayancillo, arrayán blanco, mirto 2,900 – 3,400 Shrub 0.5 – Andean Forest, 1.5 m Subpáramo and Páramo Human food (fruits) Ullucus tuberosus Caldas Basellaceae Ulluco, melloco 2,400 – 3,400 Herb Andean Forest, 0.3 – 1.2 m Subpáramo and Páramo Human food (tubers) Vaccinium meridionale Swartz Ericaceae Agraz, mortiño, uva de monte, congama Shrub 1–3m 2,400 – 3,000 Andean Forest and Subpáramo Human food (fruits) Vasconcellea cundinamarcensis V.M. Badillo Caricaceae Chilacùan, chihualcan, higuillo, tapaculo Small tree or shrub 8 – 10 m 2,400 – 3,300 Andean Forest and Subpáramo Human food (fruits) Vasconcellea goudotiana Triana & Planch. Caricaceae Papayuela, papaya de tierra fría, tapacho, tapaculo Small tree or shrub 1–6m 2,640 – 3,575 Andean Forest, Subpáramo and Páramo Human food (fruits) Vicia andicola Kunth Fabaceae (Leguminosae) Haba forrajera, alverjilla, haka, mullu, shintu 2,400 – 4,200 Herb Andean Forests, 0.4 – 0.5 m Páramo, Subpáramo and Superpáramo Animal food (forage) Tree 13 – 15 m CATALOGUE OF USEFUL PLANTS OF COLOMBIA 73 CHAPTER 4 TABLE 3. Native and non-native food species suitable for the warm and temperate thermal floors of Colombia. (*) Altitudinal range (m a.s.l.); (**) Habitat classification based on Cuatrecasas (1958). Scientific name Common name(s) Habit Altitudinal range (*) and habitat (**) Main uses Herb 0.8 – 1.0 m 200 – 1,850 Human food (seeds and Lower Neotropical leaves) and animal food rainforest and Sub(leaves) Andean Forest Anacardiaceae Marañón, merey, caracolí, cahu, aspavé Tree 20 – 25 m 20 – 1,200 Lower Neotropical Human food (fruit rainforest and Sub- peduncle and seeds) Andean Forest Anacardiaceae Anacardo, marañón, merey, caju Small tree 3 – 10 m 0 – 1,300 Lower Neotropical Human food (seeds) rainforest and SubAndean Forest Artocarpus altilis (Parkinson) Fosberg Moraceae Árbol del pan, fruta del pan, frutipan Tree 15 – 20 m 80 – 1,550 Lower Neotropical Human food (fruits) rainforest and SubAndean Forest Bactris gasipaes Kunth Arecaceae (Palmae) Chontaduro, pejibaye, pijuayo, cachipay Tree 7 – 20 m 0 – 1,500 Lower Neotropical Human food (fruits) rainforest and SubAndean Forest Bixa orellana L. Bixaceae Achiote, onoto, bija, urucú Small tree or shrub 2–6m 0 – 1,200 Lower Neotropical Human food rainforest and SubAndean Forest Borojoa patinoi Cuatrec. Rubiaceae Borojó, burijo, burojò Small tree 3–5m 0 – 600 Lower Neotropical rainforest Human food (fruits) Cajanus cajan (L.) Millsp. Fabaceae (Leguminosae) Guandú, Guandul, fríjol de palo, quinchoncho Shrub 1–3m 100 – 900 Lower Neotropical rainforest Human food (seeds) Canavalia ensiformis (L.) DC. Fabaceae (Leguminosae) Haba criolla, haba de burro, fríjol espada 40 – 900 Herb Lower Neotropical 0.6 – 1.3 m rainforest Cannaceae Achira, sagú, chisgua Herb 0.4 – 2.5 m 0 – 2,400 Lower Neotropical Human food (roots) rainforest and SubAndean Forest Caryodendron orinocense Karsten Euphorbiaceae Inchi, cacay, nuez llanera e iracana Tree 15 – 25 m 0 – 1,200 Lower Neotropical Human and animal food rainforest and Sub- (seeds) Andean Forest Erythrina edulis (L.) DC. Fabaceae (Leguminosae) Balú, chachafruto, sachafruto, frijol nopas, poroto Tree 6 – 10 m Human food (seeds) and 1,400 – 2,400 Animal food (fruits and Sub-Andean Forest leaves), forage plant Amaranthus dubius Mart. ex Thell. Anacardium excelsum (Bertero & Balb.) Skeels Anacardium occidentale L. Canna indica L. 74 Family Bledo, bledo Amaranthaceae blanco, yuyo, amaranto CATALOGUE OF USEFUL PLANTS OF COLOMBIA Human food (seeds) and animal food (fruits, seeds and forage) CHAPTER 4 Scientific name Family Common name(s) Arazá, guayabo amazónico Eugenia stipitata McVaugh Myrtaceae Ipomoea batatas (L.) Lam. Camote, batata, Convolvulaceae chaco, papa dulce. Habit Altitudinal range (*) and habitat (**) Main uses Small tree or shrub 5 – 10 m 50 – 500 Lower Neotropical rainforest Creeping or climbing grass 0 – 2.400 Human food (roots) and Lower Neotropical animal food (roots and rainforest and Subleaves) Andean Forest Human food (fruits) Arrurruz, maranta, yuquilla, sagú 0 – 800 Herb Lower Neotropical 0.8 – 1.0 m rainforest Moriche, aguaje, buritì, morete Tree 20 – 35 m 0 – 900 Lower Neotropical rainforest Human and animal food (fruits) Camu camu Shrub 4–8m 0 – 300 Lower Neotropical rainforest Human food (fruits) Fabaceae (Leguminosae) Ahipa, ajipa, jacatupé Herb 30 – 60 cm 0 – 2,400 Lower Neotropical Human food (tuberous rainforest and Sub- roots, pods and seeds) Andean Forest Plukenetia volubilis L. Euphorbiaceae Sacha-inchi, sacha-maní, maní del inca Climbing grass 0 – 2,000 Lower Neotropical Human food (seeds) rainforest and SubAndean Forest Pouteria lucuma (Ruíz & Pav.) Sapotaceae Lúcuma, fruta de oro, oro de los incas Tree 13 – 15 m 1,500 – 2,400 Human food (fruits) Sub-Andean Forest Prosopis juliflora (Sw.) DC. Fabaceae (Leguminosae) Algarroba, trupillo, mezquite, cují yaque Tree 10 – 15 m 0 – 2,100 Lower Neotropical rainforest and SubAndean Forest Human food (fruits and seeds) and animal food (fruits, seed, leaves, forage) Sechium edule (Jacq.) Sw. Cucurbitaceae Guatila, chocho, chayote, papa de pobre Creeper 1,150 – 2,500 Human and animal food (fruits and roots) Solanum sessiliflorum Dunal Solanaceae Cocona, tupiro, topiro 0 – 1,000 Herb Lower Neotropical 0.8 – 2.0 m rainforest Theobroma bicolor Humb. & Bonpl. Malvaceae Pataxte, bacao, Small tree maraca, Mocambo 7 – 12 m de Chocó 20 – 1,000 Lower Neotropical rainforest Human food (fruits and seeds) Theobroma grandiflorum (Willd. ex Spreng.) K. Schum. Malvaceae Copoasù, copoazù, cacao blanco Tree 14 – 18 m 120 – 1,000 Lower Neotropical rainforest Human food (fruits and seeds) Nacedero, naranjillo, cajeto Tree 10 – 12 m 100 – 2,150 Animal food (tender Lower Neotropical leaves and stems as rainforest and Subfodder) Andean Forest Maranta arundinacea L. Marantaceae Mauritia flexuosa L.f. Arecaceae (Palmae) Myrciaria dubia (Kunth) Myrtaceae McVaugh Pachyrhizus ahipa (Wedd.) Parodi Trichanthera gigantea Acanthaceae (Humb. & Bonpl.) Nees CATALOGUE OF USEFUL PLANTS OF COLOMBIA Human food (roots) Human food (fruits) 75 CHAPTER 4 Acknowledgements Many people contributed to the production of this chapter. The authors wish to acknowledge the help provided by the ‘Useful Plants and Fungi of Colombia’ project team. The assistance provided by Tiziana Cossu, Laura Kor, Ian Ondo, Carolina Tovar and David Hammond was greatly appreciated. Special thanks go to Kaitalin White for the valuable technical and administrative support and dedicated time. 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CATALOGUE OF USEFUL PLANTS OF COLOMBIA 77 CHAPTER 5 Leaf beetle (Chrysomelidae, Coleoptera). 78 Carolina Castellanos-Castro CATALOGUE OF USEFUL PLANTS OF COLOMBIA Chapter 5 Insecticide Plants of Colombia Jennyfer Andrea Aldana-Mejía1, Henry Yesid Bernal M.2* & Alba Nohemí Téllez A.3 1 2 3 Doutoranda em Ciências Farmacêuticas, Laboratório de Farmacognosia, Faculdade de Ciências Farmacêuticas, Universidade de São Paulo, Campus de Ribeirão Preto, Ribeirão Preto, São Paulo, Brasil. j.aldana@usp.br; j.aldana@javeriana.edu.co Ex-Profesor Asociado del Departamento de Biología, Facultad de Ciencias, Universidad Javeriana, Herbario de la Pontificia Universidad Javeriana (HPUJ), Bogotá, D.C., Colombia. henryesid@gmail.com Profesor Titular del Departamento de Química, Facultad de Ciencias, Universidad Javeriana, Grupo de Investigación en Fitoquímica (GIFUJ), Pontificia Universidad Javeriana, Carrera 7 No. 43–82, Laboratorio 206, Edificio Carlos Ortiz, S.J. Bogotá, D.C., Colombia. ntellez@javeriana.edu.co *Corresponding author: henryesid@gmail.com Keywords: insectostatic, insectifugal, insectivorous plants, phytoecdysteroids, biocides, repellants, chemosterilisers. ABSTRACT Several plant species from Colombia are recognised as having some insecticide activity, but no publication has detailed this use to date. This study aimed to identify and characterise the plant species of Colombia that are used as insecticides, assessing the state of their traditional and scientific knowledge, and emphasising the native species of the country. A literature review of insecticide plants in Colombia allowed us to recognise 16 mechanisms of action and to classify them into four major categories: i) species that provide chemical pesticides; ii) insectostatic species; iii) insecticide species by mechanical means; iv) and species with unknown insecticide action. The main mechanisms of action reported were repellent (22.3%), followed by ‘insecticide by unknown means’ type (21.2%). In total, 632 species that are used as ‘insecticides’ were identified in Colombia, of which spermatophytes represent 98.9% and only 1.1% are lycophytes and ferns. The families Fabaceae, Asteraceae (Compositae), and Solanaceae present the largest number of reports on insecticide species. A total of 408 native species of Colombia used as insecticides were arranged into seven groups according to their geographical origin, including 11 endemic species: Ageratina ampla, Ageratina vacciniaefolia, Berberis monguiensis, Berberis samacana, Berberis tabiensis, Chromolaena barranquillensis, Espeletia killipii, Lourteigia microphylla, Mandevilla mollisima, Ocotea caparrapi, and Ryania speciosa var. chocoensis. Regarding the endemic species, eight (72.7%) present a wide geographic distribution in the country, and three species (27.3%) have restricted geographic distribution in the country, mainly in the departments of Boyacá, Cundinamarca, and Caldas. None of those Colombian endemic species presented sufficient evidence of scientific knowledge, and only a single species has sufficient evidence of traditional knowledge regarding its insecticide action. RESUMEN Numerosas especies vegetales presentes en el territorio colombiano se reconocen con algún tipo de actividad insecticida; sin embargo, no existen publicaciones que detallen el estado actual de este uso. Esta investigación se orientó a la identificación y caracterización de las especies de Colombia utilizadas como insecticidas y al establecimiento del estado actual de sus conocimientos tradicional y científico, con énfasis en las especies nativas del país. Una revisión bibliográfica de las plantas insecticidas de Colombia permitió reconocer 16 mecanismos de acción, y clasificarlos en cuatro grandes categorías, definidas como: especies proveedoras de plaguicidas químicos, especies insectostáticas, especies insecticidas por medios mecánicos y especies sin medio de acción conocido. El principal mecanismo de acción reportado fue el de repelente (22,3%), seguido por insecticidas por medios desconocidos (21,2%). Se identificaron 632 especies utilizadas en Colombia como insecticidas, de las cuales el 98,9% son espermatofitos y el 1,1% son musgos, hornworts, hepáticas o licofitas. Las familias Fabaceae, Asteraceae (Compositae), Solanaceae presentaron el mayor número de especies reportadas. Las 408 especies nativas de Colombia utilizadas como insecticidas se ordenaron en siete grupos de acuerdo con su origen geográfico, destacándose la presencia de 11 especies endémicas del país que son: Ageratina ampla, Ageratina vacciniaefolia, Berberis monguiensis, Berberis samacana, Berberis tabiensis, Chromolaena barranquillensis, Espeletia killipii, Lourteigia microphylla, Mandevilla mollisima, Ocotea caparrapi y Ryania speciosa var. chocoensis. De estas especies 8 (72,7%) son de distribución geográfica amplia en el país y 3 (27,3%) especies tienen distribución geográfica restringida en el país, con prevalencia en los departamentos de Boyacá, Cundinamarca y Caldas. Ninguna de estas especies endémicas de Colombia presentó suficiente evidencia de conocimientos científicos y una sola especie posee suficiente evidencia de conocimientos tradicionales para su utilización como insecticida. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 79 CHAPTER 5 INTRODUCTION Research and field collection of plants in Colombia have been carried out for more than 200 years for national and international herbaria or botanical gardens. Nowadays, the country holds more than 30,000 registered plant species, ranking second in the world in terms of plant richness (Andrade, 2011; SIB Colombia, 2020). Many of these plant species are considered important due to the environmental services they provide upon which human societies depend. Those useful or promising plants species have applications in different fields. Three main aspects lead to the great variety of useful plants in Colombia: the great variety of ecosystems, the fantastic plant species richness, and the diversity of human groups represented by nearly 100 different ethnic groups, including indigenous Afro-descendants, mestizos, and other traditional groups. The Flora of Colombia (native and exotic) is used in almost all daily activities: as artisanal resources, biocides, coagulants, colourings, construction materials, dyes, essential oils, flocculants, food, fuel or industrial oils, honey, housing products, insecticides, insectostatics, insects traps or catalysts, medicines, oilseeds, repellents, tanning agents, timber, waxes, and wrapping, and in cultural activities, magical-religious activities, and dentistry, among other uses. A strong example of useful plants of Colombia is the medicinal species identified and documented in recent studies carried out by Bernal & Mesa (Chapter 3). Ethnobotanical knowledge has been used as a starting point for developing new products (Ribeiro et al., 2017). Terms such as “bioprospecting” have been applied to define the search for new medicinal products (Garnatje et al., 2017). In general, many species that are considered useful and from which production chains have been derived (such as biocides, food, and textiles) have come from the link between traditional knowledge and its scientific application. Ethnobotanical bioprospecting for the identification of bioactive compounds has become an important tool that combines ethnobotanical, phytochemical, phylogenetic, and molecular data. From the traditional knowledge, it has been possible to validate scientific predictions about the biological potential of plants (Garnatje et al., 2017). The need for food and raw materials on a large scale, guaranteeing the profitability and economic stability of agroindustries, has generated dependence on synthetic chemicals for pest protection, causing an increase in environmental and health issues or a decrease in yield (Bonilla, 1991). Insecticides derived mainly from pyrethrum, rotenone, ryanodines, nicotine, neem extracts, and some essential oils (garlic, rosemary, eucalyptus, among others) have taken hold of the market, especially in developing countries (Isman, 2006; Isman, 2020). However, the agricultural use of plants as a protection resource against insects has been documented for more than two millennia in communities in ancient China, Egypt, Greece and India (Isman, 2006). Nowadays, due to the application of this traditional knowledge, several botanical products have been brought to the market to control and manage agriculturally 80 important pests. The use of plants or their chemical compounds constitutes an environmentally safer alternative, which could be combined with other techniques and methods (Cerón, 1996). It can be used as a complementary strategy in regulating insect pests, even if the full replacement of pesticides is impossible (Cerón, 1996). In general, an insecticide is a substance or compound that, due to the nature of its chemical structure, is used to kill or exert a biocidal action on insects (Lawrence, 2003; Silva et al., 2002). However, most plant species used to protect or control harmful pests do not exhibit any insecticide effect in the strict sense. Their mechanism of action is due to other phenomena (repellency, deterrence of feeding, among others) that also have a protective effect against insects (Silva et al., 2002). Under the classical perspective of pest control, the plant species that present insecticidal actions due to their physical or mechanical characters are not considered to be insecticide species. In this present study, the conventional concept of ‘insecticide species’ is adapted based on the literature review, aiming to include other mechanisms of insecticide action. Here, the reviewed definition of ‘insecticide plant’ includes all plant species or their infraspecific categories (subspecies, variety, subvariety, form, or subform) with biological actions that mediate or interfere directly in the physiological and behavioural processes of insects, preventing, mitigating, or eradicating the damage that the insects produce on other plants, humans and/or animals. Several plant species from Colombia are recognised as presenting insecticide activity (by chemical or mechanical means) or insectostatic action. However, there are no publications detailing the existing knowledge of this use. Therefore, this study was oriented toward identifying and characterising the traditional and scientific knowledge relating to species used as insecticides in Colombia, emphasising the native species of the country. Finally, this study aims to help the discovery of new and better options for pest control. MATERIALS AND METHODS The information gathered on plants used as insecticides in Colombia was initially based on searches using keywords related to the subject in different bibliographic sources. The subsequent step was conceptualising the terms that refer to the control and management of insects and the role of plants in these fields. From the initial review on ‘plant species with biological activity against insects’ was possible to identify two differentiated groups in the literature: (i) ‘insecticide species’, which are those that exert a lethal effect on the target insect, similar to traditional chemical pesticides; and (ii) ‘insectostatic species’, which act on the behaviour or development of insects, exerting a long-term effect (Table 1). The information was searched from libraries, documentation centres and national and international databases specialised in the use and exploitation of plant resources, CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 5 from the perspective of pest control and management, using keywords related to the mechanisms of action found. The classification and conceptualisation of the terminology of the mechanisms of action of plants used as insecticides in Colombia were based on the literature (Table 1). The mechanisms were divided into four main groups and sixteen subgroups. In many cases, the mechanism was not defined, and this led us to consider the mention of use as “insecticide by unknown means” as an independent category. It is important to note that this is a classification made by this study, which we put forward as a proposal for further in-depth future research. TABLE 1. Classification of the mechanisms of action of insecticide plants of Colombia. Plants Supplying Insecticides or Chemical Pesticides Plant species presenting a source of molecules or chemical substances that cause a toxic or lethal effect for insects (Pedigo et al., 2021; Vergara & Madrigal, 1994) Insecticide plants (by ingestion) Plants producing chemical compounds causing a toxic effect on the digestive system of insects (Abubakar et al., 2019; Alonso, 1999; Pedigo et al., 2021) Insecticide plants (by touch) Plants producing insect-killing chemical compounds in direct contact with insects (Alonso, 1999; Pedigo et al., 2021) Fumigant insecticide plants Plants with insecticide chemical compounds acting on target pests in the vapour or gas phase, leading to death (Abubakar et al., 2019; Pedigo et al., 2021; Rajendran & Sriranjini, 2008; Spochacz et al., 2018) Phototoxic insecticide plants Plants with photosensitive chemical compounds that can accumulate in insects and, when exposed to visible radiation, become phototoxic products, or cause lethal photochemical reactions (Ahmed et al., 2018; Pascual, 1996; Spochacz et al., 2018) Insectostatic Plants Plant species with chemical compounds that do not act directly and immediately as lethal agents but can contribute to the death or control of insects in the long term, intervening in their behaviour and/or development (Enrique et al., 2008; Montesino et al., 2009; Silva et al., 2002) Repellent or insect-repellent plants Plants with chemical compounds capable of making insects move away from the surface or structure from which they emanate due to the repulsion or irritation that they cause (Vergara & Madrigal, 1994; Enrique et al., 2008; Abubakar et al., 2019) Feed-regulating plants Plants with chemical compounds affecting the insect feeding process at any stage, either by inhibiting or interrupting it (Pascual, 1996; Abubakar et al., 2019; Enrique et al., 2008) RIG (Regulating Insect Growth) Plants with chemical compounds intervening in the functioning of the hormonal systems of insects, resulting in the alteration of physiological processes related to body growth (Abubakar et al., 2019; Pedigo et al., 2021) RIG plants, seedling inducers: phytoecdysteroids Plants with chemical compounds similar to ecdysones or hormones that induce seedlings, causing acceleration or advancement of seedlings, intervention in the formation of oocytes, generation of malformations, involvement in embryonic development and sexual immaturity of the adult male (Pascual, 1996; Primo, 1991; Vergara & Madrigal, 1994) RIG plants, seedling inhibitors: anti-hormones Plants with chemical structures capable of preventing the ecdysis process from taking place, causing the lengthening of the development phases and the subsequent death of the individual without the fulfilment of its normal life cycle (Pascual, 1996; Primo, 1991; Vergara & Madrigal, 1994). RIG plants, metamorphosis inducers: juvenile antihormones Plants with antagonist compounds of the juvenile hormone (JH), capable of generating effects such as early metamorphosis, the appearance of adults, sterility of females, reduction of fertility in males, interruption of embryogenesis and inhibition of sexual pheromone (Pascual & Robledo, 1998; Silva-Aguayo, 2021; Vergara & Madrigal, 1994) CATALOGUE OF USEFUL PLANTS OF COLOMBIA 81 CHAPTER 5 RIG plants, metamorphosis inhibitors: ‘phytojuvenoids’ Plants with ‘phytojuvenoid-type’ compounds, which fulfil roles analogous to those of the juvenile hormones of insects, and therefore impede metamorphosis and sterility by suspending embryogenesis and gametogenesis processes (Pascual, 1996; Primo, 1991; Silva-Aguayo, 2021) IGI plants (Inhibiting the Growth of Insects) Plants with chemical molecules capable of inhibiting chitin synthesis in insects, preventing them from completing ecdysis (Vergara & Madrigal, 1994) Plants regulating the oviposition of insects Plants with compounds capable of suppressing or interrupting oviposition through processes that cut off the needed perception of plant-insect stimuli (but not from any hormonal functioning intervention) (Montesino et al., 2009) Chemosterilizing plants Plants with chemical compounds that exert a definitive sterilising action on target insects (Carrero, 1996; Pedigo et al., 2021; Vergara & Madrigal, 1994) Insecticide Plants by Mechanical Means Species with particular morphological adaptations acting as adherent traps that impede insect movement and lead to death by starvation or that are digested as a complementary activity in the acquisition of nutrients (García-Saavedra et al., 2007) Insectivorous plants Plant species using morphological and anatomical adaptations to carry out the mechanical process of trapping and digesting insects using proteolytic enzymes (Alvarado et al., 2011; Fernández-Pérez, 1964; García-Saavedra et al., 2007; Lawrence et al., 2003) Plants with sticky traps Plant species with morphological characteristics including special structures which work as ‘sticky traps’ where insects can pose, fall, and die (LoPresti et al., 2015) Insecticides Plants by Unkown Means Plants reported with insecticide action, without specifying the type of biological action, nor how they exert it A database was built aiming to identify, compile, and analyse the existing traditional and scientific knowledge associated with the species used in Colombia as ‘insecticide’ and ‘insectostatic’ plants, containing the following information for each species: i) kingdom, ii) sub-kingdoms, iii) division (phylum), iv) class, v) order, vi) botanical family, vii) valid scientific name, viii) synonyms, ix) common names, x) geographical distribution, xi) insecticide action mechanisms, xii) bibliographic references, xiii) indicators of ethnobotanical knowledge, and xiv) indicators of scientific knowledge. The geographical distribution for each species was organised into four large groups: 1) wild plants, 2) cultivated plants (native or exotic), 3) naturalised plants, and 4) species with uncertain geographic distribution. The wild (non-cultivated) species were organised into three groups: 1) native, indigenous, or autochthonous plants of Colombia; 2) non-native, introduced, or exotic plants, and 3) cosmopolitan plants. The native species used as insecticide plants in Colombia were organised into seven groups: 1) endemic species of Colombia (E); 2) native species of the Neotropics (N) with presence in Colombia; 3) native species of Mesoamerica (M) with presence in Colombia; 4) species native to South America (S) with presence in Colombia (in two or more non-Andean South American countries); 5) species from Mesoamerica and South America, present in Colombia (M + S); 6) Mesoamerican and Andean species present in Colombia (M + A), and 7) native plants of the Andean region (A) present in Colombia (in two or more Andean countries: Colombia, Ecuador, Peru, Venezuela, and Bolivia). 82 Parameters included in semi-structured interviews from ethnobotanical studies on useful plants, such as type of use, used organs, type of use, preparation and dosage (EspinosaJiménez et al., 2021; Ahmad et al., 2014; Gu et al., 2020), were used define the information compiled on traditional knowledge. Available information on the spectrum of action and possible toxicity were added to the above parameters. The compilation of the state of scientific knowledge was based on the search for information on characterisation, efficacy, and safety in the use of insecticide plants. The search included literature on the characterisation of insecticide plants (taxonomic, systematic, biogeographical, and ecological studies), efficacy analysis of insecticide plants (phytochemical and biological activity studies), and safety analysis (toxicity studies). The indicators of traditional knowledge related to each species were: a) insecticide actions (mention of insecticide use), b) part(s) used, c) traditional preparation mode, d) route of administration (topic or systemic), e) method of use (dose), f) method of use (posology), g) spectrum of action (low 1–3 pests, medium 4–6 pests and high 7 or more pests) and h) toxicity (low, medium, or high). The indicators of scientific knowledge related to each species were: a) geographical distribution, b) altitudinal distribution, c) ecological characterisation, d) morphological characterisation, e) molecular characterisation, f) taxonomic description and diagnosis, g) methods for identification and taxonomic determination, h) toxicity tests in insects (LD50 and concentrations), and i) tests of CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 5 insecticide use efficacy (phytochemical investigations and biological activities investigations). In the phytochemical investigations, the phytochemical profiles (extraction, fractionation or separation, qualitative characterisation, isolation, and purification) and the identification of the substances (structural elucidation) were recorded. Regarding the investigation of biological activities, in vivo activities (adults, eggs, larvae, pupae, and nymphs) and spectrum of action (low 1–3 pests, medium 4–6 pests and high 7 or more pests) were recorded. For each indicator, scores were assigned according to the absence (0) or presence (1) of bibliographic information to conduct a quantitative assessment of each level of knowledge. Considering that the indicators ‘action spectrum’, ‘toxicity’, and ‘phytochemical profile’ were subdivided into several levels according to the complexity of the information, the scoring scale was extended. It could also range from zero (no information) to 4 (high spectrum of action, high toxicity/ LD50, and isolation/purification of active substances). Some ethnobotanical (traditional form of preparation, route of administration, method of use, and toxicity) and scientific indicators (toxicity tests on insects) were considered ‘not applicable’ in the quantification of data for plant species classified as insecticides by mechanical means. A scale of scores was established to determine the level (low, medium, or high) of each species’ state of ethnobotanical knowledge. The scale was differentiated according to the category of insecticide use. For example, for plants exerting non-mechanical mechanisms of action, within a maximum range of 13, it was determined that a score of 0–4 indicates a low state of knowledge; 5–9 intermediate, and 10–13 high. For insectivorous plants and ‘sticky trap’ species, within a maximum range of five, it was defined that a total score between 0–1 represents a low state of knowledge, between 2–3 medium, and 4–5 high. At the scientific level, for species with non-mechanical mechanisms, within a maximum range of 34, it was defined that scores between 0–11 represent a low level of knowledge, 12–23 medium, and 24–34 high. For insectivorous plants and ‘sticky trap’ species, within a maximum range of 31, it was defined that scores between 0–11 represent a low state of knowledge, 12–22 intermediate, and 23–31 high. The identification and characterisation of the insecticide plants of Colombia were carried out based on the review of 432 bibliographic references on the subject, from which a corresponding list was constructed. Owing to its length, only the basic bibliography is presented in this publication. For each species presenting information on insecticide uses in Colombia, a search was carried out from the national herbaria of the country that presents online data accessible from the Global Biodiversity Information Facility (GBIF) (Table 2). National and international databases were also consulted regarding species nomenclature, geographical distribution, altitudinal distribution, and bibliography (Table 3). TABLE 2. Consulted Colombian herbaria from the Global Biodiversity Information Facility (GBIF). Herbarium Colombian Amazon Herbarium related to the Amazon Institute for Scientific Research, Sinchi Colombian National Herbarium Herbarium of the University of Antioquia Universidad Católica de Oriente Herbarium Botanical Garden of Bogotá “José Celestino Mutis” Herbarium Acronym City, department COAH (Sinchi) Bogotá, Bogotá D.C. https://sinchi.org.co/coah COL Bogotá, Bogotá D.C. http://www.biovirtual.unal.edu.co/es/ colecciones/search/plants/ HUA Medellín, Antioquia https://www.udea.edu.co/wps/portal/udea/ web/inicio/unidades-academicas/cienciasexactas-naturales/herbario#:~:text=El%20 herbario%20de%20la%20Universidad,y%20 otras%20regiones%20de%20Colombia HUCO Medellín, Antioquia https://www.uco.edu.co/herbariouco/Paginas/ Catalogo-Virtual.aspx Bogotá, Bogotá D.C. http://herbario.jbb.gov.co/especimen/simple JBB Web site Guillermo Piñeres Botanical Garden Herbarium JBGP Cartagena, Bolívar http://www.jbgp.org.co/el-herbario/ Juan María Céspedes Garden Herbarium TULV Tuluá, Valle del Cauca https://www.inciva.gov.co/patrimonio-turistico/ jard-iacute-n-bot-aacute-nico-juan-maria-ceacute-spedes Herbario Forestal Gilberto Emilio Mahecha Vega of the District University UDBC Bogotá, Bogotá D.C. http://herbario.udistrital.edu.co/herbario/index. php?option=com_content&view=article&id=4&I temid=3 CATALOGUE OF USEFUL PLANTS OF COLOMBIA 83 CHAPTER 5 TABLE 3. National and international databases consulted as online resources. Database 84 Web site BHL. Biodiversity Heritage Library https://www.biodiversitylibrary.org/ BIOVIRTUAL (Common names). Institute of Natural Sciences – National University of Colombia http://www.biovirtual.unal.edu.co/ nombrescomunes/es/ BIOVIRTUAL (Herbarium specimens). Institute of Natural Sciences – National University of Colombia http://www.biovirtual.unal.edu.co/es/ Catalogue of plants and lichens of Colombia http://catalogoplantasdecolombia.unal. edu.co/es/ CBIF. Canadian Biodiversity Information Facility https://www.cbif. gc.ca/acp CEIBA. Cataloguer of Biological Information – Instituto de Investigación de Recursos Biológicos Alexander von Humboldt-Colombia http://www.i2d.humboldt.org.co/ceiba/ Colombia Biotic Diversity, Institute of Natural Sciences, Museum of Natural History of the National University of Colombia http://colombiadiversidadbiotica.com/ Sitio_web/Bienvenida.html CoL. Catalogue of Life https://www.catalogueoflife.org/ ColPlantA. Colombian Resources for Plants https://colplanta.org/ DR. DUCKE’s. Phytochemical and Ethnobotanical Databases https://phytochem.nal.usda.gov/ phytochem/search EOL. Encyclopedia of Life. National Museum of Natural History, Smithsonian https://eol.org/ Flora Of Colombia. Monographs of the Institute of Natural Sciences of the National University of Colombia. http://ciencias.bogota.unal.edu.co/menuprincipal/publicaciones/biblioteca-digital/ flora-de-colombia/ Flora of the Royal Botanical Expedition. Royal Botanical Garden of Spain https://bibdigital.rjb.csic.es/ GBIF. Global Biodiversity Information Facility https://www.gbif.org/species/search Global Compositae Database. International Compositae Alliance https://compositae.org/ GRIN. Germplasm Resources Information Network https://www.ars-grin.gov/ ILDIS DATABASE. International Legume Database and Information Service http://www.ildis.org/ IPNI. International Plant Name Index https://www.ipni.org/ IUCN-Red List. The IUCN red list of the threatened species https://www.iucnredlist.org/ ITIS. Integrated Taxonomic Information System https://www.itis.gov/ JSTOR Global Plants Database https://plants.jstor.org/ Kew Herbarium Catalogue. Royal Botanic Gardens, Kew http://apps.kew.org/herbcat/navigator.do Kew Plant Information Portal. Napralert Database. University of Illinois at Chicago https://www.napralert.org/ NCBI. National Center for Biotechnology Information. National Library of Medicine, National Institutes of Health, USA https://www.ncbi.nlm.nih.gov/ NMNH. Smithsonian National Museum of Natural History https://collections.nmnh.si.edu/search/ botany/ PL@NTUSE. Plant uses https://uses.plantnet-project.org/en/ Main_Page RACCEFYN. Journal of the Colombian Academy of Exact, Physical and Natural Sciences https://raccefyn.co/index.php/raccefyn CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 5 Database Web site REFLORA. Virtual herbarium of the flora of Brazil. Rio de Janeiro Botanical Garden http://reflora.jbrj.gov.br/reflora/ herbarioVirtual/ConsultaPublicoHVUC/ ConsultaPublicoHVUC.do Science Direct https://www.sciendedirect.com/ SCOPUS-Elsevier. Bibliographic database http://www.scopus.com SIAC. Colombia’s environmental information system http://www.siac.gov.co/biodiversidad SIB. Colombia’s Biodiversity Information System https://sibcolombia.net/ SPECIES 2000 and ITIS. Catalogue of Life. Annual checklist. Integrated Taxonomic Information System http://www.catalogueoflife.org/annualchecklist/2019/info/about The Plant List http://www.theplantlist.org/ Tropicos. Vascular Plants of the Americas http://legacy.tropicos.org/project/VPA USDA-Plants Database. United States Department of Agriculture http://www.plants.usda.gov Useful Tropical Plants Database http://tropical.theferns.info/ WCSP. World Checklist of Select Plant Families. Royal Botanic Gardens Kew https://wcsp.science.kew.org/qsearch.do WCVP. The World Checklist of Vascular Plants. Royal Botanic Gardens Kew https://wcvp.science.kew.org/ WFO. World Flora Online Consortium http://www.worldfloraonline.org/ Eudicots RESULTS AND DISCUSSION From the literature search, a total of 632 plant species with reports of insecticide activity were identified as presenting occurrence in Colombia, including 625 species (98.9%) of spermatophytes and seven species (1.1%) of lycophytes and ferns. The complete list of species and associated information on insecticide plants in Colombia has been added to the ColPlantA portal (https://colplanta.org), Royal Botanical Garden, Kew. The species recorded as insecticide belong to 44 orders, distributed in 117 botanical families. The highest percentages of insecticide useful species were found among the families: Asteraceae (Compositae), Euphorbiaceae, Lamiaceae, Leguminosae (Fabaceae), and Solanaceae. From the 117 botanical families identified, 41 Eudicots a A 3.2% Lentibulariaceae families (35%) presented a single species with associated information as ‘insecticide use’. 3.7% Euphorbiaceae The non-monocots flowering plants represented 562 species distributed within 5.0%94 botanical families. The Lamiaceae highest percentages of species with insecticide action were represented by Leguminosae 5.3%(Fabaceae) with 92 species, Solanaceae Asteraceae (Compositae) with 66 species, Solanaceae with 30 species, and Lamiaceae with 28 species 11.7% (Figure 1a). The 60 Asteraceae species of monocots reported as insecticides were distributed in Leguminosae 18 botanical families, with Poaceae (Gramineae) standing 16.4% out with 16 species, Araceae with ten, Arecaceae (Palmae) with eight, and Zingiberaceae with five (Figure20.0% 1b). Three species 0.0% 10.0% 30.0% of gymnosperms with presence in Colombia were registered. Monocots bB Lentibulariaceae 3.2% Amaryllidaceae Euphorbiaceae 3.7% Cyperaceae Lamiaceae 5.0% Zingiberaceae Solanaceae 5.3% Arecaceae 11.7% Asteraceae 0.0% 10.0% 6.7% 8.3% 13.3% 16.7% Araceae 16.4% Leguminosae 5.0% 20.0% 26.7% Poaceae 30.0% 0.0% 10.0% 20.0% 30.0% FIGURE 1. Taxonomic coverage of the main botanical families of A Eudicots and B Monocots, as percentage of species number/family. Monocots Amaryllidaceae Cyperaceae 5.0% 6.7% Figure 1. Taxonomic coverage on the main botanical families of a in percentage of species number/ family. 85 Monocots, CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 5 Seven insecticide species representing lycophytes and ferns were identified, distributed in six botanical families (Figure 2). Lycopodiaceae is the only family reported with more than one record, including two species (28.6%), the other species are ferns, one species (14.3%) from Equisetales, and four species (57.2%) of Polypodiales. Considering the 632 species reported as insecticides in Colombia, the lack of information on the particular mechanism exerted by the species on the ‘target’ insects category is highly prevalent any level of knowledge, being the second most frequently mentioned category in the literature (Figure 3). The categories ‘repellent insecticide’ and ‘feeding regulation insecticide’ were the most common and third most common categories. However, it was Lycophytes and Ferns found that the categories comprising specific biological actions that have been studied at the scientific level (for example, insecticide related to ‘phototoxic’, ‘inducer of Dryopteridaceae 14.3% metamorphosis’, ‘inducer of moulting’, ‘chemosterilating’, and ‘growth regulator’) are those that have the least reports Dennstaedtiaceae 14.3% among the native species of Colombia. From the total number of species used as ‘insecticide’ Davalliaceae 14.3% plants in Colombia, 408 species are considered native to the country, including 405 (99.3%) spermatophytes and three (0.7%) ferns and lycophytes. The native species belong to Equisetaceae 14.3% 38 orders, distributed in 90 botanical families. The native species to Colombia grouped by geographical distribution Lycopodiaceae 28.6% are represented by 222 (54.8%) Neotropical species, 77 (19.0%) Mesoamerican and South American species, 39 0.0% 20.0% 40.0% (9.6%) Andean species, 28 (6.9%) Mesoamerican species, 27 (6.7%) South American species, 11 (2.7%) species Figure coverage on the themain mainbotanical botanical families of lycophytes and ferns, endemic to Colombia, as in well as one (0.2%) Mesoamerican FIGURE2.2. Taxonomic Taxonomic coverage of on families percentage of species number/ family. and Andean native species (Figure 4). of lycophyte and ferns, in percentage of species number/family. 0.0% 5.0% 10.0% 15.0% 20.0% 25.0% Repellent or insect repellent Insecticide (unknown measures) Food regulator Insecticide (touch) Fumigant Insecticide (ingestion) Oviposition regulator Growth inhibitor Insectivore Provider of chemical pesticides Seedling inhibitor Sticky trap Inhibitor of metamorphosis Growth regulator Chemosterilizer Seedling inductor Metamorphosis Inductor Phototoxic FIGURE 3. mechanisms of action forofinsecticidal plants with occurrence in Colombia. Figure 3.Registered Registered mechanisms action for insecticidal plants with occurrence in Colombia 86 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 5 2.7% Neotropics natives 0.2% 6.7% Mesoamerica and Suramerica natives 6.9% Andean natives 9.6% 54.8% Mesoamerica natives South America natives 19.0% Native endemics Mesoamerica and Andean natives 4. Geographical distributionofofthe the native native wild species of Colombia. Figure FIGURE 4. Geographical distribution wildinsecticide insecticide species to Colombia. Among the native species used as ‘insecticides’, 11 species were identified as endemic to Colombia: Ageratina ampla, Ageratina vacciniaefolia, Berberis monguiensis, Berberis samacana, Berberis tabiensis, Chromolaena barranquillensis, Espeletia killipii, Lourteigia microphylla, Mandevilla mollisima, Ocotea caparrapi, and Ryania speciosa var. chocoensis (Table 4). The majority of species (72.7%) present a wide geographic distribution in Colombia, and only three species (27.3%) have restricted geographic distribution in the country, with prevalence especially in the departments of Boyacá, Cundinamarca, and Caldas (Table 4). Among the native species of Colombia used as ‘insecticide’, none presented high levels of traditional and scientific knowledge about their insecticide activity. However, the endemic species Ageratina ampla is notable for presenting a high level of traditional knowledge, and it is also included in the category of insecticides by mechanical means, acting as a ‘sticky trap’ for insects, one of the least reported mechanisms reported in this study (Table 4). TABLE 4. List of species endemic to Colombia. 1 WGDS = Wide geographic distribution species; RGDS = Restricted georaphic distribution species; m a.s.l. (m) = above sea level (metres). 2 H = High; I = Intermediate; L = Low. State of 1 Ageratina ampla (Benth.) R.M. King & H. Rob. Compositae (Asteraceae) Eupatorium amplum Benth. 2 Ageratina vacciniaefolia (Benth.) R.M. King & H. Rob. Compositae (Asteraceae) Eupatorium confertifolium Klatt Eupatorium vacciniaefolium Benth. Synonyms Common names Jarillo, jarrillo Altitudinal range Geographical (m a.s.l.) and distribution and habitat departments1 Mechanism of action Scientific Scientific name and family Traditional No. knowledge2 1,750 – 3,400 Sub-Andean forest, Andean forest and Subpáramo WGDS Boyacá, Caldas, Cundinamarca, Meta, Risaralda, Tolima and Valle del Cauca Sticky trap H L 2,120 – 3,600 Sub-Andean forest, Andean forest, Subpáramo and Páramo WGDS Arauca, Boyacá, Casanare, Cauca, Cundinamarca, Meta and Santander Insecticide (unknown means) L L CATALOGUE OF USEFUL PLANTS OF COLOMBIA 87 CHAPTER 5 State of No. Scientific name and family Traditional Scientific 3 Berberis monguiensis L.A. Camargo Berberidaceae 3,120 Sub-páramo RGDS Boyacá Growth inhibitor, feed regulator L L 4 Berberis samacana L.A. Camargo Berberidaceae 2,425 – 2,980 Andean forest RGDS Boyacá Feed regulator L L 5 Berberis tabiensis L.A. Camargo Berberidaceae Uña de gato 1,750 – 3,670 Sub-Andean forest, Andean forest, Subpáramo and Páramo WGDS Antioquia, Boyacá, Chocó, Cundinamarca, La Guajira and Santander Insecticide (by touch) L I 6 Chromolaena barranquillensis Eupatorium (Hieron.) R.M. barranquillense King & H.Rob. Hieron Compositae (Asteraceae) Falso guaco, saragüay, rosita vieja, santa cruz, rosa vieja, santa maría 0 – 600 Lower Neotropical Rainforest WGDS Atlántico, Bolívar, Chocó, Córdoba, La Guajira, Magdalena and San Andrés Isla Repellent L L Frailejón, frailejón de Chisacá 2,120 – 4,010 Páramo, Subpáramo and sometimes found in the Andean forest RGDS Cundinamarca and Caldas Chemical pesticide supplier, feed regulator L I 1,990 – 3,600 Andes WGDS Boyacá, Cundinamarca, Magdalena, Tolima Feed regulator L L WGDS Antioquia, Boyacá, Caldas, Cauca, Cundinamarca, Huila, Magdalena, Nariño, Putumayo, Santander, Tolima, Valle Feed regulator L L 7 Espeletia killipii Cuatrec. Compositae (Asteraceae) 8 Lourteigia microphylla (L.f.) R.M.King & H.Rob. Compositae (Asteraceae) 9 88 Altitudinal range Geographical (m a.s.l.) and distribution and habitat departments1 knowledge2 Mandevilla mollissima (Kunth) K.Schum. Apocynaceae Synonyms Espeletia killipii Cuatrec. var. chicasana Cuatrec. Espeletia killipii Cuatrec. var. killipii Conoclinium microphyllum DC. Eupatorium microphyllum L.f. Echites mollissimus Kunth Exothostemon mollissimum (Kunth) G.Don Common names Chilca, jarilla, patinegra, salvia, trébol aromatizador Bejuco amarillo 880 – 2,000 Andes CATALOGUE OF USEFUL PLANTS OF COLOMBIA Mechanism of action CHAPTER 5 State of Ocotea caparrapi (Sand.-Groot 10 ex Nates) Dugand Lauraceae Ryania speciosa M.Vahl. var. chocoensis 11 (Triana & Planch.) Monach. Salicaceae Synonyms Nectandra caparrapi Sand.Groot ex Nates. Nectandra oleifera Posada Arango ex Nates. Oreodaphne oleifera Posada Arango. Common names Aceite de caparrapí, aceite caparrapí real, aceite de palo, aceituno, aguarrás, aymendron, amacey, caparrapí, calaba, canelo, canelo real hoja chica, canelo real caparrapí, comino, laurel canelo, palo de aceite, palo de caparrapí Ryania chocoensis Triana & Planch. Patrisa chocoensis (Triana & Plach.) Warb. Patrisia chocoensis (Triana & Plach.) Warb. From the total of native species of Colombia used as ‘insecticides’, only Nicotiana tabacum was associated with a high state of traditional knowledge and Ocotea caparrapi, an endemic species used as an ‘insecticide’ plant, presented an intermediate level of traditional knowledge. According to García (1992), a repellent action against mosquitoes is produced from burning the leaves of Ocotea caparrapi. However, this species is mostly recognised as medicinal, and there are no reports on the specific dose, dosage, and toxicity, neither any indication of use as an ‘insecticide’. Further endemic species of Colombia were mainly classified as ‘low’ as regards the level of traditional knowledge, and only a few records of use as an ‘insecticide’ were found for Ageratina vacciniaefolia, Chromolaena barranquillensis, and Ryania speciosa var. chocoensis. It is important to note that no research linking traditional and scientific knowledge for the endemic species of Colombia used as ‘insecticide’ was found. The species that presented the highest scores for traditional knowledge had ‘low’ scientific knowledge. Indeed, none of the endemic species presented a high level of scientific knowledge, and only Berberis tabiensis and Espeletia killipii presented intermediate levels. For example, there is evidence that the dichloromethane fraction of a stem extract from Berberis tabiensis is rich in the alkaloid tabienin B, and that it exhibits activity as a ‘contact insecticide’ Altitudinal range Geographical (m a.s.l.) and distribution and habitat departments1 Mechanism of action Scientific Scientific name and family Traditional No. knowledge2 1,000 – 2,380 Sub-Andean forests WGDS Antioquia, Cundinamarca and Santander Repellent I L 10 – 550 Lower Neotropical Rainforest WGDS Antioquia, Bolívar, Cauca, Chocó, Santander, Tolima and Valle del Cauca Insecticide (by touch) L L against Culex quinquefasciatus (Diptera: Culicidae) larvae, with an EC50 (effective concentration) of 78 mg/µL (Núñez, 2010). On the other hand, an extract of the leaves of E. killipii in petroleum ether, from which two of the grandifloric acid and kaurenal diterpenes were isolated and identified, showed insecticide activity against Premnotrypes vorax (Coleoptera: Curculionidae), an important pest of the potato – Solanum tuberosum (Marín, 1997). Acosta & Torrenegra (2011) also reported E. killipii as a regulating plant for feeding Lepidoptera larvae (Pterygote Subclass). In this case, the plant produces a leaf exudate rich in grandifloric acid, which has an antifeedant effect on insects, as a defence mechanism the attack by lepidopteran larvae (Acosta & Torrenegra 2011). A study on insecticide and antifungal activities and chemical constituents of the essential oils of Ocotea longifolia and O. macrophylla recorded a significant fumigant activity against Sitophilus zeamais (LC50 280, 5 µL/L air) (Prieto et al., 2010). Other species such as Ryania speciosa var. chocoensis, although presenting a low state of scientific knowledge with a lack of phytochemical characterisation, presented biological activity in vivo against Toxoptera citricidus (Hemiptera: Aphididae), with LC50 (lethal concentration) of 236 mg/L and 68 mg/L of the ethanolic extracts from stems and leaves after 36 hours of contact (Méndez et al., 2011). On the other hand, species with a high state of scientific knowledge, such as Annona CATALOGUE OF USEFUL PLANTS OF COLOMBIA 89 CHAPTER 5 cherimola, Annona squamosa, and Ageratum conyzoides, have intermediate to low levels of traditional knowledge. These results show that there are still gaps in the links between the different states of knowledge about native species that are used as insecticides. The study of insecticide plants has grown over the past few years. Since 1980, more than 20% of the publications on pesticides have been related to chemical pesticides made from plant resources (Isman & Grieneisen, 2014). Recently, there has also been increased pressure worldwide for the production of organic products as people are more aware of the risk of pesticides poisoning food and water and affecting ecosystems. Not only for countries in development but for any country, it has been very important to search for better methods of pest control that overcome the need for chemical pesticides (especially those targeting a wide range of pests) but do not present negative effects for the environment and human health (Amoabeng et al., 2014; Isman, 2020). Although the mechanisms of action of insecticide plants are diverse, the literature review shows that studies have mainly focused on the potential of some plant secondary metabolites with biological activity at a scientific level. The use of plant extracts that have insecticide activity is a highly explored alternative due to the possibility of synergistic effects between the active substances, the reduction in resistance to isolated active compounds, the easy access to the plant material, the simple extraction processes and the good cost-benefit when compared with conventional pesticides (Silva et al., 2002; Amoabeng et al., 2014; Miresmailli & Isman, 2014). A peculiarity of this study is the inclusion of the category of plants used as insecticides by mechanical means (insectivorous plants and ‘sticky traps’). Those species are still poorly studied compared to other categories. An important example of the importance of understanding and prioritising native insecticide plants of Colombia is the endemic species Ageratina ampla (Compositae-Asteraceae), which is reported as a ‘sticky trap’ for insects and presents sufficient evidence of traditional knowledge. The recognition of this species as an auxiliary/complementary method in the control of insects should be explored, considering that the mechanism of action is free from the disadvantages seen for conventional pesticides. Likewise, the recognition of endemic species that belong to botanical genera or families which already had secondary metabolites identified as insecticides is also a way forward in the search for alternative pesticides. For example, alkaloids such as ryanodine isolated from species of the genus Ryania, berberine and other alkaloids from Berberis, and acetogenins identified in species of genus Annona, are natural compounds with high potential for product development due to their biocidal capacity (Isman 2020; Méndez et al., 2011; Moreira et al., 2007; Rattan 2010). CONCLUSIONS The analysis of the insecticide mechanisms shows that the insecticide actions known from traditional knowledge 90 (repellency, fumigant activity, and toxicity by ingestion) have been brought forward for evaluation and scientific verification. The results showed that these actions are among the categories presenting the highest number of mentions in the literature. In general, the lack of knowledge on the mechanisms of action of many of these species is predominant, indicating an unclear validation of some of these insecticide uses. There is a predominant lack of knowledge on insecticide evidence for Colombian plant species, especially endemic species, at traditional and scientific knowledge levels. Scientific knowledge is more lacking than traditional knowledge, and the identification and prioritisation of useful species in projects of research, development, and innovation (R + D + I) still need to be deepened to allow the chemical synthesis or the sustainable use of plants as raw material to produce insecticide preparations or products. Likewise, there are still other barriers to be overcome in the development of plant-based insecticides, such as the lack of information on geographic characterisation, management and cultivation; the standardisation of the chemical composition of raw materials for the manufacture of insecticide products; production at large-scale; and the endorsement of controlling agencies to regulate the use and commercialisation (Isman, 2020; Miresmailli & Isman, 2014), as well as the study and alternative use of insecticide/insectifugal species that have non-traditional or unknown insecticide mechanisms. Chapter’s endnote: It is important to note that for this study, we did not follow the taxonomic criteria of some authors or databases (such as the World Checklist of Selected Plant Families or Plants of the World Online) for several of the insecticide species native to Colombia. As a result of our research, we found that the synonymies of Ocotea caparrapi with Mespilodaphne cymbarum (Ocotea cymbarum) and of Chromolaena barranquillensis with Chromolaena odorata were not applicable, as we found those species pairs to be divergent. Publications are in preparation to define the taxonomic, nomenclatural, geographical, and ecological distribution status of these species, which we consider to be endemic to Colombia. Acknowledgements The authors would like to thank Colciencias for funding this research through the Young Researchers and Innovators program “Virginia Gutiérrez de Pineda”, Convocatoria Nacional No. 525– 2011, and the Pontificia Universidad Javeriana, Bogota for allowing the development of research on its University campus. References Abubakar Y, Tijjani H, Egbuna, C, Adetunji C, Kala S, Kryeziu T, Patrick-Iwuanyanwu K (2019) Chapter 3. Pesticides, history, and classification. In: Egbuna C, Sawicka B, eds., Natural Remedies for Pest, Disease and Weed Control. Part I: Green approach to pest and disease control. Academic Press pp. 29–42. Acosta N, Torrenegra R (2011) Extracción, separación e identificación de la sustancia activa producida por Espeletia killipii como defensa CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 5 ante el ataque de larvas de lepidópteros. Rev. Fac. Cienc. Básicas, 7(1):126133. Acero L (2005) Plantas útiles de la cuenca del Orinoco. 2nd Edition. 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CATALOGUE OF USEFUL PLANTS OF COLOMBIA 91 CHAPTER 6 Symphonia globulifera L.f (Clusiaceae) has several uses in Colombia, such as Environmental, Fuel, Human Food, Material and Medicinal. 92 Dairon Cárdenas-López CATALOGUE OF USEFUL PLANTS OF COLOMBIA Chapter 6 Useful Plants of the Colombian Amazon Dairon Cárdenas-López†1, Nicolás Castaño-Arboleda1*, Nórida Marín-Canchala1 & Andrés A. Barona-Colmenares1 1 Herbario Amazónico Colombiano COAH, Instituto Amazónico de Investigaciones Científicas SINCHI. *Corresponding author: ncastano@sinchi.org.co Keywords: food, fuel, medicinal, ornamental, and timber ABSTRACT This study presents a synthesis of the current state of knowledge about the Useful Plants of Colombia, based on information gathered from field expeditions and records added from the Colombian Amazon Herbarium (Herbario Amazónico Colombiano (COAH)) database, which is managed by the Amazonian Scientific Research Institute (Instituto Amazónico de Investigaciones Científicas SINCHI). The useful plants were identified and classified according to their different categories of uses, their origin (native or endemic), and their extinction risk assessments. In total, 2,286 useful plants were recorded in the Colombian Amazon, representing 177 botanical families and 896 genera. Among the 12 categories of uses identified, the medicinal (883) and food (690) plants presented the highest number of useful species. Challenges to achieving the sustainable use of useful species in the Colombian Amazon are discussed from the economic botany perspective. RESUMEN Se presenta una síntesis del conocimiento de las plantas útiles de la Amazonia colombiana, a partir de los registros de la base de datos del Herbario Amazónico Colombiano (COAH) del Instituto Amazónico de Investigaciones Científicas SINCHI e información colectada en campo. Se registran 2.286 especies útiles, pertenecientes a 896 géneros y 177 familias y se identifican especies útiles nativas, endémicas, introducidas y con alguna categoría de amenaza. Se reconocen 12 categorías de uso en las que medicinal (883) y alimenticia (690) son las categorías que registran mayor número de especies. Se discuten los retos para alcanzar el aprovechamiento sostenible de las especies útiles en la Amazonia colombiana bajo la perspectiva de la botánica económica. INTRODUCTION The Colombian Amazon is an excellent expression of tropical forest biodiversity. Such high diversity has been attributed to the geological history resulting from the different tectonic movements that resulted in the Andean uplift, which involved different tectonic movements, changes in the direction of rivers, and sedimentation processes (Hoorn et al., 2010). Differences among tree communities have been suggested for geological units of the Colombian Amazon forest, the border of Orinoco savannas and the eastern border of the Andes (Cárdenas et al., 2017). The Guyana shield is represented by extensions of savannas (northeast part) and rocky outcrops (e.g., tepuis; as Serranía de Chiribiquete, Serranía de Naquén, Serranía de Araracuara, and some others distributed in the departments of Vaupés, Guainía, and Guaviare). Hernández (1984) estimates that more than 50% of Colombian living forms are represented there. According to Infante-Betancour & Rangel (2018), in the Colombian Amazon lowlands (Amazonia sensu stricto), there are 7,288 plant species recorded, whereas the Colombian Amazon Herbarium (COAH) holds records for 8,329 documented plant species (7,535 vascular and 794 non-vascular plants). Also, The Worldwide Fund for Nature (WWF) has identified several diverse ecoregions in the Amazon, such as the “Napo River Humid Forests”, the richest biota in the world, the western arc of the Amazon, and in particular the areas near the Andes foothills (Dinerstein et al., 1995; Cardoso et al., 2017). The Amazonian forests are considered a warehouse for multiple natural products, including a great variety of foods, medicinal supplies, and craft production items; they also represent an important refuge for plant species that constitutes a genetic bank for humanity (Leyva, 1998). However, a critical point in making the most of this high diversity is the low density of individuals presented by most plant species (Gentry, 1989). The first records on the use of Colombian Amazon plant species come from the notes of several naturalists about the local use of species, indigenous names for plants species, mythology and superstitions, and other aspects of indigenous life. The German Karl Friedrich Philipp von Martius was the first botanist to work in the Colombian Amazon, arriving by the Amazon River and crossing from the Caquetá River to the Araracuara rapids around 1820 (Schultes & Raffauf, 1994). His ethnobotanical notes were recorded in the monumental Flora Brasiliensis (Martius, 1829). For the 20th century, it is important to highlight several studies developed by Richard Evans Schultes (1970, 1979, 1983) and Schultes & Raffauf CATALOGUE OF USEFUL PLANTS OF COLOMBIA 93 CHAPTER 6 (1990, 1994), which provided valuable information on phytochemical and pharmacological aspects of Amazonian plants, also enhancing the richness of plant diversity and the lack of awareness of the possibilities for taking advantage of these resources. Enrique Pérez-Arbeláez’s masterpiece “Plantas Útiles de Colombia” was published for the first time in 1935, with many of the Amazonian plant species known at that time included. Likewise, stand out studies by J. M. Idrobo and A. Fernández were carried out in the 1950s using botanical collections with recorded information on plant uses in indigenous communities, mainly in the Igara-Paraná River sector. Later, within the framework of the Amazon Grammetric Radar Project (Colombian mapping Project – PRORADAM), Acero (1979) produced an essential contribution to the knowledge of the plant resources of the region, reporting 100 species with timber potential, whilst Pabón (1982) started the publication of the magazine “Colombia Amazónica”, reporting 327 useful plant species from the same region. On the other hand, Glenbosky (1983) carried out an inventory of medicinal plants used by the indigenous Tikuna of the Amazonian trapezium region, recording 184 species. Also, in 1983, la Rotta published a study describing the information recorded on 128 plant species used by the Andoque people, and later, la Rotta et al. (1987) carried out a similar study with the Miraña people from the middle Caquetá River. More recently, the Flora team of the Amazonian Scientific Research Institute (SINCHI) has carried out floristic inventories in the Colombian Amazon, obtaining an important reference collection and valuable information on the use of vegetation in several localities of the Amazonas, Guaviare, Putumayo, Guainía, and Vaupés departments (Cárdenas et al., 2002, 2012; Cárdenas & López, 2000; Cárdenas & Politis, 2000; Castaño et al., 2007). This chapter presents a synthesis of the knowledge on useful species in the Colombian Amazon Forest. MATERIALS AND METHODS Study area Several delimitations have been proposed for the Colombian Amazon through the years, based on different approaches and following biogeographic, hydrographic and/or politicaladministrative criteria (Domínguez, 1985; Rudas-Ll., 2009). The delimitation could include the Amazon and the Orinoco River basins, with the Vichada River as its northern limit. Physiognomic, climatic, and edaphic aspects were also included (Herrera, 1999; Domínguez, 1985; Rudas-Ll., 2009), as was the boundary of the Orinoco and Amazon River basins (Hernández-C. et al., 1992). From the east and south, the boundaries were defined by the neighbouring countries of Brazil, Ecuador, Peru, and Venezuela. To the west and north- FIGURE 1. Distribution map of the Colombian Amazon, highlighting its departments and conservation units. 94 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 6 west, the Amazon basin borders the Andes foothills (Andean Amazonian transition), the rocky outcrops of the Guiana Shield, the Amazon sensu stricto or Amazonian lowlands, and the Serranía de La Macarena (Rangel-Ch & Infante-Betancour 2018). The western limit for the Colombian Amazon is the elevation of 1,000 m of the eastern slope of the eastern mountain range (Cordillera oriental), where elements of the Amazonian tropical humid forest predominate (Figure 1). It is also essential to take into account that from the political point of view, the Amazon Forest covers the southern part of the Vichada department; the southeast of the Meta department; the entire territory of the departments of Amazonas, Caquetá, Guainía, Guaviare, Putumayo and Vaupés, as well as the Cauca Boot and the Amazonian slopes of Nariño department (which includes the middle part of the rivers Guamuez, Sucio, San Miguel, and Aguarico) (Salazar & Riaño 2016). Methods The potential uses of plant species were defined based on the information gathered from existing biological records in the database of COAH herbarium, obtained since 1983, and data from the vegetation surveys carried out in different projects at SINCHI. All species for which there are botanical records in the herbarium were considered, including those cultivated in orchards, gardens, and fields in the region. It is noteworthy that in the Amazon piedmont, many of the species are documented from photographic records of species growing in the residential gardens and orchards of settlers coming from the country’s interior. Subsequently, all the scientific names were validated using the Taxonomic Name Resolution Service tool v4.0 (Boyle, 2013). The records with taxonomic issues were reviewed, and those accepted in the Catalogue of Plants and Lichens of Colombia were considered (Bernal et al., 2016). Also, priority was given to names assigned recently by expert botanists. The record of threatened species was defined according to the categories assigned in two national resolutions (Resolución 1912/2017) (MADS, 2017), which establish the lists of wild threatened species of Colombia. The identification of endemic species was based on the Catalogue of Plants and Lichens of Colombia (Bernal et al., 2016) and supplemented with new records added to the COAH collection. The categories of use are those used on the COAH database, and they are not mutually exclusive, as follows: for handcrafted paper, wood for splint, trays, and containers. • Medicinal: species with curative and properties, according to local populations. preventive • Ornamental: species employed (or potentially employed) as an ornament and in the decoration of spaces. • Psychotropic: species that produce effects on the nervous system. • Timber: species used for house construction in the region and as commercial lumber. • Toxic: species used as poisons for hunting, or which are recognized as harmful to humans. RESULTS AND DISCUSSION The useful plants of the Colombian Amazon are represented by 2,289 species belonging to 896 genera and 177 families. Of these species, 2,211 correspond to angiosperms, 14 gymnosperms, 56 ferns, and five lycophytes. Some families have marked dominance: 60% of the registered speciesare concentrated within the 30 richest families. Among the most diverse families, we can highlight those that have important uses as timber (Lecythidaceae and Meliaceae), ornamentals (Orchidaceae), food and handcrafts (Arecaceae), or species with multiple uses as timber, food, and medicinals (Rubiaceae, Fabaceae) (Table 1). The genera with the highest species richness were Piper (Piperaceae) (49 spp.), Inga (Fabaceae) (41 spp.), Miconia (Melastomataceae) (38 spp.), Protium (Burseraceae) (32 spp.), and Solanum (Solanaceae) (27 spp.) (Figure 2). In contrast to what was observed for families, the richness of useful species at the generic level was more heterogeneous. The 30 most diverse genera account for only 22.8% of the species, while about 74% of the genera have only one or two useful species. TABLE 1. Botanical families with the highest number of reports of useful plants in the Colombian Amazon. Family Species Genera Fabaceae 210 78 Rubiaceae 107 45 Arecaceae 80 31 Melastomataceae 72 22 Annonaceae 70 14 Malvaceae 64 25 Euphorbiaceae 62 27 Araceae 62 18 Asteraceae 58 38 Apocynaceae 58 24 • Fuel: species used as firewood and for lighting. Piperaceae 58 2 • Handcrafts: species used as dyes, fibres for basketry, pulp Lauraceae 56 10 Moraceae 55 18 • Construction: species used in the structure, roofing, floors, and mooring of houses. • Cultural: species used in rituals and activities related to indigenous ancestral culture. • Dye: species that produce dyes or natural colours used in foods or crafts. • Food: Forest and cultivated species, used as edibles. • Forage: plant species used to feed animals. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 95 CHAPTER 6 TABLE 2. Numbers of useful plant species in the Colombian Amazon by category of use. 60 Number of spcies 50 Category of use Species 40 Medicinal 883 30 Food 690 20 Timber 401 10 Ornamental 392 0 Fuel 275 Handcrafts 233 Construction 211 Forage 81 Toxic 77 Cultural 65 Dye 39 Psychotropic 22 FIGURE 2. Plant genera with the highest number of useful species in the Colombian Amazon. Fig u r e 2 . Pla nt g e ne ra wit h t he hig he s t num b e r o f us e ful s p e c ie s in t he Co lo m b iaUseful n Am aspecies z o n. of the Colombian Amazon by category of use The species with potential uses recorded in the Colombian Amazon were classified according to 12 categories. They corresponded to 2,286 species representing 28% of the total number of species registered in this region (8,329 species). In this case, the Colombian Amazon presents a high proportion of species compared to the number of useful species for the entire country, considering that Pérez-Arbeláez (1978), in his publication “Plantas Útiles de Colombia”, considered the existence of about 1,850 species of useful plants in Colombia in the middle of the 20th century. On the other hand, the 252 species recorded by the ‘Useful Plants of the Orinoco Basin’ is very low if we consider that it includes species from cold, temperate, and warm zones of the Orinoco River basin in Colombia (Acero, 2005). However, it is important to highlight that for the Peruvian Amazon, 3,140 useful species were reported (Brack, 1993), representing a larger proportion of useful species than that reported in the present study for the Colombian Amazon. The regional richness of useful plants of the Colombian Amazon is extensive considering that it could represent about 50% of the useful plants recorded for entire South American countries. For example, 5,172 useful plant species were reported for Ecuador as the product of a several-year initiative with the participation of about 40 researchers (de la Torre et al., 2008). Those results show the importance of the Amazon for those countries where indigenous communities most widely use those species. The categories with the largest number of species were medicinals with 884 species, foods with 691 species including those introduced, and timber with 402 species, including a few introduced species (Table 2). A list of species by category is presented below, starting with the categories including the largest number of useful species: • Construction: 211 species were recorded, including those used in building structures [many genera of Annonaceae (Duguetia, Guatteria, Oxandra, and Xylopia); Lauraceae 96 (Aniba); Lecythidaceae (Eschweilera and Grias); and Myristicaceae (Osteophloeum and Virola)], roofing [some Annonaceae and Arecaceae (Iriartea deltoidea (bombona), Lepidocaryum tenue (puy), and Mauritia flexuosa (canangucha)], floors (several families), and the moorings of houses [such as Araceae (Heteropsis)]. • Cultural: this category includes 65 species that are important in local communities and represents the contextual basis for each culture. Several useful plants are key elements that identify each ethnic group (Garibaldi & Turner, 2004). They include, for example, yagé – Banisteriopsis caapi (Malpighiaceae), coca – Erythroxylum coca (Erythroxylaceae), tobacco – Nicotiana tabacum (Solanaceae), ortiga – Urera baccifera (Urticaceae), as well as many other plants used in rituals and daily activities depending on the different cultural backgrounds (Schultes & Raffauf, 1994). • Dyes/colourants: 39 species have been used as a dye, with a few used in rituals and others in the preparation of typical foods, such as the broadly used saino – Goupia glabra (Goupiaceae), Picramnia sellowi (Picramniaceae), azafrán de tierra – Escobedia grandiflora (Orobanchaceae), huito – Genipa americana (Rubiaceae), and achiote – Bixa orellana (Bixaceae). • Food: 690 species were recorded, including important forest species such as asaí – Euterpe precatoria (Arecaceae), madroño – Garcinia madruno (Clusiaceae), canangucha – Mauritia flexuosa (Arecaceae), milpesos – Oenocarpus bataua (Arecaceae), and cucuy – Macoubea guianensis (Apocynaceae). The presence of a significant number of introduced species is also highlighted (84 species) such as guanábana – Annona muricata CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 6 (Annonaceae), coco – Cocos nucifera (Arecaceae), coffee – Coffea arabica (Rubiaceae), mango – Mangifera indica (Anacardiaceae), and pomarrosa – Syzygium malaccense (Myrtaceae). Invasive species in this category include the african palm – Elaeis guineensis (Arecaceae) with a high potential of invasion in the Amazon region (Cárdenas et al., 2011). Still, its use is currently promoted by government agencies. However, it is known that it is contributing to the transformation of natural areas and consequently impacting the natural populations of native species, some of them threatened. An important part of the food category corresponds to the Non-Conventional Edible Plants (PANCs) – defined as native, not cultivated, and underutilized species (Marín, in press), which correspond to 605 registered species, representing a significant proportion of species included in the food category. • Forage: this category includes 81 species used to feed animals, which mainly represent the families Poaceae (20 species), Fabaceae (17 species), Cyperaceae (10 species), and Asteraceae (eight species). Some of them have been incorporated by the extensive livestock management in the Amazon region, such as: yaguará – Hyparrhenia rufa (Poaceae), guayacana – Imperata brasiliensis (Poaceae), caminadora – Rottboellia cochinchinensis (Poaceae), and several species from braquiarias – Urochloa (Poaceae). A few other species that are widely used as forage in Amazon region are: mataratón – Gliricidia sepium (Fabaceae), leucaena – Leucaena leucocephala (Fabaceae), botón de oro – Tithonia diversifolia (Asteraceae), and quiebrabarrigo – Trichanthera gigantea (Acanthaceae). • Fuel: 343 species from the Colombian Amazon are used as fuel, given the need to generate fire to carry out daily activities such as lighting the darkness of a maloca at night, preparing food or baking ceramics, among others. Previously, 295 species were recorded as being used as fuel in the Colombian Amazon region (Cárdenas et al., 2012), and 226 species were recorded as fuels in Ecuador (Palacios, 2008). Both studies emphasized that these species present special characteristics, such as suitability as firewood, carbon content and the presence of exudates that make them good fuels or combustion initiators (Palacios, 2008, Cárdenas et al., 2012). The species most used as fuel in the Colombian Amazon region are cucharo – Arawakia weddelliana (Clusiaceae), costillo – Aspidosperma excelsum (Apocynaceae), turí – Eschweilera rufifolia (Lecythidaceae), chaparro or motilón – Hieronyma alchorneoides (Phyllanthaceae), palo negro – Piptocoma discolor (Asteraceae), anime or copal – Protium apiculatum (Burseraceae), chimbe or bizcocho – Pseudosenefeldera inclinata (Euphorbiaceae), and seal or carate – Vismia baccifera (Hypericaceae). 1 • Handcrafts: 233 species were recorded in this category. Linares (1994) reported only 248 species in a preliminary inventory of the plants used to make handicrafts in Colombia, a number lower than expected considering the variety of ecosystems and landscapes in Colombia. A total of 114 species form a very important component of the artisanal species of Colombia, vegetable fibres that are extracted to produce traditional Colombian handcrafts (‘Artesanías de Colombia’), a fundamental basis of a national trade (Linares et al., 2008). The main species in this category are: fibre sources: cumare – Astrocaryum chambira (Arecaceae), papelillo – Couratari guianensis (Lecythidaceae), yanchama – Ficus maxima (Moraceae), chiquichiqui – Leopoldinia piassaba (Arecaceae), palo de algodón – Pseudobombax munguba (Malvaceae); wood carving: palisangre – Brosimum rubescens (Moraceae), granadillo – Platymiscium pinnatum (Fabaceae), and capinurí – Pseudolmedia laevis (Moraceae); seed sources: lágrimas de Job – Coix lacryma-jobi (Poaceae) and chochos – Ormosia spp. (Fabaceae); fruits used to make containers: totumo – Crescentia amazonica (Bignoniaceae); and fibres sources for basketry: different species of yaré – Heteropsis spp. (Araceae) are very popular throughout the region. • Medicinals: 883 species were recorded, with the following species standing out: andiroba – Carapa guianensis (Meliaceae), copaiba – Copaifera reticulata (Fabaceae), sangre drago – Croton lechleri (Euphorbiaceae), chuchuhuasa – Maytenus laevis (Celastraceae), bálsamo – Myrocarpus venezuelensis (Fabaceae), curarina – Potalia resinífera (Gentianaceae), caraño – Trattinnickia burserifolia (Burseraceae), uña de gato – Uncaria tomentosa (Rubiaceae). This category also includes some introduced species (95 spp.), such as paico – Chenopodium ambrosioides1 (Amaranthaceae), limón – Citrus limon (Rutaceae), totumo – Crescentia cujete (Bignoniaceae), and balsamina – Momordica charantia (Cucurbitaceae). Several of those species are shared with the Peruvian Amazon, where Brack (1993) reported 1,044 species with medicinal uses. Likewise, in Ecuador, 3,118 species were documented as “plants used to cure, alleviate, and treat human diseases” (de la Torre et al., 2008). In the Amazon, the medicinal species represent an important resource used by local communities, and the region presents a very high number of medicinal species according to the traditional knowledge, even though in most cases, there is still a lack of phytochemical and clinical studies documenting their true medicinal indications and effects (Taylor 2005). Studies by SINCHI found tannins, saponins, steroids, and triterpenoids, as well as terpenic lactones and coumarins, in caraño – Trattinnickia burserifolia (Burseraceae) (Cárdenas et al., 2002). In the Amazon region, this species is used for Chenopodium ambrosioides is accepted by Tropicos and Catálogo de plantas y líquenes de Colombia (Bernal et al., 2016). CATALOGUE OF USEFUL PLANTS OF COLOMBIA 97 CHAPTER 6 kidney diseases and as an antibiotic for skin problems (da Paz et al., 2004, Guimarães et al., 2014). This species is also used by the eastern Nukak (indigenous communities) from the Colombian Amazon for kidney diseases and as antibiotics (Cárdenas & Politis, 2000). For tigre guasca o carare – Aristolochia ruiziana (Aristolochiaceae), this study evidenced the presence of alkaloids in low quantity, steroids, and free triterpenoids (Cárdenas et al., 2002). In the region of Mocoa, this species is used to relieve cramps (Cárdenas et al., 2002). The same uses are reported from the scientific literature and attributed to its marked antispasmodic properties (against muscle spasms), which are potentially related to the presence of alkaloids and free triterpenoids (Cárdenas et al., 2002). The study also found flavonoids, leucoanthocyanidins, and tannins for chimbe – Senefeldera inclinata (Euphorbiaceae), besides a small number of steroids and triterpenoids (Cárdenas et al., 2002). Schultes and Raffauf (1990) have reported the direct use of S. inclinata scratched bark by the Achuar (Jibaros Mayna) – an indigenous ethnicity from Peru – as a treatment for toothaches. Alkaloids, tannins, saponins, and terpenic lactones were found in curarina – Potalia resinifera (Gentianaceae) (Cárdenas et al., 2002). In Putumayo and other regions of the Amazon basin, this plant is used in plasters as a type of palliative for snakebite (Schultes & Raffauf, 1990). However, no pharmacological studies have been found to explain this activity. • Ornamentals: 392 species from the Colombian Amazon were recorded in this category of use, including 189 introduced species from several different countries. In this category, all species with current or potential ornamental uses were included. Although most of the native species are not used in the region for this purpose in the interior market of Colombia, there is an important demand for ornamental plants. This group includes many palms, heliconias, bromeliads, zamias, and orchids, in addition to specific species such as Goeppertia lanata (Marantaceae), Guacamaya superba (Rapateaceae), Schizaea elegans (Schizaeaceae), Schoenocephalium martianum and Schoenocephalium teretifolim (Rapateaceae), Victoria amazonica (Nymphaeaceae), and Vellozia tubiflora (Velloziaceae). This group of species could constitute an economic alternative for the region involving a program of propagation of promising species. This category has been influenced by the establishment of populated urban centres, farms, farmsteads, and dwellings in the Amazon region, as species were gradually brought from different parts of the world to recreate and decorate the environment with plants that adorned the places of origin of the new settlers. In this process, several species have been introduced, such as gold cup – Allamanda cathartica (Apocynaceae), araucaria – Araucaria columnaris 2 98 (Araucariaceae), bismarkia – Bismarckia nobilis (Arecaceae), flanboyan – Delonix regia (Fabaceae), royal palm – Roystonea regia (Arecaceae), and african tulipan – Spathodea campanulata (Bignoniaceae), among others. • Psychotropic: 22 species were recorded, such as yopo – Anadenanthera peregrina (Fabaceae), yagé – Banisteriopsis caapi (Malpighiaceae), coca – Erythroxylum coca (Erythroxylaceae), mamitavea – Iryanthera crassifolia (Myristicaceae), and tobacco – Nicotiana tabacum (Solanaceae), several of those 22 species have been identified and used by different indigenous peoples of the Amazon (Schultes & Raffauf, 1994). • Timber: 401 species were included in this category, a number close to the 468 forest species studied in the diagnosis of Flor de Agosto (Perú) and Tarapacá (Colombia) forests (Unda, 1993). Several of those species are among the 30 most-mobilized timber species from the natural forests of Colombia (Blanco, 2020). They are ranked among highly among the 875 species reported as timber by the Regional Autonomous Corporations of Colombia and Sustainable Development (Cárdenas & Salinas, 2007). Some of the most important species used as timber are cachicamo – Calophyllum brasiliense (Calophyllaceae), andiroba – Carapa guianensis (Meliaceae), abarco amazónico – Cariniana domestica (Lecythidaceae), palo de arco – Handroanthus serratifolius (Bignoniaceae), itauba – Mezilaurus itauba2 (Lauraceae), acapú – Minquartia guianensis (Olacaceae), achapo – Cedrelinga cateniformis and granadillo – Platymiscium pinnatum (Fabaceae). Some of those species were assessed as threatened according to the National List of threatened species (Cárdenas & Salinas 2007), such as the Critically Endangered (CR) palorosa – Aniba rosodora (Lauraceae), and the Endangered (EN) cedro – Cedrela odorata (Meliaceae), and canelo de los andaquíes – Mespilodaphne quixos (Lauraceae). There are also 14 species used as timber that have been introduced into the Colombian Amazon, such as melina – Gmelina arborea (Lamiaceae), ciprés – Hesperocyparis lusitanica (Cupressaceae), kaya – Khaya anthotheca (Meliaceae), pino caribea – Pinus caribaea (Pinaceae), and teca – Tectona grandis (Lamiaceae), some of them incorporated into agroforestry in the Guaviare and Caquetá departments. • Toxic: 77 toxicspecies have been recorded, including especially the curares [Abuta grandifolia, Curarea tecunarum (Menispermaceae), and Strychnos toxifera (Loganiaceae)]. Species with toxic properties are recognized under the name of curare, although their extract is drawn from different species and mainly used in hunting arrows (pers. obs.). Several cultivated species were recorded, including those from the Identified by Henk Van der Werff and accepted by Tropicos CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 6 fields and small orchards of indigenous communities, and especially those predominantly incorporated in agroforestry arrangements such as the food species: anón amazónico – Annona mucosa, guanábana – Annona muricata (Annonaceae), maní – Arachis hypogaea (Fabaceae), chontaduro – Bactris gasipaes (Arecaceae), papaya – Carica papaya (Caricaceae), ahuyama – Cucurbita maxima (Cucurbitaceae), yuca – Manihot esculenta (Euphorbiaceae), aguacate – Persea americana (Lauraceae), cocona – Solanum sessiliflorum (Solanaceae), copoasú – Theobroma grandiflorum (Malvaceae), and maíz – Zea mays (Poaceae). Likewise, some recorded cultivated species are exclusively medicinal, such as albahaca – Ocimum basilicum (Lamiaceae), or psychotropic, such as coca – Erythroxylum coca (Erythroxylaceae) and tobacco – Nicotiana tabacum (Solanaceae). Among the cultivated species, there are also species from other continents, incorporated in the near and remote past into the crops of the American indigenous peoples, such as mafafa – Colocasia esculenta (Araceae) from South Asia, plátano – Musa paradisiaca (Musaceae) from Asia, caña de azúcar – Saccharum officinarum (Poaceae) from New Guinea, and several species of citrus – Citrus spp. also from Asia (Purseglove, 1968). Endemic Useful Species The reports of Colombian endemic species are important to ensure the protection of those species that are only found in the country. And therefore, a protection strategy that guarantees their conservation is needed. Between the recorded useful plants of the Colombian Amazon, there are nine endemic species, including Guatteria araracuarae (Annonaceae) registered only in the region of Araracuara and used by the natives to make artisanal fishing tools; Astrocaryum cuatrecasasianum3 (Arecaceae) registered in the Amazonian piedmont between the departments of Caquetá and Putumayo, used by the local indigenous people as a food product; Ouratea chiribiquetensis (Ochnaceae) recorded exclusively in the Natural National Park Serranía de Chiribiquete, where it is used as fuel (firewood) by the local communities; and Schoenocephalium martianum (Rapateaceae) recorded in the region of Araracuara and used as ornamental. Threatened Useful Species Among the useful plants recorded for the Colombian Amazon, there are 19 species assessed in categories of risk of extinction (according to the IUCN criteria) and included on the National Resolution no 1912 de 2017 produced by the Ministry of Environment and Sustainable Development. Five of these species were assessed as Critically Endangered (CR) [e.g., Roystonea oleracea (Arecaceae), Aniba perutilis (Lauraceae), and Aniba rosodora (Lauraceae)]; seven as 3 Endangered (EN) mainly due to the overexploitation of wood: these included Cedrela odorata (Meliaceae), one of the species with the highest demand in the world (Cárdenas & Salinas 2007); Pachira quinata3 (Malvaceae); and Elaeis oleifera (Arecaceae), not very common in Amazonia, but mainly affected at the national level by fragmentation of the natural habitat. Also, seven species were assessed as Vulnerable (VU), including Caryocar amygdaliferum (Caryocaraceae), Podocarpus guatemalensis (Podocarpaceae), Syagrus sancona (Arecaceae), and Zamia amazonum (Zamiaceae). Among the threatened species at the national level, there are some introduced species that are considered to be transplants in the Amazon region. These include species assessed as Critically Endangered (CR), such as mahogany – Swietenia macrophylla (Meliaceae), abarco – Cariniana pyriformis (Lecythidaceae), and guayacán de bola – Bulnesia arborea3 (Zygophyllaceae), a timber species assessed as Endangered (EN) in the Colombian Amazon and used as ornamental (Cárdenas & Salinas, 2007). The Brazil nut – Bertholletia excelsa (Lecythidaceae) is a food species assessed as Vulnerable (VU). Although there are very few records in Colombia, these are found in the surroundings of human settlements, which is why they are surmised to be introduced individuals as natural populations of the species have never been reported in the national territory. Introduced Useful Species Biological invasions are regarded as the second cause of biodiversity loss and one of the five main drivers of change globally (McNeely et al., 2001). Any species introduction can become a biological invasion as some of the introduced species generate changes in the composition of species, displacement of native species and loss of biodiversity (Cárdenas et al. 2011). In this study, 323 introduced useful species were recorded, grouped in 85 families and 238 genera. This is a large number considering that 202 introduced species were reported previously for the region by Infante-Betancour & Rangel-Ch. (2018), and only 274 introduced plant species were recorded for the entire country according to Cárdenas et al. (2011). The families with the highest number of useful introduced plants were Fabaceae (26 species), Poaceae (21 species), Euphorbiaceae (17 species), and Lamiaceae (14 species). The main genera presenting the highest number of useful introduced species are Citrus (Rutaceae) and Euphorbia (Euphorbiaceae) (six species each), Hibiscus (Malvaceae) and Jatropha (Euphorbiaceae) (five species each), and Allium (Amaryllidaceae), Canna (Cannaceae), Ficus (Moraceae), Musa (Musaceae), Pelargonium (Geraniaceae), and Urochloa (Poaceae) (four species each). The categories of uses with the highest number of species introduced to the Colombian Amazon were Ornamental (191 species), Medicinal (102 species), Food (87 species), and Forage (19 species). Species accepted by Bernal et al., 2016 CATALOGUE OF USEFUL PLANTS OF COLOMBIA 99 CHAPTER 6 Useful species in large landscapes of the Colombian Amazon The following section presents the useful species related to large landscapes of the Colombian Amazon (sensu Duivenvoorden & Lips 1993). Some useful species are exclusive to the Terraces of the great rivers, where the greatest human activity of indigenous communities is concentrated. These include marañón de monte – Anacardium giganteum (Anacardiaceae), uflé – Anaueria brasiliensis4 (Lauraceae), barbasco – Caryocar nuciferum (Caryocaraceae), madroño – Garcinia madruno (Clusiaceae), siringa – Hevea spruceana (Euphorbiaceae), juansoquillo – Parahancornia oblonga (Apocynaceae), toñeka – Parinari montana (Chrysobalanaceae), and juansoco de alacrán – Rhigospira quadrangularis (Apocynaceae). Also, unique useful species were recorded for the dissected surfaces of sedimentary origin, such as cuchara caspi – Ambelania occidentalis (Apocynaceae), palo rosa – Aniba rosodora (Lauraceae), cachicamo o lagarto caspi – Calophyllum brasiliense (Calophyllaceae), sangre drago – Croton lechleri (Euphorbiaceae), charapilla – Dipteryx odorata (Fabaceae), chicle – Lacmellea edulis (Apocynaceae), Picramnia latifolia (Picramniaceae), and umarí – Poraqueiba sericea (Metteniusaceae). The Alluvial floodplains of the Andean and Amazonian rivers also have species that only occur there, including Anthodiscus amazonicus (Caryocaraceae), costillo – Aspidosperma excelsum (Apocynaceae), yagé – Banisteriopsis caapi (Malpighiaceae), lagarto o yacaré – Calophyllum longifolium (Clusiaceae), copaiba – Copaifera reticulata (Fabaceae), curare – Curarea tecunarum (Menispermaceae), oreja de chimbe – Erisma uncinatum (Vocysiaceae), palo de arco – Handroanthus serratifolius (Bignoniaceae), catahua – Hura crepitans (Euphorbiaceae), palo de algodón – Pseudobombax munguba (Malvaceae), Omphalea diandra (Euphorbiaceae), and uña de gato – Uncaria tomentosa (Rubiaceae). The rocky areas of sedimentary and igneousmetamorphic origin, recognized as the rocky outcrops in the northern part of the studied area, have relatively few records of only 13% (85 species) of the total useful species registered. Some of the useful species recorded for these ecosystems are Acanthella sprucei (Melastomataceae), Aechmea rubiginosa (Bromeliaceae), Epidendrum nocturnum (Orchidaceae), Ouratea chiribiquetensis (Ochnaceae), Pachira fuscolepidota (Malvaceae), Parahancornia surrogata (Apocynaceae), Molongum lucidum (Apocynaceae), Navia acaulis (Bromeliaceae), Schoenocephalium martianum (Rapateaceae), and Vellozia tubiflora (Velloziaceae). On the other hand, there are species with wide distribution in different landscapes such as cagüi – Caryocar glabrum (Caryocaraceae), fariñero – Clathrotropis macrocarpa (Fabaceae), surba o juansoco – Couma macrocarpa (Apocynaceae), fono – Eschweilera coriacea (Lecythidaceae), asaí – Euterpe precatoria (Arecaceae), Guatteria decurrens (Annonaceae), milpesos 4 100 Species accepted by Bernal et al., 2016 – Oenocarpus bataua (Arecaceae), and curarina – Potalia resinifera (Gentianaceae). All of them are characterised by their in high densities within the different landscapes, and some are included among the 227 most abundant species in the Amazon rainforest (Ter Steege et al., 2013). CONCLUSIONS In general terms, the knowledge on the useful plants of the Colombian Amazon has been tackled from an ethnobotanical perspective, which is the science focused on the relationship between plants and humans. However, it is important to approach this knowledge from the perspective of economic botany, the science dedicated to useful plants, which considers taxonomic, pharmacological, ecological, and economic aspects, allowing us to know the precise identification, potentialities for forest production, and supply and demand conditions for products in the Colombian markets (Ricker & Daly, 1997). It is also necessary to address this knowledge from the bromatological and phytochemical point of view to determine the species’ properties. 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CATALOGUE OF USEFUL PLANTS OF COLOMBIA 101 CHAPTER 7 Snake carved from the twisted stem of a liana of Entada polystachya, collected in Antioquia, Colombia and sent by William R. Jervis to Kew in the 1850s. 102 Economic Botany Collection, EBC 59030 CATALOGUE OF USEFUL PLANTS OF COLOMBIA Chapter 7 Colombian plants in Kew’s Economic Botany Collection Hernando Echeverri-Sanchez1,2, Fabio Ávila1, Natalí Sánchez-Garzón1 & Mark Nesbitt1* 1 2 Royal Botanic Gardens Kew, Richmond, Surrey, TW9 3AE, UK Department of Anthropology, University College London, 14 Taviton St, Bloomsbury, London WC1H 0BW, UK *Corresponding author: m.nesbitt@kew.org Keywords: ethnobotany, ethnobiology, museum, history, empire ABSTRACT Kew’s Economic Botany Collection was first founded as the Museum of Economic Botany in 1847. Among a total collection of 100,000 plant raw materials and plant products, it contains 320 specimens from Colombia. Recent research has enabled botanical names to be updated and provenance data retrieved through archival research. The Colombian specimens are best understood as representing changes in the scientific relationship between Colombia and Kew from the mid-nineteenth century to today. We identify three stages in bioprospecting. In the nineteenth century, medicines, dyes, and other commercial materials were sent to Kew for evaluation and subsequent display, often by British residents in Colombia. The Royal Pharmaceutical Society Museum, now part of the Kew collection, also includes medicinal plants from Colombia. In the early twentieth century, the Colombian government employed Morley Thomas Dawe to explore Colombia’s agricultural and botanical potential, leading to the donation of 70 specimens to Kew. Later in the twentieth century, Colombian botanists donated timber specimens. In the twenty-first century, the Colombia Bio project (between Colombia and Kew) led to the acquisition of new objects, emphasising the development of sustainable livelihoods and the preservation of biocultural knowledge. RESUMEN La Colección de Botánica Económica de Kew fue fundada como Museo de Botánica Económica en 1847. Con un total de 100.000 materias primas vegetales y productos vegetales en la colección, 320 especímenes son de Colombia. Investigaciones recientes han permitido actualizar los nombres botánicos y recuperar datos de procedencia mediante la investigación de archivos. Los especímenes colombianos representan cambios en la relación científica entre Colombia y Kew desde mediados del siglo XIX hasta la actualidad. Identificamos tres etapas en la bioprospección. En el siglo XIX, diversos medicamentos, tintes y otros materiales comerciales se enviaron a Kew para su evaluación, a menudo por parte de residentes británicos en Colombia. El Museo de la Real Sociedad Farmacéutica, ahora parte de la colección Kew, también incluye plantas medicinales de Colombia. A principios del siglo XX, el gobierno colombiano empleó a Morley Thomas Dawe para explorar el potencial agrícola y botánico de Colombia, lo que llevó a la donación de 70 especímenes a Kew. Posteriormente en el siglo XX se evidencia algún contacto con botánicos colombianos a través de donaciones de madera. En el siglo XXI, el proyecto Colombia Bio condujo a una intensa interacción entre Kew y Colombia, y la adquisición de objetos que enfatizan el desarrollo de medios de vida sostenibles y la preservación del conocimiento biocultural. INTRODUCTION The Economic Botany Collection (EBC) at the Royal Botanic Gardens, Kew (Kew) is a large, wide-ranging biocultural collection of over 100,000 materials and plant products spanning food products, tools, artefacts, medicines, poisons, wood, ornaments and textiles, among other objects, covering all aspects of plant uses in human life. The EBC is the successor to Kew’s Museum of Economic Botany, founded in 1847. Such museums of economic botany were widespread in the second half of the nineteenth century, connecting producers of raw materials with manufacturers. Economic botany collections are identifiable by their emphasis on usefulness to humans, by their distinctive combination of raw materials and products, and by the consistent application of botanical (Latin) names to the specimens (Cornish & Nesbitt, 2014; Nesbitt & Cornish, 2016). Economic botany was a fundamental part of the colonial system. By the 19th century, vegetable products were considered strategic elements, and oversight of their trade was a state priority (Drayton, 2000). As different colonial and commercial powers sought to control the economic potential of tropical biodiversity, they funded the creation and establishment of museums of economic botany. It is likely that similar materials, such as cinchona barks, were held by the ‘Botanical House’ in Bogotá: the base of the Real Expedición Botánica del Nuevo Reino de Granada. These specimens were removed to Madrid in 1816 (DíazPiedrahÍta, 1986). Although Colombia, with its recognised biological wealth, seems like a strategic source for economic botany specimens, it is surprisingly under-represented in the EBC. Unlike other countries of the region such as Mexico CATALOGUE OF USEFUL PLANTS OF COLOMBIA 103 CHAPTER 7 (850 items; Rico Arce et al., 2013) and Brazil (2,700 items), the Colombian component of the EBC is only 320 items (Echeverri-Sanchez et al., in press). As part of the larger Colombia Bio project, the authors of this chapter have catalogued, digitised, and studied these items and uploaded them to the Plants of the World Online (http:// www.plantsoftheworldonline.org/) and ColPlantA platforms (https://colplanta.org/). Updated records are also available on the EBC database, searchable online at: https://www. kew.org/science/collections-and-resources/collections/ economic-botany-collection. The Colombian element in EBC is small in number and is uneven in coverage of the many ways in which plants are used in the country. Although the EBC underrepresents the enormous botanical diversity of the country, it offers a useful perspective on British scientific and commercial interest in Colombia from the Victorian period to the current day. BRITISH – COLOMBIAN RELATIONS As a government institution since 1840, Kew’s relationship with Colombia reflected the official stance of the British government. In the second half of the nineteenth century, this was characterised by a tentative interest in the country’s natural wealth but little or no interest in formal administrative control. The British banking industry sought to make loans to the new nations of South America, establishing a debt system that, on many occasions, exerted a profound influence on these countries. This type of ‘soft’ power or, as historians now term it, ‘informal empire’, was considered more effective than formal political and economic control (Cain & Hopkins, 2016). However, British powers represented economic interests in the region, especially in countries with the greatest economic potential, such as Argentina, Brazil, and Chile (Thompson, 2008; Cain & Hopkins, 2016). Ensuring debt servicing of British loans and opening the Latin economies to English investors was London’s main policy in South America. In Colombia, informal political and economic control over the territory was rarely plausible. However, although Britain was one of the greatest investors in the young Andean nation, political instability, rurality, and geographical obstacles made it unattractive for English banks (Bushnell, 1993). Colombia was constantly trying to pay off large debts throughout its history and was quickly considered too risky for international investment (Deas, 1993, 2008). At the same time, the country had a relatively small luxury goods market and its rugged geography limited transportation. Foreign companies that tried to enter the domestic market often went bankrupt (Deas, 1993). Even large infrastructure investments, such as the Colombian rail network, failed. For these reasons, Colombia was one of the few countries in Latin American that did not produce large English fortunes, never being at the centre of British military and colonial interests. The only product that could generate wealth in the country was gold, and as a result, several mines were granted by 104 the Colombian government to pay interest on loans. These mines would be English cultural enclaves until the middle of the 20th century (Mejía Rivera, 2016; González Rodríguez, 2005). People working in the mines were one of the main sources of early collections in the EBC. They housed British doctors and engineers interested in local history and with the ability and interest to communicate directly with Kew. A search to find new medicines or industrial products was of great interest to local British enterprises. Scientists, merchants, and diplomats would send materials to Kew to explore their potential as a commodity. As the geopolitics of empire and trade changed during the 20th century, objects collected for the EBC reflect these changes. Founded in part as a tool to classify and control the biodiversity of tropical colonies, the purpose of such institutions has transformed profoundly. The differences between the objects collected at the beginning of the 20th century and those at the end of the century demonstrate the gradual change. When the British colonial project ended, these scientific institutions began to focus on documenting and understanding biocultural diversity to enable its preservation and use in local livelihoods. As such, items collected in the past 40 years are focused on preserving biocultural diversity, with fuller documentation of the techniques and knowledge associated with them. THE NINETEENTH CENTURY: SEARCHING FOR COMMODITIES By the time the EBC was constituted in the 1840s, the mercantilist trade dynamics of the nineteenth century had led to a focus on Europe as an importer of raw materials, and an exporter of manufactured goods, with an emphasis on trade within a country’s imperial territories. For countries outside the imperial system, entering this trade network required commodities and internal wealth that would incentivise foreign investment, which often came with heavy political and economic interventionism. The young South American nations were not part of any global imperial system at the time, following their recent independence from Spain and Portugal. They were often integrated into the north Atlantic trade network through a specific commodity or through banking and financial transactions. Throughout the history of the continent, several products caused exponential economic growth for local communities. Most of these products were raw resources produced from the local biodiversity, such as rubber, quinine, sugar, tobacco, alongside other lesser-known products, such as dyes and tropical fruits. But Colombia was not part of the British colonial sphere, so there was no interest in investing in national agricultural production. Instead, the country was understood as a possible source of resources that could break the existing monopolies, such as Brazil with rubber and Peru with quinine. The race to find species that produce quinine and rubber, evidenced in the collection, demonstrates the interest in these products in the British markets. At the same time, it highlights the efforts of the Colombian government to look for new products for global trade, leading in the early CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 7 Some specimens highlight a local interest in finding alternative products that could generate income. An early example of this is Té de Mutis (Symplocos theiformis), with several samples in the EBC collection. The earliest specimen (Catalogue number EBC 50445, Figure 1a) was donated to Kew in 1893 by London’s Pharmaceutical Society. This donation was an early attempt – involving the Spanish botanist Jose Celestino Mutis, resident in Bogotá – to commercialise an alternative to the lucrative tea (Camellia sinensis) trade. However, as a biodiverse region, there was much interest in the potential of local medicines found in the forest and jungles of the new nation. Several items in the collection, given by W.R. Jervis, demonstrate the role of British imperialism and trade in the formation of botanical medicine bioprospecting. Jervis was probably William R. Jervis, a doctor in the English mines of Marmato (Mejía Rivera, 2016; González Rodríguez, 2005). Though we have little information on him, Jervis was a proficient collector with over 30 items collected between 1825 and 1850. He was one of several British citizens who emigrated to Antioquia during the first half of the nineteenth century, when a small population of foreigners managed the British-owned gold mines. Jervis collected several specimens of cinchona bark (Cinchona spp., Figure 1b) and balsam (Myroxylon sp. and Myrospermum sp.), among other medicinal plants. The two examples of curare (Strychnos toxifera) from Colombia (EBC 49123 and EBC 49125) were sent to Kew by Jervis in 1853. Curare is used to poison darts used with blowguns or sometimes with arrows for hunting animals. It is prepared from the bark of Strychnos stems, mixed with a plant gum (usually from Couma macrocarpa latex) so that it will stick to the dart or arrow. Strychnos and other tubocurarinecontaining plants are used throughout the Amazon basin. It is unclear how Jervis obtained this specimen, which might have come from the Amazon region (far from Marmato). Nevertheless, several Strychnos species grow in Andean and inter-Andean valleys. The 1850s were when curare played an important role in understanding the usefulness of paralysis-inducing drugs in medical treatment. The EBC contains about 1,000 examples of Cinchona bark but only 32 specimens from Colombia. Cinchona bark A B twentieth century to financing European experts to visit Colombia and analyse the local biodiversity (see Morley Thomas Dawe, below) Unlike other Latin American nations, Colombia was relatively poor. Its economy was based mostly on gold extraction, with smaller subsistence farming throughout the Andean region and larger plantations on the coast. In the absence of important export products such as guano in Peru, cocoa in Venezuela, or rubber in Brazil, Colombia had little to offer to international markets. Those products that gained a certain international value had a limited effect on the national economy, thanks to mismanagement and civil conflicts, as in the cases of the tobacco boom of 1850 (Bushnell, 1993) and quinine in the 1860s (Tirado Mejida, 1971). It was only with coffee in the late 19th century that Colombia effectively entered world trade. This market was directed towards the United States rather than Great Britain but began consolidating Colombia’s influence throughout the continent. MEDICINES FIGURE 1. Medicines. A Té de Mutis (leaves of Symplocos theiformis), donated to Kew in 1893 by the Pharmaceutical Society of Great Britain (EBC 50445). B Bark of Cinchona pitayensis (Wedd.) Wedd., from Antioquia, Yarumal, sent to Kew by W.R. Jervis, 1854 (EBC 52393). CATALOGUE OF USEFUL PLANTS OF COLOMBIA 105 CHAPTER 7 A B C FIGURE 2. A Materials. Top. Leaves, twigs, and dyed fabric of ‘Bignonia tinctoria’ (this is an unplaced name according to Plants of the World Online; it is probably related to Fridericia chica (Bonpl.) L.G. Lohmann), given by Robert Thomson to the Pharmaceutical Society of Great Britain c. 1890 (EBC 47638). B Fibre from leaves of Furcraea sp., given to Kew as ‘picta’ by London brokers Messrs. Wigglesworth & Co in 1919 (EBC 29981). C Bag waterproofed with Castilla elastica Cerv. Sent to Kew in 1919 by Morley Thomas Dawe, described by him as ‘a rubber sack made by the Indians of that region [Darien Country]. They buy the material in the coastal towns or villages and waterproof it in the forests with the latex of Castilla elastica’ (EBC 43012). 106 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 7 was the only effective treatment for malaria in much of the world from 1630 to 1940. It was used as powdered bark until the extraction of the quinine alkaloids, which was perfected in the 1820s (Walker & Nesbitt, 2019; Rønsted et al., 2020). Since Peru had established a near-monopoly over Cinchona bark exports, Colombia had little effect on the international market (Crawford, 2016). However, there was a smaller-scale Cinchona bark industry in Colombia, which was characterised by a series of short-lived booms and busts over three periods: 1849–1852, 1867–1873, and 1877–1882 (Sastoque, 2011; Sandoval & Echandia, 1986; Tirado Mejida, 1971). Most of the quinine was harvested from the wild, and the samples found in the collection represent species common in Colombian cloud forests such as Cinchona pitayensis and Cinchona pubescens. Most of these specimens were donated to Kew by the Royal Pharmacological Society, which collected many samples from the London trade. From the 32 samples of the quinineproducing plants catalogued as Colombian, they represent five species of Cinchona, three species of Ciliosemina, and one each of the species of Carapichea and Ladenbergia (incl. Cascarilla). Colombia never had a sustained quinine industry, and when production shifted to Asia from the 1860s, Latin America lost its role in producing this important medicine. MATERIALS Dyes were of great interest to early Colombian and British bio prospectors. An interesting example is the chica dyes (Fridericia chica) held by the Pharmaceutical Society of Great Britain and transferred to Kew in 1983 (EBC 47638, Figure 2a). These were brought to London by Robert Thomson, a British gardener who visited Bogotá in the 1880s-1890s (Cendales Paredes, 2012). Even though this dye produced a beautiful red colour, correspondence in the Kew archives highlights that the colour does not last. This fact was the reason it was never a commercial alternative to cochineal dyes. Rubber was another neotropical product of great interest for the British empire. Although there are many rubber products in the EBC, there are no Colombian items made with Hevea brasiliensis. Instead, five objects are made of the rubber ‘substitute’ known as caoutchouc or Indian rubber (Castilla elastica, Figure 2c). Since H. brasiliensis only grew wild in the Amazon basin, until the development of Asian plantations late in the nineteenth century, Brazil had a monopoly overexploiting this important commodity. Due to its strategic value, there was a constant incentive to search for alternatives that could compete against the Brazilian industry. Although C. elastica has been used as a source of rubber by indigenous people since pre-Colombian times, it was never a realistic alternative due to its lower quality and production limitations (Hosler et al., 1999). There was also much interest in materials to make strong ropes, which were considered strategic in an age before plastics. In the early twentieth century, the British naturalist and collector Morley Thomas Dawe had a particular interest in the fique or Furcraea genus plants, the source of excellent rope materials (Figure 2b). In 1916 he wrote, “There are probably few vegetable products, the subject of domestic industry, particularly in the fibre world, which have so wide an application in everyday use as fique in Colombia” (Dawe, 1916). Even though the production of fique in Colombia was mostly to supply a growing internal industry, the trade potential of this product was not lost to Dawe, who, in that same bulletin, highlights the cost of production and export to ports such as New York. TIMBER A key factor in Colombia’s international trade was its access to transport. This fact is particularly the case with bulk materials such as timber. As late as 1916, its railways were poorly developed, with the main transport route being the Magdalena River. An easy connection between the country’s Pacific and Atlantic coasts awaited the opening of the Panama Canal in 1914 (Simmons, 1916). Simmons reported an extremely challenging environment for timber exports, with stands of two desirable timber trees, Spanish cedar (Cedrela odorata) and Colombia mahogany (Swietenia macrophylla), so scattered as to be economically inviable – at least for export. The wood component (xylarium) of the EBC contains about 42,000 specimens, with only 77 from Colombia. The history of the xylarium falls into two parts: collecting timber species from 1847 to the 1930s and then a broadening of interest to all woody species to support broad taxonomic surveys of wood anatomy (Cornish et al. 2014). The Colombian woods fall into two groups: a first period (1890– 1920), mostly of Dawe’s collections, and a second one (1960–1980) based on academic cooperation with the Universidad Nacional of Colombia (Figure 3). The first group contains three important timbers even today: Andean cedar (cedro de Bogotá, Cedrela montana), native pine (pino chaquiro, Prumnopitys montana), and laurel (comino crespo or chachajo, Aniba perutilis). Santiago Cortés (1897) highlighted these species as important for the timber industry, and all are now categorised as at threat of extinction. The second group of woody plants represented in the collection (and currently used in the country) belongs to 1960–1980 accessions. These items include commonly used plants in rural areas and are highly harvested throughout the country. Examples of this group are Couma macrocarpa (juansoco), Terminalia amazonia (arenillo), Cariniana pyriformis (abarco), and Bulnesia arborea (guayacán), which are also threatened (BGCI, 2021). Nowadays, according to MADS (2021), it is estimated that nearly 21,528 cubic metres of these trees were harvested between January 2020 and March 2021. RICHARD SPRUCE’S ETHNOBOTANICAL COLLECTIONS The scope of Kew’s Museum of Economic Botany was, from its foundation, defined as ‘all kinds of useful and curious Vegetable Products, which neither the living plants of the Garden nor the specimens in the Herbarium could exhibit’ CATALOGUE OF USEFUL PLANTS OF COLOMBIA 107 CHAPTER 7 (Hooker, 1855: 3; Cornish, 2012). In other words, not all specimens were purely utilitarian. This scope is reflected in the c. 350 ethnobotanical specimens collected from indigenous peoples in the Amazon region by the botanist Richard Spruce, mainly in the years 1850–1855 (Martins et al., 2021). Some of these are clearly of economic potential, as in the fruits or the caapi vine (Banisteriopsis caapi), a component of ayahuasca, which was of medicinal interest in Europe. Spruce also collected other items of less obvious usefulness in a European context, such as ritual weapons or clothing, with the same scrupulous level of attention to their botanical identity. Most of Spruce’s ethnobotanical specimens were collected in the Rio Negro region of the northwest Amazon, along the Rio Uaupés, probably into Colombia. These Colombian specimens include poisoned arrows with shafts made from grass (EBC 38648, Gynerium sagittatum) and a palm leaf (Mauritiella armata, EBC 35883). He also collected items on the border between Venezuela and Colombia. THE TWENTIETH CENTURY: COLOMBIA’S NATIONAL AGENDA A B FIGURE 3. Wood specimens. A Aniba perutilis Hemsl., trade names ‘Comino liso, ‘Plain Comino’. Sent to Kew in 1893 by William Gordon, British Vice-Consul, Medellin (EBC 14468). B Caraipa llanorum Cuatrec. Sent to Kew in 1976 by the National University of Colombia (EBC 1130). 108 One of the most interesting themes drawn from the EBC is the transformation of British global science: from the commercial focus of the late colonial period to a postcolonial scientific interest in biodiversity. Botanists, diplomats, and experts would send items to Kew that reflected their interests and what they believed was the interest of Kew as an imperial scientific institute. From the late twentieth century onwards, these sources dry up, and donations are made almost exclusively by professional botanists operating within the Convention on Biological Diversity parameters, working in a collaborative and conservation-based framework for science. The work of Morley Thomas Dawe, who was a consultant for the Colombian Ministry of Agriculture, is emblematic of the shift from individual or commercial collectors to the state agency, albeit still working within a framework of bioprospecting. Dawe travelled the country between 1915 and 1920, drawing on his expertise as a trained botanist and long-time collaborator with Kew and exploring Colombia’s botanical potential (Figures 2c and 4). Though Colombia had a policy of highlighting its botanical potential on the global stage since its independence (del Castillo, 2010), it relied mostly on foreign experts. Due to his pre-existing relationship with Kew, Dawe sent over 70 items (representing a quarter of the entire Colombian EBC collection). His expeditions were mostly in the northern and western regions of Colombia, traversing the different ecosystems of the Caribbean, Andean, and Pacific regions. He later published lengthy reports (such as Dawe 1918, 1921a, 1921b). This scenario highlights the mediating role he had between the UK and Colombia. As with the fique collections, these studies were fundamentally economic, looking for commercial and agricultural alternatives to benefit Colombia and the UK. The lack of Colombian specimens (except for timber) during the rest of the twentieth century reflects both a wider decline in acquisitions during this time, as the development of the Museum of Economic Botany ceased to be a priority for Kew, and a parallel lack of engagement of Kew botanists with Colombia. Colombia entered the commercial sphere of the United States, and American botanists played a major role in the exploration of the Colombian flora. Typical of the sporadic acquisitions of Colombian items during this period was the basketry purchased by Kew botanist Lauren Gardiner at the World Palm Symposium, held in 2015 in the municipality of Montenegro, Quindío CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 7 FIGURE 4. Model with figures and utensils made from corky outgrowths of the bark of ‘hobor’ tree, Spondias mombin. Sent by Morley Thomas Dawe in 1916 (EBC 37950). Department (Figure 5). The Symposium included an organised market for attendees to buy traditional crafts and other palm products. This reflected the shift during the 20th century from an emphasis on exporting raw materials to benefit large corporations or manufacturing industries overseas, to projects that emphasise high-value products, the retention of income in local communities in Colombia, and the sustainable use of biological resources. THE TWENTY-FIRST CENTURY: THE COLOMBIA BIO PROJECT In the 21st century, Kew has developed a new relationship with Colombian researchers through collaboration with the national Colombia Bio project. This Colombian government initiative is based on the sustainable use of local biodiversity. Kew has collaborated with Colombia Bio in several projects, of which the ‘Biodiversity Expeditions’ in Boyacá provided several items for the collection. Although there are many similarities between these expeditions and those conducted by Dawe one hundred years earlier, Colombia Bio is based in a well-established community of botanical researchers in Colombia. The items collected during the Colombia Bio expeditions, such as the sample of the basketry in the Boyacá area (EBC 100447-100449), demonstrate a different vision and perspective: one of self-reflection and identification of the country’s own ecological and cultural heritage. CONCLUSIONS Our initial study of the Colombian component of the EBC – and there is much more to be done – demonstrates the importance of such case studies of countries that (during our study period) did not form part of one of the European empires. We emphasise the complex routes by which specimens reached Kew, with the strong involvement of commercial interests and actors within Colombian society in the nineteenth century. While economic botany in the nineteenth century clearly continues to function as a tool for extracting wealth from South America, even after independence, this is a complex process in which Kew’s engagement was fitful. From the early twentieth century onwards, the main actors are the Colombian government and, later, Colombian scientists. In both the early twentieth and twenty-first centuries, the emphasis shifts towards supporting local economic development. Now, we can see an explicit focus on conservation and ethnobotanical methodologies as the bridge between biodiversity conservation and its use. The Economic Botany Collection is thus an archive both of traditional knowledge (for detailed case studies, see Echeverri-Sanchez et al., 2021) and an archive of Colombia and Kew’s changing engagement with the world. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 109 CHAPTER 7 FIGURE 5. Bag woven by Reinaldo da Silva, of the Tikuna people of the Colombian Amazon, with fibres from the unexpanded leaves of the chambira palm, Astrocaryum chambira Burret. This palm, the most important source of fibre in western Amazonia, was in process of domestication by the time of the European conquest (Garcia et al. 2015). Bought by Lauren Gardiner at the World Palm Symposium, Montenegro, Quindío, Colombia, 2015 (EBC 99453). 110 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 7 Acknowledgements We thank Frances Cook for curatorial assistance in the EBC, Ellie Graves and Federico Fabriani for advice on plant identification, and Carmen Amalia Díaz Peña and William Milliken for helpful comments on the manuscript. References BGCI (2021) GlobalTree Portal. Available at: https://www.bgci.org/ resources/bgci-databases/global-tree-assessment-portal/ Bushnell D (1993) The making of modern Colombia: a nation in spite of itself. Berkeley, University of California Press. 335p. Cain PJ, Hopkins AG (2016) British imperialism: 1688-2015. 3rd Edition. London, Routledge. 794p. Cendales Paredes C (2012) Robert Thomson (1840-1908): a Kew gardener in Bogotá, Colombia. J Gard Hist, 40: 239-52. Cornish C (2012) Useful and curious: a totem pole at Kew’s timber museum. J Mus Ethnogr, 25:138-151. 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CATALOGUE OF USEFUL PLANTS OF COLOMBIA 111 CHAPTER 8 CENIVAM Garden. 112 Elena Stashenko CATALOGUE OF USEFUL PLANTS OF COLOMBIA Chapter 8 Bioeconomy: a sustainable and responsible option for making the most of Colombian plant resources Elizabeth Hodson de Jaramillo1* & Elena Stashenko2* 1 2 Professor Emeritus, Pontificia Universidad Javeriana, Bogotá, Colombia. ehodson8@outlook.com Laureate Professor, General Director CENIVAM – Industrial University of Santander, Colombia. elenastashenko@gmail.com *Corresponding authors: ehodson8@outlook.com, elenastashenko@gmail.com Keywords: circular bioeconomy, biodiversity, sustainability, SDGs, value-added bioproducts ABSTRACT Environmental deterioration, degradation of natural resources, and climate change are the greatest current challenges our planet faces. The environmental and social imperative of meeting the Sustainable Development Goals in a more environmentally friendly economy is not an option, but it is humanity’s greatest responsibility today. The most obvious and clever response to this challenge is through the circular bioeconomy model for the sustainable production and use of resources and their transformation into value-added bioproducts. The bioeconomy strategies focus on value chains with integrated cycles, which increase efficiency and recycling through products and co-products in different biological systems. For a megadiverse country such as Colombia, sustainable bioeconomy requires a completely different way of producing: circular instead of linear, and using science intensively to add value to biological resources and biological processes. Basic concepts of circular bioeconomy, value chains and biotechnology are given, and some examples of bioeconomy-focused biotechnological developments in the Research Centre for Agro-industrialisation of Tropical Aromatic and Medicinal Plant Species CENIVAM are presented. RESUMEN El deterioro ambiental, la degradación de recursos naturales y el cambio climático son el mayor reto actual. El imperativo ambiental y social de atender los ODS en una economía más respetuosa con el ambiente no es una opción, es la mayor responsabilidad actual de la humanidad. Probablemente la respuesta más obvia e inteligente a este reto es a través del modelo de bioeconomía circular para la producción y aprovechamiento sostenible de los recursos y su transformación en bioproductos con valor agregado. Las estrategias de la bioeconomía se basan en cadenas de valor con ciclos secuenciales integrados, lo cual incrementa la eficiencia y reciclaje a través de productos y co-productos en diferentes sistemas biológicos. Para un país megadiverso como Colombia, la bioeconomía sostenible requiere una forma totalmente diferente de producir: sistemas circulares en vez de lineales y el uso intensivo de conocimiento para dar valor agregado a los recursos y procesos biológicos. Se presentan conceptos básicos de bioeconomía circular, cadenas de valor y biotecnología y se plantean algunos ejemplos de desarrollos biotecnológicos con enfoque de bioeconomía obtenidos en el Centro de Investigación para la Agroindustrialización de Especies Aromáticas y Medicinales Tropicales CENIVAM. INTRODUCTION The world is facing systemic climate, biodiversity, social, economic and health crises (IACGB, 2020). Biodiversity loss and land use are drivers correlated with zoonotic diseases such as COVID-19 and other emerging infectious diseases (Dasgupta, 2021). Environmental deterioration, degradation of natural resources, and climate change are the utmost current challenge. The global economic system must undergo radical transformation processes to improve sustainability (Lewandowski, 2018). The greatest endowment humankind has, and depends on, is nature that provides food, water, shelter, recreation, and resources to improve health and well-being, while maintaining nutrient cycles, soil, and water health, regulating climate, and increasing resilience. Inadequate human management has led to severe negative impacts, degradation, and decline of natural resources, challenging nature’s productivity, resilience, and adaptability, and threatening global economies. Ecosystem health and integrity have been affected by the extraction of natural resources, including the waste generated from this process (Dasgupta, 2021). The wise use of Science and Technology (S&T) to face social and environmental threats is an ethical and social imperative. Scientific innovations play a crucial role in maintaining healthy soils and water, and in protecting or restoring natural habitats, conserving biodiversity, and increasing agrobiodiversity in local communities (von Braun et al., 2021). The fulfilment of the Sustainable Development Goals (SDGs) in an economy that is more respectful of the environment is not an option, but it is the greatest CATALOGUE OF USEFUL PLANTS OF COLOMBIA 113 CHAPTER 8 current responsibility of humanity. It is fundamental to promote human, environmental, social, and economic health. Probably the clearest and smartest approach to this challenge is through a circular bioeconomy model. Circular bioeconomies aim to use resources sustainably by mitigating waste streams through their transformation into side-line products that have added value such as food, feed, biobased products, biomaterials, and bioenergy. The goal is to produce more with less through waste recovery and to make use of all available S&T developments to harness the benefits of biodiversity by promoting resilient value chains (Misión Internacional de Sabios, 2021). Transformation to a sustainable system requires changes in food production and processing (the major driver of terrestrial biodiversity decline), changes in decarbonising our energy systems (the principal contributor to anthropogenic climate change), and drastically altering patterns and behaviours in production and consumption to better align with both environmental objectives and the entire global supply chain network (Dasgupta, 2021). The concept of bioeconomy as an alternative path to sustainable development rests in its potential for renovation. A circular bioeconomy has a major role in re-aligning growth with the biosphere (IACGB, 2020). As stated by the European Commission in their updated Bioeconomy Strategy (European Commission, 2018), “We live in a world of limited resources […]. We must improve and innovate the way we produce and consume food, products, and materials within healthy ecosystems through a sustainable bioeconomy.” The bioeconomy includes, in addition to the traditional agriculture, forestry, fisheries and aquaculture sectors, several downstream processing and service industries, such as food, paper, textiles, building and construction, chemistry, cosmetics, and biopharma. Key converging technologies, such as bio-, nano- and information technologies, are crucial to the knowledge-based bioeconomy, which applies biobased processes and principles in industrial applications for the obtention of value-added bioproducts (IACGB, 2020). The bioeconomy embraces principles of renewability and circularity and provides an opportunity to create a system based on sustainable economic growth, using science to add value to biological resources and biological processes while reducing resource consumption and protecting and regenerating ecosystem (IACGB-GBS 2020). The objective of a cascading and circular economy is to increase resource efficiency, reduce demand for fresh materials, and add knowledge and technology to transform and create new industrial value, create jobs, and improve the welfare of the local community (Trigo et al., 2021). BIOECONOMY IN COLOMBIA – WHY? Colombia’s immense biodiversity constitutes the most exceptional asset. Worldwide, it ranks first in orchids, second in general plants, and third in palms. The conservation and sustainable use of its resources represent its development driver. The main proposal of the International Experts Mission 2019 (Misión Internacional de Sabios, 2021) is 114 that the country’s conceptual and political framework for the country’s sustainable development is the bioeconomy, with a territorial approach within the framework of the SDGs. The Ministry of Science, Technology, and Innovation launched the Colombia Bioeconomy Mission (December 2020) and opened a call for proposals in bioeconomy for the development of new technologies and products based on the use of biomass, biodiversity, and ecosystem services, and additionally to encourage sustainable agriculture in regions (https://minciencias.gov.co/convocatorias/innovacion-yproductividad/convocatoria-para-el-apoyo-programas-yproyectos-idi-que). One of the goals is to consolidate science and innovation as the axis of growth in the generation of progress and social equity. This goal must beadjusted to the characteristics, resources, conditions and capacities of each region in the country so that the adoption of the circular bioeconomy model promotes job creation, income growth, and the wellbeing of the entire community. The bioeconomy path chosen by each region must be defined in participatory bottom-up processes to adjust to local conditions and requirements and to contribute to common prosperity. Therefore, the main objective of this chapter is to present a review of the fundamentals of the circular bioeconomy, to promote the sustainable management of biodiversity in Colombia and to present, as a case study in agribusiness, some of the advances that have been obtained in the Centre for Agroindustrialisation of Aromatic Plant Species and Tropical Medicines CENIVAM of the Industrial University of Santander. THE CIRCULAR BIOECONOMY – WHAT IS IT? The circular bioeconomy is considered the efficient management of biobased renewable resources to sustainably produce, technologically process and trade resources from land, fisheries and aquaculture, and to transform them into bioproducts while promoting new industries and jobs (Tan & Lamers, 2021). The circular bioeconomy model for territorial efficient conversion of resources into value added products is considered the best choice for achieving SDGs. This model directly covers at least 14 of the 17 SDGs, making important contributions to the health and nutrition of a growing population, to the sustainable provision of food, energy, water and raw materials, and to soil protection, climate, and environmental protection (IACGB-GBS 2020). This concept excludes the false dichotomy between productive development and sustainability. According to FAO (2021), sustainable and circular bioeconomic development implies balancing different SDGs because strategies are cross-cutting and multi-sectoral, and consequently face greater challenges than sustainable development strategies that target just a single sector. As stated by FAO (2021): “The bioeconomy sought unlocks the potential of nature to provide bioresources, bioprocesses and biodiversity and respects the planetary boundaries”. The circular bioeconomy has also been proposed as the new strategy for reactivation and reindustrialisation (IACGB, 2020), strengthening the eco-efficiency of processes and the use of recycled carbon to reduce current dependencies on CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 8 fossil carbon. It is transversal, regional, and interdisciplinary (Misión Internacional de Sabios, 2021). The circular bioeconomy includes all kinds of biomaterial streams with innumerable application paths, and reconciles local and global integration by promoting the creation of diversified, innovative, and decentralised value-chains and adapting old ones (IACGB-GBS 2020). This model is a disruptive process that needs new knowledge-based processes for innovation towards the eco-efficient circular sustainable management of renewable resources. The goal is to reposition agriculture and the management of rural areas as a part of a balanced development process that generates employment by giving added value to the primary uses of resources and waste byproducts. The bioeconomy is the opportunity for territorial development and a complement to existing productive activities, diversifying and integrating them into value chains and networks that look forward to make responsible use of natural resources and locally generated waste in parallel, to maintain an appropriate balance with nature. In this system, the value of products, materials and resources is maintained in the economy for as long as possible, and waste generation is reduced to a minimum (Misión Internacional de Sabios, 2021). It not only adopts circularity by cascading materials and co-products into several uses in different value streams, but also increases the efficiency in each step of the value chain and minimises the residues generated (Tan & Lamers, 2021). SUSTAINABLE, RESILIENT VALUE CHAINS – THE EFFICIENT MODEL Bioeconomy is not inherently sustainable because it is based on natural resources; it requires the articulated integration of a sustainable supply chain, sustainable, appropriate transformation processes, and sustainable high-value bioproducts, biomaterials or bioenergy with appropriate trade systems (Tan & Lamers, 2021). The circular bioeconomy focuses on the efficient and sustainable valuation of biomass in comprehensive multi-product production chains, using waste and optimising the value of biomass in a cascade (Trigo et al., 2021). We can look to future scientific and technological developments to increase the portfolio of bioproducts developed from the local biodiversity, keeping with a circular economy approach (Hodson et al., 2021). The greatest potential of the circular bioeconomy lies in its wide diversity and ability to adjust to territorial conditions and resources, promoting unique developments that make it possible to establish niches in global markets and to promote local and regional initiatives that create synergies among the stakeholders (IACGB-GBS, 2020). Nevertheless, to promote the advances and bioscience innovations and to direct them to being a factor of sustainability is not enough. It is fundamental to adapt them to local conditions, resources, and capacities, to make them available and accessible to the regional community, especially smallholders, and to use them to enhance local and traditional knowledge (von Braun et al., 2021). This approach reconciles local and global integration because it needs to define policies to strengthen clever international and local value chains, which requires a wise balance of global and local objectives. There is a need to maintain permanent investments in bioeconomy development to establish new, innovative, diversified, and decentralised value-chains and adjust old ones to ensure the sustainability, resilience, and adaptability of the selected supply chains. The bioeconomy maintenance lies in its diversity, adaptability, and close interactions with local and rural communities (IACGB-GBS, 2020). In terms of sustainability and resilience, using a greater range of plant diversity is critically important. Numerous plant species have not been domesticated and show successful traits for cultivation, such as adaptation to stresses and high nutritional content. They offer new possibilities to improve food and nutritional safety and as raw materials for innumerable processes and bioproducts. They contain significant genetic diversity to increase agrobiodiversity and contribute to resilience and more sustainable agriculture to in the face of climate change (von Wettberg et al., 2020). The value chain is a business efficiency model that makes it possible to find comparative and competitive advantages based on the particular activities of each system. For the bioeconomy approach, these value chains include natural ingredients used for several industries: food, feed, cosmetics, nutritional supplements, tropical flowers and foliage, plant extracts, sustainable agricultural products, and ecotourism, among many others. The prioritised value chains contribute in different ways to the conservation of biodiversity and territorial development. BIOTECHNOLOGY – A KEY TO CREATING BIOECONOMIC VALUE The bioeconomic model is the convergence of science towards biotechnology, focusing intently on the management of natural resources for the industrial use, transformation, and manipulation of biomass. It uses the present latent value of this biomass to achieve new growth and social benefits in thefuture (renewable fuels, bioplastics, agricultural inputs, and medicinal products). Biotechnology is the technological instrument of the bioeconomy. Innovation is essential when modifying inefficient, polluting and resource-based industrial processes to improve efficiency and respect for the environment (Misión Internacional de Sabios 2021). Biotechnology supports the development of bio-based industries to facilitate the sustainability of the sector and to reduce environmental impacts. A sustainable bioeconomy requires sustainable supply chains and appropriate transforming biotechnologies for processing renewable resources into high-value-added bioproducts, materials and bioenergy (Tan & Lamers, 2021). Biotechnology is one of the keys enabling the circular bioeconomy to function, offering solutions for efficient and sustainable production of plant and microbial biomass for bioenergy production; for bioremediation and environmental protection; for transformation through green chemistry; and as an integral part of agri-food processes in value chains CATALOGUE OF USEFUL PLANTS OF COLOMBIA 115 CHAPTER 8 among other processes (Trigo et al., 2021). In addition, biotechnology also brings solutions to improve post-harvest processing and thus optimise the bioeconomy sector, as a whole. For instance, in the forestry sector, biotechnology offers solutions to reduce the lignin content of trees or crops, thereby reducing the highly polluting processing of the wood in the paper or biofuel industry (Lewandowski, 2018). Biotechnology plays a central role in these value chains and technological transformation processes. It plays a vital role in numerous biochemical processes such as the fermentation of residues for animal feed or bioenergy, production of biotechnological fertilisers or soil amendments, microbial acidification in silage, enzymatic saccharification of starch and lignocellulose, obtention of food and feed additives, and production of supplements, vitamins, amino acids, biopolymers, cosmetics, and pharmaceutical bioproducts. Engineering the composting and residual biomass transformation by microorganisms is one example of the crucial value added by applying biotechnology to the ongoing issue of residue management (Kircher, 2021). Biotechnology applies science to disrupt business-as-usual processes by focusing on value added through renewability, circularity, and cascaded resource use. This ‘biologisation’ of the economy comprises social and high-tech innovations and the development of completely new solutions, services, and products that combine sustainability with increased consumer benefit and thus enable new lifestyle concepts (IACGB-GBS, 2020). Every present-day economic system considers sustainability essential. It mimics biological systems in what nature has done for millions of years: being dynamically balanced and sustainable. The central core of this model is the integration of all the steps of the processes in value chains or networks and not only processes of valueadding at each step. These chains must take advantage of cascade technologies to optimise processes across multiple products and must generate circularity (Misión Internacional de Sabios, 2021). AGROINDUSTRY IN COLOMBIA Colombian agriculture faces a series of very sensitive challenges to its future productivity, fundamentally related to the dramatic decline of the rural population (55% in 1955 and currently barely reaching 24%), its progressive ageing and the lack of labour, accompanied by few incentives for young people to stay in rural areas and associate their future with the countryside. Agro-industry in Colombia focuses on a basket of staple and cash crops; including many grains (rice, sorghum, corn, wheat, barley, quinoa, and others), oils and oilseeds (palm oil, coconut oil, cashew, soy), crops for beverages (coffee, cocoa, tea), fresh vegetables (potatoes, onions, bananas, yams, cassava, beans, peas, carrots, etc.), wood products, rubber and fibres (fique fibre plant, cotton), fresh fruits (pineapple, avocado, blackberry, mango, lulo, passion fruit, guava, banana, citrus, etc.), fresh aromatic spices (bay leaf, turmeric, ginger, basil, thyme, rosemary, etc.), and cut flowers (rose, carnation, and others). Although not all the land in Colombia is arable, the 116 current area occupied by these crops is less than 9%, and large areas remain that could support the introduction of promising new crops. While banana cultivation occupies 915,987 ha (DANE, 2014), that of aromatic plants is only 65,846 ha (0.78% of the total cultivated area). Agricultural agro-industrial projects in large areas can cultivate aromatic plants to obtain essential oils (EOs). Still, considering the topography and agricultural infrastructure of the country, production from smallholdings – through associations or cooperatives of growers and their collective work as small farm owners – is paramount to achieving socio-economic and environmental sustainability. This could prevent the displacement of growers and consequently preserve family farms. In many cases, men must abandon their families and move to other country regions as day labourers (for example, to collect coffee). However, by having new productive projects on their farms, they could obtain additional income, which will allow families to remain together. THE EXAMPLE OF CENIVAM The scientific research carried out by the Centre for Research on Tropical Aromatic and Medicinal Plant Species (CENIVAM) based at the Industrial University of Santander, UIS (Bucaramanga, Colombia) is focused on the transfer of scientific and technological knowledge to the field through the development of an Agro-industry of EOs and natural ingredients from aromatic and medicinal plants cultivated in small farms, by growers united in associations. The development route that needs to be covered is still quite long, as illustrated by the import/export activities of Colombia in the EO market, which represented in 2019 a mere 0.0058% of the US$ 5,6 billion-dollar global market (International Trade Center, 2021). CENIVAM is a multidisciplinary research network of groups from several Colombian public and private universities that joined efforts to study Colombia’s biodiversity, focusing on aromatic and medicinal plants, their EOs and extracts. The approach performs research activities under three major investigation blocks: bioprospection, biotransformation, and bioactivity. The long-term expected results are developing both natural ingredients and final commercial products that use their biological properties. Under a permit from Colombia’s Ministry of Environment and Sustainable Development, more than 25 botanical outings and expeditions were organised to obtain plant material from different regions in the country. Verbenaceae (350 accessions), Asteraceae (241 accessions), Labiatae (235 accessions), Piperaceae (139 accessions), and Melastomataceae (27 accessions) are the main plant families studied, thus far yielding over a thousand EOs and extracts for further chemical and biochemical characterisation. During the past fifteen years, CENIVAM has conducted multiple studies on native and introduced aromatic and medicinal plants, their secondary metabolites, EOs and extracts, their biological activities, and their applications in final products. For the instrumental analysis of volatile CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 8 FIGURE 1. Lippia origanoides and the Chicamocha river. Elena Stashenko fractions, EOs, and extracts, gas chromatography (GC) is used in one-dimensional or two-dimensional versions (GCxGC), in capillary columns of different polarities, using universal, selective, or very specific detection systems (Choi, 2003). High-resolution mass spectrometry detection systems (HR-TOF, Orbitrap) coupled to gas or liquid chromatographs are nowadays an excellent alternative for exact mass determination (elemental composition analysis), and for the selective and very sensitive detection of secondary metabolites in complex floral scent mixtures (Stashenko & Martínez, 2007, 2008), plant volatiles, EOs, and extracts obtained with organic solvents or supercritical fluids (CO2). This research network has carried out more than 5,000 biological tests during 2005-2020 on EOs and extracts obtained from more than a thousand botanical accessions collected in the field in different regions of Colombia. Around 45-47% of EOs or extracts possessed some interesting biological activity. Since the cytotoxicity of EOs and extracts may prevent their use in some pharmaceutical applications, this aspect is normally included among the first assays of the bioactivity investigation. Acute toxicity tests using brine shrimp with Artemia franciscana for a large set of EOs studied in CENIVAM showed that over 30% of these oils did not have any degree of toxicity (Olivero et al., 2009). The virucidal activity of some essential oils against dengue (Meneses et al., 2009a,b; Ocazionez et al., 2010) and yellow fever (Meneses et al., 2009a,b) was studied at the Research Center for Tropical Illnesses, CINTROP. Of the EOs tested, 83% were active against the dengue virus, and 66% showed positive results in inactivating the yellow fever virus. These results are a very interesting contribution, especially considering that there are not many data in the literature on the EO activity against yellow fever and related viruses. Chagas and leishmania are additional tropical diseases that deserve attention. Bioactivity assays against CATALOGUE OF USEFUL PLANTS OF COLOMBIA 117 CHAPTER 8 Trypanosoma cruzi (epimastigotes and amastigotes) and Leishmania chagasi (promastigotes and amastigotes) were carried out at CINTROP using cell lineages commonly employed for these purposes (Vero and THP cells). From the results, 48% of the EOs examined were active against T. cruzi. The EOs were also tested against L. chagasi (promastigotes and amastigotes) and 19% were found to be active (Escobar et al., 2010). Antifungal (Mesa-Arango et al., 2010) and antibacterial (Caceres et al., 2020) activities have also been extensively studied. An important consideration to keep in mind during the study of secondary metabolites from aromatic and medicinal plants is the existence of plant chemotypes when plants with identical morphologies have different secondary metabolic profiles. Notable examples are Lippia alba and L. origanoides, two Verbenaceae species that have attracted lots of attention due to their biological properties. Seven chemotypes have been reported for L. alba, three found in Colombia (Lopez et al., 2011; Mesa-Arango et al., 2010). Up to five different chemotypes have been distinguished for L. origanoides, three of them present in Colombia (Stashenko et al., 2010; Sarrazin et al., 2015). Only two of the L. origanoides chemotypes are rich in thymol and carvacrol. Their high content confers antimicrobial properties to their EOs to make them important ingredients for a successful commercial chicken feed product. Mountain oregano, the common name used for L. origanoides, is a species that constitutes a very good example of a tropical plant with many potential applications in the pharmaceutical and food industries. It is a very good source of flavonoids (Stashenko et al., 2013; Arias et al., 2020), which may explain its activity against breast cancer (Raman et al., 2017, 2018), as well as the efficacy of its EO as an insect repellent that may be used to protect grains from beetle attack during storage (Alcala-Orozco et al., 2019; Caballero-Gallardo et al., 2012). The antigenotoxicity of L. origanoides EOs (Vicuña et al., 2010) makes them important ingredients of photoprotective cosmetics, which may as well take advantage of their antioxidant properties (Stashenko et al., 2008). An example of circular economy application is the patent for a methodology that makes integral use of L. origanoides by producing its EO, fractions enriched in phenylpropanoids, extract, and hydrosol, in a combined process (Stashenko et al., 2019). The post-distillation biomass is then used as biofuel and composting material, while the hydrosol can be used in pest control. Various pilot research projects in the field have been financed in the past 15 years by Colciencias (currently Minciencias, Ministry of Science, Technology, and Innovation), the Ministry of Agriculture and Rural Development, the Governments of Santander and Arauca, and the Fund of the General System of Royalties (SGR, UIS - Government of Santander). More than 200 rural families have been involved in aromatic plant cultivation and their distillation to produce EOs. The combined knowledge of chemical composition and biological activity serves as the basis for the sustainable use of biodiversity in developing new consumer products 118 for the cosmetics, hygiene, food, and pharmaceutical industries. Pilot EO production units have been implemented in some municipalities in Santander, Arauca and Cundinamarca in Colombia, where farmer associations have been trained in good agricultural practices, postharvest treatments, and the operation of stills designed at UIS for EO production using either hydrodistillation or steam distillation. Thanks to these pilot units, farmers have begun producing Cymbopogon nardus, C. martinii, and Lippia origanoides EOs. New developments have started to extend the cultivation and EO production to additional species, such as Cananga odorata (ylang ylang), Pogostemon cablin (patchouli), Vanilla planifolia (vanilla), Lippia alba, Rosmarinus officinalis (rosemary), Thymus vulgaris (thyme), and Pelargonium graveolens (geranium). In addition, many promising aromatic and medicinal plants are under investigation, and the intense study of their biological and chemical profiles should lead to the design of new Colombian products based on natural ingredients. CONCLUSIONS — PROSPECTS AND CHALLENGES TO A COLOMBIAN BIOECONOMY The sustainable development of the bioeconomy requires solid scientific support for all the value chain steps. This was the main justification for presenting the CENIVAM example, which shows contextualised developments of final commercial products for the essential oils agro-industry in which the circular economy principles are implemented. Crossing bioprospecting results with those from analytical chemistry and molecular biology (bioactivity assays) led to the detection of promising vegetal species to be used as ingredients for new consumer products. Field projects have established the conditions for biomass production under good agricultural practices and for their rural processing to afford essential oils, extracts, biocontrol solutions, composting material, and biofuel. The modular agroindustrial projects that currently in place in several Colombian municipalities serve as a reference for developing new enterprises that add value to the crops, diversify the rural economy, and creating career opportunities for farm workers and professionals while using resources comprehensively with low waste generation. Multiple challenges remain from diverse areas, such as establishing the commercial chains with reinvestment into the countryside, the achievement of origin recognition, road improvement, internet coverage, water supplies, and stable legislation. Acknowledgements Economic support from the Colombian General System of Royalties and the Colombian Ministry for Science, Technology, and Innovation is gratefully acknowledged. References Alcala-Orozco M, Caballero-Gallardo K, Stashenko E, Olivero-Verbel J (2019) Repellent and fumigant actions of the essential oils from CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 8 Elettaria cardamomum (L.) 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(Arecaceae)—Infructescence. 120 Juan Carlos Copete CATALOGUE OF USEFUL PLANTS OF COLOMBIA Chapter 9 Sustainable value chains and development pathways for natural ingredients in Colombia: the case of naidí (Euterpe oleracea Mart.) Mabel Tatiana Rojas1*, Tiziana Ulian2*, Carlos Alberto Cortés1, Germán Eduardo Torres-Morales1, David Hammond2, Felipe García1 & Mauricio Diazgranados2 1 2 Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Bogotá, Colombia. Royal Botanic Gardens, Kew. *Corresponding authors: mrojas@humboldt.org.co, t.ulian@kew.org Keywords: biodiversity, bioeconomy, Euterpe, green growth, value-chains, sustainable development ABSTRACT Colombia is the second most biodiverse country globally, potentially having a comparative advantage over less biodiverse countries in shifting to a more sustainable, biobased economy. However, Colombia accounts for only 0.27% of natural ingredients’ exports in Latin America; hence its potential is still locked. As a case study, we applied a value-chain analysis to the fruit of the naidí palm (Euterpe oleracea) growing along Colombia’s Pacific Coast. We conducted semi-structured interviews involving community-based organisations, private companies, NGOs, international organisations, and universities to assess the full range of activities and relationships involved in its value creation. We found that the factors characterising its value chain consist of trust relationships between small-scale producers, a low distribution of profits among local collectors, limited profitability supported by external subsidies, and low regional and national demand. This value chain could be improved in the short- and medium-term by increasing the capacity of cold chains and storage centres, fostering economic incentives for implementing agroforestry systems, developing strategies for promoting traditional diets by using its natural ingredients, and improving technology and business capabilities to enhance its commercialisation. RESUMEN Colombia es el segundo país con más biodiversidad a nivel mundial, y potencialmente tiene una ventaja comparativa sobre los países con menor biodiversidad para cambiar a una economía de base biológica más sostenible. Sin embargo, Colombia representa solo el 0,27% de las exportaciones de ingredientes naturales en América Latina; de ahí que su potencial aún esté bloqueado. Como caso de estudio, aplicamos un análisis de la cadena de valor al fruto de la palma naidí (Euterpe oleracea) que crece a lo largo de la costa del Pacífico de Colombia. Realizamos entrevistas semiestructuradas en las que participaron organizaciones comunitarias, empresas privadas, ONGs, organizaciones internacionales y universidades, para evaluar la gama completa de actividades y relaciones involucradas en su creación de valor. Encontramos que los factores que caracterizan su cadena de valor son: las relaciones de confianza entre los pequeños productores, una baja distribución de utilidades entre los recolectores locales, una rentabilidad limitada sustentada en subsidios externos, y una baja demanda regional y nacional. Esto podría mejorarse en el corto y mediano plazo aumentando la capacidad de las cadenas de frío y los centros de almacenamiento, fomentando incentivos económicos para la implementación de sistemas agroforestales, desarrollando estrategias para promover dietas tradicionales utilizando sus ingredientes naturales, y mejorando la tecnología y las capacidades comerciales mejorando su comercialización. INTRODUCTION Colombia is the second most biodiverse country globally, hosting around 10% of the planet’s biodiversity (Rincón Bermúdez et al., 2009). It has at least 28,947 vascular plants, of which ca. 7,472 have a documented use (Negrao et al., 2022). Of these, 79% have medicinal uses, 32% are harvested for materials (including personal care products and textiles), 28% provide environmental services (soil improvers, ornamentals, plants for erosion, and pollution control), and 26% are sources for human food (Diazgranados et al., 2021). This high diversity of useful plants represents a potential comparative advantage over other countries, as it can be considered an asset in developing Colombia’s bioeconomy. Development of the Colombian bioeconomy and Green Growth Recently, the concept of bioeconomy has received much attention at the international level as a viable system for transforming goods and services to overcome social, economic, techno-scientific, energetic, and environmental sustainability challenges (Staffas et al., 2013; Moñux et al., 2018). In 2018, more than 40 countries had policies or CATALOGUE OF USEFUL PLANTS OF COLOMBIA 121 CHAPTER 9 strategies in place to develop their bioeconomies based on the transformation of goods and services from the use of biological resources (Dietz et al., 2018). These were led by strategic priorities within each country, economic specialisation, biomass allocation, historical investments in research and development and labour productivity (Bracco et al., 2018). For Colombia, bioeconomic development has been identified as a priority to achieve the 17 UN Sustainable Development Goals (SDGs). Similarly, the Organisation for Economic Co-operation and Development (OECD) and the Economic Commission for Latin America and the Caribbean (ECLAC) have encouraged the country to design an agenda to unlock the potential of its biodiversity for sustainable development (OECD, 2009; Albrecht et al., 2015). Currently, the government is implementing a national bioeconomy through a Green Growth strategy (CONPES, 2018), taking advantage of the nation’s large natural capital. After developing the Green Business Strategy, the concept of bioeconomy was promoted by the Green Growth Mission of the National Planning Department (Departamento Nacional de Planeación, DNP) in 2015. This Mission has identified that the development of a national bioeconomy is fundamental for achieving sustainable growth, defining it as “a model that efficiently and sustainably manages biodiversity and biomass to generate new value-added products, processes, and services, based on knowledge and innovation” (DNP, 2018, p. 28). It is expected that by 2030 the bioeconomy will account for 10% of the Colombian Gross Domestic Product (GDP). Around 100,000 new jobs will be created in different subsectors, including biocommerce, natural ingredients, generation of new materials, biorefineries, bioenergy, agri-food systems, and advanced biotechnology (Misión Internacional de Sabios, 2019). Connections between bioeconomy and natural ingredients Growing a bioeconomy in Colombia involves identifying and producing natural ingredients via biotech and non-biotech pathways as part of a process of value-creation and innovation that earmarks the replacement of synthetic compounds with those of natural origin (Biointropic, 2019). Supplying natural ingredients is thus essential in differentiating Colombian products in global value chains (ONUDI, 2015), where the application of sustainable business models is increasingly needed to maintain market access and consumer interest. In this study, we adopted the following definition of natural ingredients: “The solid or liquid raw material extracted from the physical, and sometimes simple chemical processing of animal sources, plants and other living, native organisms, respecting the principles of Biotrade, used in the development of products for therapeutic purposes, of hygiene or beauty of the body and of products that through its ingestion nourish the organism” (Fondo Biocomercio, 2009, p. 14). These are classified into nine groups by the Fondo Biocomercio (2009): 1) tannins or dyestuffs; 2) active ingredients for therapeutic purposes; 3) condiments-spices and value added-fruits, sweeteners, thickeners, and flavourings; 4) essential oils; 5) fats and waxes; 6) natural flavouring substances; 7) saps, 122 gums, resins, and oleoresins; 8) juice, pulps, extracts, juice concentrates; and 9) flours and starches. Status of natural ingredients in Colombia According to Rojas et al. (2021), Colombia exported a total of 650 tons of natural ingredients worth 3.4 million dollars in 2019. The group with the largest share of the country’s exports of natural ingredients (49%) was juices and plant extracts. However, the group with the highest level of business dynamism between 2014-2019 was natural colourings, with a Compound Annual Growth Rate (CAGR) of 11.95%. In parallel, the country imported 5,552 tons of natural ingredients worth 60 million dollars (Rojas et al., 2021). This scenario means that the country has a trade deficit of 57.1 million dollars, which is significantly higher than expected when considering the rich country’s biodiversity and potential for producing natural ingredients, reflecting the lack of development in the natural ingredients sector. Colombia accounts for only 0.27% of Latin America’s exports of natural ingredients (Rojas et al., 2021). Hence, its potential remains unfulfilled in helping to meet the Colombian government’s commitment to Green Growth. Papellet et al. (2020) argue that this is due to a lack of knowledge of technical product standards and regulations, and the species management requirements needed to satisfy consumer demand. Partly this is due to a lack of information regarding the biology and ecology of species of interest and to limited research in biotechnology, such as bioprospecting (Centro Red de Innovación, 2013; Biointropic, 2019). This gap is reflected in the small number of bio-innovative companies. As of 2018, the country had only 90 identified green businesses and 84 registered bioproducts (DNP, 2018), with even fewer established value chains in the country involving local communities. Achiote (Bixa orellana), naidí (Euterpe oleracea), Jagua (Genipa americana), and sacha inchi (Plukenetia volubilis) are exemplary exceptions (Biointropic, 2019). Studies suggest an absence of institutional leadership, the inefficient use of natural capital, low investment in research and development, and regulatory barriers as the main causes hindering the development of bioeconomic initiatives in the country (DNP, 2018; Moñux et al., 2018). Administrative inefficiencies in obtaining sustainable harvest permits for non-timber forest products and contracts to access genetic resources for commercial purposes may also play a particularly important role (Biointropic, 2019; Papell et al., 2020). In this context, it is essential that biodiversity becomes a differentiating factor in Colombia’s bioeconomy to overcome socio-economic challenges at the regional and national levels. This chapter aims to assess the value chains of natural ingredients in Colombia and to explore the potential for developing Colombia’s bioeconomy and supporting sustainable livelihoods. The objective is to provide a framework for developing sustainable value chains of natural ingredients and to create pathways for sustainable development in Colombia. To do this, we applied a value CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 9 chain analysis to the fruit of the naidí palm (Euterpe oleracea). We examined limitations, opportunities, and success factors weighing on sustainable value chain development for natural ingredients in Colombia. This work has been carried out in the framework of the ‘Useful Plants and Fungi of Colombia’ project, led by the Royal Botanic Gardens, Kew, in partnership with the Alexander von Humboldt Biological Resources Research Institute (see Chapter 1). the chains. Efficiency will transcend production processes and entail information flows among the actors in the chain and the ability of support services and enabling conditions to overcome challenges (Lundy et al., 2007). Within this broader context, paying mind to internal factors, such as adaptive schemes of governance and social, human and natural capital that underpin livelihoods, will increase resilience to exogenous shocks and improve sustainability (Ellis & Allison, 2001). MATERIAL AND METHODS A value chain is understood as a strategic network among several organisations. According to Hobbs et al. (2000), this approach differs from a supply chain in several ways: value chain participants have a long-term strategic vision; recognise interdependence and align working together to solve common problems, sharing risks and benefits; value chains are demand-driven, not supply-driven, and thus focus on consumer needs. Value chains do not exist by themselves but are part of the socio-economic and institutional system, and therefore external forces affect Study area and actors We conducted a value chain analysis of the processed fruit of the naidí palm (Euterpe oleracea) growing along Colombia’s Pacific Coast. The analysis was carried out in the municipalities of Buenaventura (Valle del Cauca), Bajo Baudó (Chocó) and Vigía del Fuerte (Antioquia). We documented the source of natural ingredients (from raw material to the final product), the processes for capturing and creating value, and emerging interactions and relationships of autonomy and interdependence among TABLE 1. Direct actors involved and interviewed for the value chain analysis of naidí palm fruit products. Actor Description Naidí Pacífico SAS Community-based company comprising 7 community councils (CC): two of them (CC Cajambre and CC Bahía Málaga) are in Buenaventura (Valle del Cauca), and the rest (CC Sivirú, CC Piliza, CC San Andrés de Usaragá, CC Guineal and CC Concosta) in the municipality of Bajo Baudó, comprising the Departments of Chocó and Cauca. This company was established in 2015, with sales and production management needing training and creating a legally constituted business model. Planeta CHB SAS A company created in February 2016, with ten partners belonging to the municipalities of Vigía del Fuerte (Antioquia) and Bojayá (Chocó), adjacent to the Atrato River. Of the ten partners, seven communities are represented, and three of these have been part of the organisational process of local Community Councils. Corpocampo A wholesale company focused on selling natural and functional ingredients from Non-timber Forest Products on international markets. The naidí-based products benefit 1,200 families in Guapi (Cauca), Buenaventura (Valle del Cauca), Tumaco (Nariño) and Puerto Asís (Putumayo), who manage 15,600 hectares of forest with sustainable agroforestry models. This company won the Norwegian ‘Business for Peace’ award in 2018, is a member of the ‘Business Call to Action’ initiative led by the United Nations Development Program (UNDP), and was nominated for recognition by them. Fondo Acción Colombian private fund with 20 years of experience in sustainable investments in environmental and childhood projects. MUCHOCOL A retail distribution company that eliminates intermediaries and reaches consumers directly. Universidad de Los Andes Faculty of Economics University that offers a broad research background on collective property in the Colombian Pacific and on governance schemes in this territory. Partnerships for Forests (P4F) Programme funded in Latin America by the British Government (Department for Business, Energy & Industrial Strategy [BEIS]). This initiative aims to protect the world’s tropical forests through productive and commercial projects that combat deforestation and that, in turn, highlight the value of natural forests. Source: Adapted from Rojas et al. (2021). CATALOGUE OF USEFUL PLANTS OF COLOMBIA 123 CHAPTER 9 FIGURE 1. Mapping the value chain of the natural ingredients of the naidí palm. 124 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 9 stakeholders. A distinction is made between the different direct actors in the chain, those involved in the harvesting, processing, and marketing of products. In parallel, we consider indirect actors, who are not exposed directly to risk attached to the product and may provide ancillary services to chain actors in different links, and regulatory agencies (CIAT, 2014). Then, we explore some of the value chain’s complexity and opportunities, which could help the development and commercialisation of natural ingredients, assessing challenges and opportunities. This approach can be viewed as a tool for identifying constraints, key public policy issues, and assistance interventions (Belcher & Schreckenberg, 2007; Rich et al., 2009). Data collection and analysis We conducted seven semi-structured interviews with direct and indirect actors in the value chain (Table 1; https:// figshare.com/s/07441b0d412923c79d2a). We refer to direct actors as those directly involved in the harvesting, production and commercialisation of goods in the value chain, and conversely, to indirect actors as those who provide support and operational services to direct actors. We explored whether the distribution of the value of the products was equitable along the value chain; that is, whether the costs and benefits of investments are shared across actors in each link. Simple commercialisation margins and consumer price ratios (Marshall et al., 2006) were taken as a reference for measuring benefit distribution and equity within the value chain. First, the calculation of the commercialisation margin was obtained from the price received by each player in the value chain, from harvesters to consumers, using the following equation: Commercialisation Margin = ((PS - PP)/ PC)*100 (Equation 1) where PS is the sale price, PP is the purchase price, and PC is the final retail price. Commercialisation margin depends on the final consumer price, where the numerator of the equation reflects the gross income of each link (sale less purchase price). The commercialisation margin reflects the proportion of the gross income per link and its share on the final price. Alternatively, the proportion of the final price taken by the different actors in the value chain (Marshall et al., 2006) can be calculated from information on the product’s final price (Equation 2). Consumer Price Ratio = (PS / PC)*100 (Equation 2) RESULTS Fruit of the naidí palm is transformed first into frozen pulp and then into the lyophilised powder used in the food and the cosmetic industry (Figure 1). We recorded a diversity of health care products, such as body lotions and shampoos. Furthermore, within the value chain, we assessed the direct stakeholders, their geographical location, the flows of products and their relationships (black arrows), as well as physical and monetary units of exchange and production (boxes with black frames). In parallel, indirect stakeholders, despite not taking direct action within the stages of production, transformation, and commercialisation, act as “enablers” of the entire chain (Figure 1). Finally, we looked at the transport used in each link (Figure 1). Actor assessment Link 1: Fruit harvest In the municipalities of Bajo Baudó (Chocó) and Buenaventura (Valle del Cauca), naidí fruit is wild harvested from a total area of 56,000 hectares. This activity supports the livelihoods of 186 households who harvest the raw material for self-consumption and local sale. Currently, the Community Council of Río Cajambre concentrates most of the production and families (totalling 99), as well as the storage and processing of naidí pulp. The harvest of the fruit is organised by families and the Community Councils that own Naidí Pacífico SAS. This community-based company schedules the dates to receive the fruit, notifying families to take them to the collection centres, paying approximately 1,000 pesos per kilogram. Transport to the interior of the collective territory is 100% by motorboat, owned by the Community Councils and through Naidí Pacifico SAS, the boat circulates in the harvest areas. In parallel, Planeta CHB SAS already has granted harvesting permits in the northern part of Chocó and Urabá antioqueño, harvesting naidí fruit from around 26,000 hectares of ‘naidizales’, benefiting 50 families. Planeta CHB also pays around 1,000 pesos per kilogram of fresh fruit. For this company, wild harvest is not sufficient to meet the demand. Hence this is complemented with the cultivation of the palm in agroforestry systems. On the other hand, Corpocampo buys a kilogram of naidí’s fresh fruit for 1,0001,300 pesos; price depends on the cleanliness, selection, the degree of ripeness of the fruit among other qualities. They also commit harvesters to generate conservation agreements and ‘zero deforestation’; for example, this company requires that a percentage of fruits are left to guarantee food for other species that naturally disperse the seeds of the naidí palm, which could represent about 20% of the forest production. Harvesting is carried out mainly by men since it is timeconsuming being in the ‘naidizales’. The soil is very soft, and harvesters work all day with the water at the waist and abundant insects and moisture. It is also an activity that requires climbing the palms with a safety team (harness), removing the bunches, and lowering them to the floor on a string. On the floor, there is usually an assistant who receives the bunch and takes it to a basket. Naidí is a species that blooms and fruits all year round, with peaks between January-February and May-July, offering a harvest of around 4 tons/peak of naidí pulp (data only available for Buenaventura and Bajo Baudó). CATALOGUE OF USEFUL PLANTS OF COLOMBIA 125 CHAPTER 9 Link 2: Storage and local primary transformation Primary transformation is undertaken in the territory by Naidí Pacífico SAS, which ensures that the naidí harvest can be stored all year round thanks to the installed capacity for harvesting, processing, and freezing. Between January and July 2020, Naidí Pacífico SAS managed to produce 3.6 tons of pulp, mainly from the Community Council of Río Cajambre (Buenaventura). However, in 2020, not even 80% of the harvested fresh fruit produced by this Community Council was transformed into frozen pulp. In parallel, there are two other collection and storage centres in Sivirú (Chocó) and Pizarro (Nariño), with smaller freezers. Soon, with the Partnerships for Forests (P4F) project and Fondo Acción, these two collection centres also plan to start processing plants. Moreover, an industrial scaling phase should soon be underway, with in 10 tons of pulp production forecasted for 2021. Naidí Pacifico SAS sells the kilogram placed in Buenaventura at 12,500 pesos; however, if the client wants the product in other cities of Colombia, an additional value (transport and packaging) is charged. In parallel, there are other local alternatives for processing the naidí fruit. In the Community Council of Río Cajambre, ‘arrechón’ or naidí wine is produced by independent producers. The production and distribution of this product are already legalised, participating in the farmers market of Universidad Santiago de Cali and the music festival of the Pacific ‘Petronio Álvarez’ (Cali, Valle del Cauca). For Planeta CHB SAS, the processing factory of the company is in the urban centre of Vigía del Fuerte (Antioquia). There, the mature fruit of the naidí, the tasks of pulping, packing, and freezing are carried out. In this process, upright freezers are available, which, due to space limitations, have a total freezing capacity of 3.6 tons of pulp. Most of the employed staff are women. Planeta CHB sells a kilogram of frozen pulp to Bogotá and Medellín for 12,000 pesos, and locally, for 10,000 pesos. Likewise, Corpocampo’s processing plants have been built in Buenaventura (Valle del Cauca), Tumaco (Nariño), and La Hormiga (Putumayo) since the Guapi (Cauca) plant had to close due to public order issues. Link 3: Commercialisation Local commercialisation is concentrated in the municipality of Buenaventura (see Table 2), which inherited the traditional use of the species for daily consumption as ‘pepiao’, an exotic traditional recipe that soaks the palm fruit in warm water for manual maceration until a liquid of thick consistency is obtained. Finally, local inhabitants add milk powder, sugar or honey and a pinch of salt. In this part of the country, several independent producers transform the pulp into juices, ‘bolis’ (traditional frozen juices and ice creams), cakes, and wines. At the national level (Table 3), the naidí pulp is commercialised by retailers in Bogotá and Medellín, where it occupies a niche market, characterised by high-end products (e.g., organic food and novel foods). In parallel, wholesalers focus on selling the lyophilised naidí in international markets. TABLE 2. Local-level commercialisation circuits. Community-based companies Naidí Pacífico SAS Planeta CHB SAS The product (pulp) is transported by motorised boat from Cajambre to Buenaventura (the distribution centre), where the company’s main demand comes from hotels, restaurants, and cafes. This distributor type is very important for Naidí Pacifico SAS since 80% of demand is concentrated in this sector. In Medio Atrato (both Vigía del Fuerte and Bojayá), local communities have no reports of historical consumption. However, Planeta CHB SAS sells the frozen pulp to local communities that come to the transformation plant and have a local dispatch system. As in the case of Naidí Pacífico SAS, this has allowed Planeta CHB SAS to recover sales in this time of pandemic. According to Fondo Acción, in 2020, 20–25% of the pulp was consumed in Buenaventura, by people and restaurants. In this municipality, transport is easier as it is distributed by motorbike, door to door. In Cali, they rely on family or friends to distribute the product since it is only two hours by land. Source: Own elaboration based on Rojas et al. (2021). 126 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 9 TABLE 3. National commercialisation circuits. Community-based companies Naidí Pacífico SAS This company sells approximately 500 kilograms of frozen pulp per month through its sales division, mainly to Bogotá, with several customers, among which are retail distributors such as MUCHOCOL and a restaurant called ‘Masa’. Retailers Wholesalers Planeta CHB SAS MUCHOCOL Corpocampo Since 2017, this company has delivered frozen naidí pulp to Medellín by light aircraft, which courier services distribute. Each aircraft can carry 5,000 plastic bags packed in different sizes (200 g, 500 g, 1 kg and 5 kg). MUCHOCOL buys approximately 100 kilograms per month of frozen pulp from Naidí Pacífico SAS, distributed equally to Medellín and Bogotá. The total is distributed as follows: 60 kg in presentations of 500 g and 40 kg in presentations of 100 g. Moreover, in April 2020, they were willing to expand their market to the city of Cali; the company had everything ready, but the restrictions of the pandemic did not allow it. In the short term, Planeta CHB’s goal is to have a distribution point in Medellin, Bogotá, and Urabá. At this point, they have not been able to have a retail distribution channel in supermarkets and speciality stores because they are still organising all the information required on the labels and the brand. Before the pandemic, MUCHOCOL bought the frozen pulp in Buenaventura and made shipments in cargo planes. However, during 2020, they have sought alternatives to make shipments from Buenaventura, making an agreement with the Magdalena SA fleet to distribute to Bogotá (12 hours journey). This company buys pulp from Naidi Pacifico SAS, which in Bogotá represents a cost of 16,500 pesos per kilogram, which is 73% higher than the purchase price of Corpocampo, and 101% higher than the purchase price at Inzunai. MUCHOCOL adds packaging and logistics costs to the selling price until the producer has an income higher than the minimum wage. In turn, MUCHOCOL sells frozen pulp for 20,444 pesos/kg to members and 30,000/kg to the general public. Corpocampo has specialised retailers in Bogotá, Barranquilla, Cartagena, Cali, Medellín, Bucaramanga and Ibagué. Before the pandemic, they distributed their product line in 100 restaurants; now, they distribute their products directly to consumers through their website and WhatsApp. In addition, it is a wholesale company that distributes to other wholesalers such as Éxito, Jumbo, Olímpica and Makro. In 2014, they began exporting a range of products, concentrating on the European (France), Mexican, Chilean, Japanese and Lebanese markets. In this sense, Corpocampo is one of the few actors in the chain that has issued organic certifications (ECOCERT and USDA Organic). In turn, Corpocampo has the most diverse and value-added product line. They have a line of sorbets, frozen pulp (24,083 pesos/kg), lyophilised powder (190,400 pesos/kg), capsules, jams and jellies, as well as honey from Melipona colonies and related products. Source: Own elaboration based on Rojas et al. (2021). CATALOGUE OF USEFUL PLANTS OF COLOMBIA 127 CHAPTER 9 Link 4: Consumers a. MUCHOCOL: establishes its demand through campaigns of responsible consumption in social networks to strengthen or gain consumers’ loyalty. Within their discourse, they emphasise social justice: “Communities are represented on a plate.” b. P4F: given that consolidated producers such as Naidí Pacifico SAS or Planeta CHB SAS have not yet come to produce under economies of scale, niche customers (specialities) are willing to pay a premium for the social and environmental history of these products. c. Naidí Pacífico SAS claims that consumption trends in large cities are based on the naidí bowl (pulp with a variety of nuts and fresh fruits) and fruit juice. d. Corpocampo affirm the importance of the consumer in the value chains and how these actors can influence defining and encouraging different living methods for the peasant, indigenous and Afro-Colombian communities that harvest naidí fruit. Supporting actors 1. Inputs, machinery, tools and infrastructure a. US Agency for International Development (USAID): through the project ‘Territorios de oportunidad Guapi-Timbiquí’, they offer machinery, tools, and infrastructure. b. P4F: make investments in equipment, and these are concentrated in the Community Council of Río Cajambre (Buenaventura). Emphasis is placed on expanding freezing capacity in Buenaventura (Valle del Cauca) and Pizarro (Nariño). c. Corpocampo: supplies organic fertilisers and plant material (seedlings and seeds). 2. Logistics and operations a. MUCHOCOL: assume shipping costs (packaging and distribution) of 4,000 pesos/kg to distribute frozen naidí pulp from Buenaventura to the main cities of Colombia. a. Corpocampo: guarantee fixed purchase prices and the collection of the naidí fruit ‘door to door’ from harvesters. 3. Finance, technical and research a. MUCHOCOL and Bancolombia: ‘venda y aprenda’ platform to strengthen digital markets, business, and personal finance issues; technological development was an incentive for producers to commercialise their products, generate their marketing and expand to other domestic markets. Training in invoicing and maintenance of production and sales records, as well as in matters of verification of equipment and transformation processes. 128 b. Fondo Acción: this institution has worked since 2016 in the Colombian Pacific under a USAID umbrella project (BIOREDD), developing the entire carbon bond line as an incentive to conserve ‘naidizales’. In this region, Fondo Acción supports not only Naidí Pacifico SAS but mainly the Community Councils within the REDD+ projects, in components of governance and productive reconversion, among others. c. Corpocampo: empowers communities to make better use of the forest. Ensures that when the fruit is harvested, there is no palm heart extraction, and when the peaks of production are low, the palm growth is managed. This company has a field team dedicated to these tasks. d. P4F: funded the exchange of experiences between Naidí Pacífico SAS and Belém do Pará (Brazil), the State with the highest production and consumption of naidí in the world. This organisation also hired these experts to go to the Colombian Pacific with the aim that Naidí Pacifico SAS could optimise the harvesting processes (including technical data sheets), pulp processing, operations logistics, investments in equipment, and human capital. In parallel, P4F outsourced a consultancy to design a participatory marketing strategy, business plan, and training in social media management. e. AMPLO: a consultancy company hired by P4F that develops projects and attracts capital to fund them, as inputs to the business plan for naidí pulp. Supports and determines the selling prices of the pulp. f. Chamber of commerce of Buenaventura: according to Fondo Acción, this actor has been training leaders and the community in general to formulate business plans. g. Planeta CHB SAS: trains communities in protocols for the sustainable harvest and management of wild species. They established an agreement with the National Learning Service (SENA) in Apartadó (Antioquia) to carry out forestry training, emphasising good harvesting and processing practices. Two hundred twenty people from 34 communities have been trained. h. USAID: business plan development for naidí in 2016. However, interviews with actors in the value chain state that the business plan, specifically for the lyophilisate of the pulp made by Naidi Pacifico SAS, was never carried out. i. Selvacéutica: a technology-based company that produces NTFPs from the Chocó Biogeographic region for the natural cosmetics and phytotherapy subsectors. It has a line of hair products (nutrition and growth treatment) made from naidí fruit. The company has a community innovation component that contributes to developing competitive biodiversitybased products through the implementation of R&D and knowledge management strategies. CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 9 TABLE 4. Quantitative value chain analysis seeks to analyse the distribution of (monetary) value in each link of the value chain for natural ingredients of the naidí palm, in this case, the frozen pulp. Value chain link Harvest and/or gathering Storage and processing (Naidí Pacífico SAS) Retail (MUCHOCOL) Price (COP $) / kg Purchase 0 Sales 1,000 Purchase 1,000 Sales 12,500 Purchase 16,500 Sales 30,000 Gross income (COP $) Consumer price ratio (%) PULP Commercialisation margins (%) PULP 1,000 3.3% 3.3% 11,500 41.7% 38.3% 13,500 55.0% 45.0% Source: Rojas et al. (2021). 4. Regulatory a. Corpoamazonía: Corpocampo states that this institution has enabled processes in Putumayo, as has Vision Amazonia (an initiative from the Ministry of Environment), to facilitate the acquisition of permits for the sustainable harvest and use of Non-Timber Forest Products (NTFP). b. Corporación autónoma del Valle del Cauca (CVC): technical and logistical support to Naidí Pacífico SAS in obtaining permits to sustainably use and harvest NTFP for commercial purposes. c. CODECHOCÓ y CorpoUrabá: technical and logistical support to Planeta CHB SAS in obtaining harvesting permits. Quantitative analysis of the value chain Quantitative value chain analysis seeks to analyse the distribution of (monetary) value in each link (Marshall et al., 2006). We made this analysis for the value chain of the natural ingredients of the naidí palm, considering that in the case of the frozen pulp, retailers such as MUCHOCOL fix the product’s final price. In this regard, the small-scale harvesters capture only 3.33% of the price of the final product (kilogram of frozen pulp), representing a low distribution of profit to harvesters (Table 4). However, small-scale harvesters and Community Councils own Naidí Pacífico SAS, which manages to obtain a 41.67% share in the final price. Future research should explore how dividends and profits are distributed within the Community Councils to consider a more robust approach. Finally, and as MUCHOCOL stated in the interview, the highest margins of commercialisation, and therefore of profits, are concentrated in the retail link (45%). 1 The purchase price of the naidí fruit shows that any hidden costs of harvesting to communities have not been included. It was assumed that this purchase price was $0 (sensu Valderrama, 2011). However, no associated costs were considered related to the request for sustainable harvest permits or investments in labour, equipment, and tools. Also not accounted for are the diverse intrinsic and non-monetary values of the naidí fruit for local communities. Additionally, the gross income (difference between the purchase and sale price in each link) may not reflect the costing reality of how the naidí fruit is transformed into pulp since approximately 3 kg of fruit is needed to generate 1 kg of pulp. DISCUSSION Governance of the value chain The Colombian Pacific is intimately intertwined with the first links of the value chain. In this region of Colombia, 90% of the population is Afro-descendant, 60% of its inhabitants live in urban areas, and 40% are along some 240 rivers (Grueso et al., 1997). Although Afro-descendant communities have inhabited these lands for centuries, the right of formal ownership via Community Councils1 was not recognised as the highest authority over these territories until the Constitution of 1991 (Peña et al., 2017). The ownership of the land in this region of Colombia belongs to the Community Councils and not to individuals. However, in some cases, within the collective property, there might exist private property boundaries. In addition, within the Community Councils, like property ownership, decisionmaking is undertaken collectively by the community. In the context of the naidí value chain, for Naidí Pacifico SAS, the regulation of collective ownership of the land has allowed biodiversity-based products to be harvested, transformed, and commercialised. This fact is also supported by Decree Currently in Colombia there are 176 Community Councils with collective titles, representing 5% of the national territory. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 129 CHAPTER 9 690 of 2021, stating that only the owners of the land or those acting on response to a request made by authorisation of the owners have a valid right to sustainably use wild flora and NTFP. This means that third parties that are not part of the Community Councils cannot extract biodiversity-based products from these territories for commercial purposes. In this regard, both Naidí Pacífico SAS and Planeta CHB SAS were founded by members of different Community Councils. Therefore, the articulation of processes and the approval of commercial initiatives by the communities has been inclusive from the beginning. Besides, the request for sustainable harvest permits for wild flora to the corresponding local environmental authorities and the socialisation of productive alternatives are developed more organically. Two key aspects can be highlighted by further analysing the governance of naidí’s value chain. For example, the main suppliers of pulp are characterised by operating under a community-based business system. This kind of structure makes relationships and trust fundamental, revealing a horizontal integration. It means that the first links in the naidí value chain can be characterised by being relational: internal transactions are made possible by high levels of trust. However, in the next links of the value chain, where the transformation of the naidí fruit into a natural ingredient occurs, local producers begin to rely heavily on different commercialisation channels (retailers or wholesalers). Conversely, Naidí Pacífico SAS and Planeta CHB depend on very few buyers to satisfy their demand. This type of relationship makes community-based vulnerable to potential volatilities, such as the effects of COVID-19, because they do not have a broader demand. In addition, this also causes them to relate with distributors in a vertical or managerial manner due to their low bargaining power. Livelihoods engaged in the value chain The fruit of the naidí palm has great importance not only for local economies but also for the livelihoods of rural communities that inhabit the Chocó biogeographic region. According to the interviews undertaken, the cultural roots around the fruit are part of the essence of the people who inhabit the forest. Since childhood, local communities have eaten the fruit without its being transformed into pulp in the form of ‘pepiao’ (cooked naidí fruit mixed with powdered milk and sugar), traditional in local consumption and relevant for food security. The fruit has also been consumed historically and traditionally in juices and in the preparation of ‘viche’, which is a fermented drink based on the naidí palm’s fruit. Similarly, the consumption of naidí in several communities has been used for medicinal purposes by the ‘comadronas’ (midwives) for the consumption of pregnant women and to treat anaemia and joint pains. The Community Councils that makeup Naidí Pacifico SAS know very well the properties of the fruit (high content of antioxidants and vitamins), as well as the positive effects of adding it to the diet for personal health, being one of the main products of the family basket during harvest peaks. The sustainable use of the natural ingredients of the naidí palm 130 can be considered substitutes for illicit crops through the guaranteed purchase of raw materials and the generation of complementary income in agroforestry systems, which contribute to the food security of the farms. Additionally, in the municipalities where Planeta CHB is located (Vigía del Fuerte and Bojayá), the naidí palm has been used to construct housing and in traditional fishing techniques. However, neither the heart’s palm nor pulp has been part of the diet in this region, despite the more than 150,000 hectares in these two municipalities of natural forest dominated by naidí. Faced with this situation, the company has started a social appropriation process so that, in hand with the sustainable harvest, local inhabitants begin to consume it. Notwithstanding, other livelihoods of the Medio Atrato region cannot be denied. Income from the different traditional economic activities such as agriculture in the Atrato River basin, fishing in the rainy season, mining in the upper part of the Atrato River, and timber harvesting are also of great importance in this area because they are used to purchase food such as bananas, meat, oil, soft drinks, and potable water from the north of Urabá. Limitations Colombia has approximately 55 million hectares of forest. However, according to the Partnership for Forest (P4F) (Table 1), less than 1% is used sustainably. This situation is not different for the Colombian Pacific region, which has historically been very vulnerable in terms of limited infrastructure, limited geographic access and public services, and has a high level of poverty (Grueso et al., 1997). These, among other elements, represent limitations for developing the NTFP value chain, and in the case of naidí, they are summarised below. • High logistics costs: the lack of road infrastructure and the difficulty in accessing the naidizales (mainly by motorboat) make some areas impossible to use. These situations make transportation costs in the Colombian Pacific very high, making the products that come from Chocó more expensive. • Supply of public services: in municipalities such as Cajambre, public services are limited, which increases the production costs of maintaining the cold chain (temperature control to maintain product quality and safety) and makes it difficult to invest in other areas such as Good Agricultural Practices certifications (GAP). • Incomplete population data for the species: projections cannot be made to guarantee that the process of use and management of the palms are sustainable and that the practices are standardised. • Local knowledge about the palm: for Corpocampo, one of the great limitations in the use of the naidí is the lack of knowledge of the local populations about the palm. For example, they do not know its value and, therefore, they cut the palm as a means of subsistence. CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 9 • Armed conflict: some groups still charge bribes (called “vaccines”) to extract raw materials from the port of Buenaventura, which directly affects the distribution channels. • Pressure from other economic activities: the extraction of wood, a practice carried out for more than 200 years according to Planeta CHB, and illegal mining in the south of the Atrato river, among others, put pressure on the use of the naidí palm. • Limited domestic market: the low cultural appropriation at local and national levels means that the production of companies such as Planeta CHB is not continuous over time, which makes it difficult to maintain contracted personnel throughout the year, forcing the employer to hire new workers each time, and repeat training them. • High certification costs: due to the conservationist vision of forest management, use protocols and permits for the use of NTFPs could cost between 25 and 30 thousand dollars (in P4F estimates). This amount is not affordable for local communities. The same happens with other regulations, such as the work at height regulations. • Discontinuity of cooperation projects: Subsidies and incentives remain essential to guarantee value chains because the commercialisation volumes are insufficient. Without cooperation projects, community-based companies had difficulties providing technical training, which not only affects production but ends up demotivating the people involved, and new personnel need to be hired. • COVID-19 pandemic: deeply affected community enterprises, many of which already depended on a single retailer or wholesaler. People working both in the harvest and in the plant were left without the opportunity to work. Success factors Although the value chain for naidí pulp is still in the process of consolidation and commercialisation, several success factors are highlighted here that can be replicable for other natural ingredients. For instance, the differential factor has been the human factor, i.e., the relationship of trust between small-scale harvesters and food producers. This has allowed the creation of community-based organisations under jointstock companies. In this regard, the Naidí Pacifico SAS is a successful example of how a community-based company, owned by Community Councils (hence families), contributes to local economic growth and territorial development. In addition, the inclusion of diverse segments of the population (youth, women, elderly, and ex-combatants) has been vital to boosting this value chain. Other actors in the value chain, such as Planeta CHB, argue that the needs of local inhabitants must be prioritised. Then later, it is possible to create conservation awareness on the resource since it has value, and, therefore, people take care of it. On the other hand, Corpocampo, despite not being a community-based company, also relies on local communities for the provision of the naidí pulp and canned palm hearts. This company has encouraged the constant support of local communities to generate awareness of the forest’s intrinsic value and to promote social inclusion. Hence, working with local communities is an enabling condition of the value chain that, in the case of Corpocampo, generates sustainability in the supply of raw materials and supports the monitoring and maintenance of the harvest areas. In addition, the company establishes conservation agreements with farmers to undertake forest enrichment activities in degraded areas. This system not only encourages the naidí value chain from its raw material but also gives a sense of belonging to the employees, helping to preserve nature and sustain their livelihoods. The scale and scope of Corpocampo show how incentives within the value chain can foster collaborations within local communities and can create a clear and common purpose in relation to the commercialisation of the naidí palm. Most of the community-based companies addressed in this study emphasised that for these initiatives to be successful, they must offer concrete alternatives of work and complementary income to families at risk of falling into illegal or extractive economies. Therefore, families and individuals should be offered alternatives that are equally profitable, allow the conservation of biodiversity, and to help protect the forest. In this sense, it is essential that downstream valuechain actors downstream value-chain actors guarantee the supply of raw materials. In addition, the assistance of NGOs, national educational services, companies, national public programmes and international cooperation for harvesting, processing, scaling-up and marketing products and biocultural assets are key contributing factors for the sustainability of the value chain. Finally, another successful factor has been large companies leading the market development of the products. In this regard, this kind of company has the financial assets and resources necessary to develop sustained long-term actions and the capacity to share risks with small-scale producers. Opportunities In general, diverse opportunities were identified for scaling up the value chain of the naidí pulp by the interviewed actors. Some of these opportunities include the following approaches or milestones: • Demand-led approach: in the traditional development discourse, assistance programs have usually focused on technical training for producers, forgetting demand consolidation; that is, market creation. Therefore, community-based companies such as Naidí Pacífico SAS and Planeta CHB must take advantage of the market development process that Corpocampo and Crepes and Waffles have done in the national and international markets. • Fund training: it is important to implement agrobiodiverse systems and to develop business capacities in the territories to increase the bargaining power of local communities to access funding. Agro-biodiverse systems such as Agroforestry employing the naidí palm CATALOGUE OF USEFUL PLANTS OF COLOMBIA 131 CHAPTER 9 as a pioneer species for natural regeneration are key with other cash crops that support food security. • Development of logistic chains: subsidise the expansion of cold chains and storage centres to increase production capacity under the maximum yield established by the protocols for sustainably harvesting the fruits of the naidí palm. • Reach economies of scale: not only focus on niche markets, but develop productive and technological capacities to reduce price, reach more customers and be more competitive in the market. • Community-based monitoring: although project execution time is relatively short, monitoring is an important activity to incorporate environmental authorities and the community to generate alternative sources of income and sustainability in the initiatives. • Further efforts in research: implement demonstration plots to determine potential and improve the management of the naidí palm and its NTFP. • Development of local markets and social appropriation of the species: more educational processes are needed to socialise the nutritional virtues of naidí and pulp consumption. For instance, roadshows can be an alternative way of informing rural and urban communities about the naidí palm. In this regard, the promotion of gastronomic tourism around this species is also key. • Diversify products and clients: not relying on a single customer for distribution and diversifying production, for instance, the use of bioproducts of the naidí palm such as fungi that grow in the compost and fermentation of the shell of the palm hearts or residues of the fruit after the pulping process. • Foray into the international market: the transformation of naidí pulp to functional ingredients such as lyophilised powder has great potential for exports to countries such as the United States (State of California), Mexico, and Chile. Therefore, Procolombia should deliver aid to small (community-based) producers and entrepreneurs to access international markets. CONCLUSIONS The historical incidence of the armed conflict, extractive and illegal economies, a lack of public services and limited access to some areas weigh on the value chain of the naidí palm in Colombia. Until alternatives to dignified survival are guaranteed, local inhabitants will not be able to decide how to sustain their livelihoods, especially in areas where food dependence and poverty are very high. The natural ingredients of the naidí palm offer a real economic alternative for the local communities and have the potential to relieve the pressure from other extractive and illegal economic activities, to provide additional income and to diversify the food security of the inhabitants of the ‘naidizales’. However, the pathway for commercial recognition and integration still has a long way 132 to go, as the value chains of the naidí and other NTFPs have not yet been fully appreciated by the national government or incorporated into the country’s business policy. Conversely, the lack of market expansion and the low cultural ownership of the naidí palm at a national level have been decisive factors in generating higher sales, and consequently, better working conditions, or in the search for certifications. Ultimately, the demand is challenging, as the harvest area is approximately 300 thousand hectares, which is still not all currently accessed because commercialisation is still low. Moreover, our analysis revealed that the value chain of the natural ingredients from the naidí palm is not profitable yet unless it receives aid or subsidies. Smallscale producers depend highly on external funds, mainly through international cooperation. Thus, their income generation remains economically unsustainable. More studies are needed to improve the quality control standards of products and to carry out long-term economic, financial and sustainability analyses. Similarly, it is still impossible to achieve a fair and equitable distribution of profits without an in-depth analysis of production costs and considering the scale of businesses. For example, companies like Corpocampo operate under economies of scale, while other companies operate at a small scale. The higher prices may reflect market inefficiencies. However, it is possible to take actions in the short- and medium-term, such as: 1) upgrading the expansion of cold chains and storage centres; 2) fostering incentives for the implementation of agroforestry systems and further investigation on hidden harvesting costs; 3) developing strategies for promoting local diets and cultural acceptance of the species, including investigating the cultural practices; and 4) transfering technology and business capabilities for the commercialisation of natural ingredients of the naidí palm. At the same time, complementing the harvesting of the naidí from the wild in the forest with its cultivation through the development of agroforestry systems would go a long way towards stimulating bioeconomic growth (Papell et al. 2020). More in general, companies in Colombia should be encouraged to focus on the cultivation of native species rather than non-native major crops. Finally, to boost sustainable value chains of natural ingredients in Colombia, the focus should be concentrated on small landscape units to generate diverse mechanisms of self-consumption and complementary sources of income derived from the high diversity of native plant species. Likewise, it would be a mistake to prioritise only the value chain of the naidí pulp while ignoring that families traditionally consume a variety of products, such as chontaduro, borojó, vanilla, and milpesos, and rely on other productive activities, such as fishing, the legal and sustainable use of wood and ecotourism. 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Royal Botanic Gardens, Kew; Instituto de Investigaciones en Recursos Biológicos Alexander von Humbold. 125p. doi: 10.34885/201 Staffas L, Gustavsson M, McCormick K (2013) Strategies and policies for the bioeconomy and bio-based economy: an analysis of official national approaches. Sustainability, 5(6):2751–2769. doi: 10.3390/ su5062751. Valderrama N (2011) Value Chain Investigations on Four Colombian Palm Species. MSc. Thesis. School of Forest Science and Resource Management, Technische Universität München. 113p. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 133 CHAPTER 10 Guacamaya superba (Flor de Inírida de Invierno) in the white sand savannas of Inírida. 134 Mateo Fernandez Lucero CATALOGUE OF USEFUL PLANTS OF COLOMBIA Chapter 10 A taxonomic summary of useful plants in Colombia Mauricio Diazgranados1* 1 Royal Botanic Gardens, Kew, UK *Corresponding author: m.diazgranados@kew.org Keywords: biodiversity, bioprospecting, economic botany, ethnobotany, richness. ABSTRACT As part of the Useful Plants and Fungi of Colombia project, we compiled a comprehensive checklist of useful plants for this country. This chapter provides notes on the taxonomic coverage of this checklist, highlighting the richest plant groups in useful plant species, categories of use, and phylogenetic coverage. It also provides notes on the evolutionary origin of species and comparisons with the Colombian flora as a whole. Potential taxonomic gaps (i.e., missing use reports for plant groups containing useful species) are also identified. The checklist of useful plants comprises 7,472 species, 2,140 genera, and 258 families, representing all major plant groups. More than 50% of the species belong to 18 families. Most of the species are native (78.6%), and 6.4% are endemic to Colombia. At least 14% of the useful species are commercially cultivated, and 8.6% are introduced and naturalised. The richest families in useful plant species are Fabaceae (665 spp.), Asteraceae (418), and Poaceae (338). The richest genera in useful plant species are Solanum (88), Inga (84), and Passiflora (78). The categories of use with the most associated plant species are medicines (5,108 spp.), human food (3,806), and materials (2,363). Before this project, Colombia did not have any updated national list of useful plants, with only a few lists of species by use category (i.e., medicinal, edible, and fibres), often with conflicting taxonomy and incomplete geographic coverage. The annotated checklist published in this volume provides records for more than 4,000 useful species not included in any of the previous lists, integrating all use categories with updated taxonomy, covering the country’s geographic extent, and linking the records with the expanded species profiles in ColPlantA. RESUMEN Como parte del proyecto Plantas y Hongos Útiles de Colombia, un equipo multinacional de investigadores compiló una lista de verificación completa de plantas útiles para este país. Este capítulo proporciona notas sobre la cobertura taxonómica de esta lista de verificación, incluyendo menciones de los taxones más ricos y categorías de uso, cobertura filogenética, notas sobre el origen de las especies y comparaciones con la flora colombiana. También se identifican posibles vacíos taxonómicos (es decir, reportes de uso faltantes de taxones que contienen especies útiles). La lista de verificación de plantas útiles comprende 7.472 especies de plantas con usos documentados, clasificadas en 2.140 géneros y 258 familias, distribuidas equitativamente en la filogenia de plantas. Más del 50% de las especies útiles se clasifican en 18 familias. La mayoría de las especies reportadas son nativas (78,6%) y el 6,4% son endémicas de Colombia. Al menos el 14,4% de las especies útiles se cultivan comercialmente y el 8,6% son introducidas y naturalizadas. Las familias más ricas en especies son Fabaceae (665 spp.), Asteraceae (418) y Poaceae (338), mientras que los géneros más ricos son Solanum (88), Inga (84) y Passiflora (78). Las categorías de uso con más especies son medicamentos (5,108 spp.), alimentación humana (3,806) y materiales (2,363). Antes de este proyecto, Colombia no contaba con ningún listado nacional actualizado de plantas útiles; en cambio, había algunas listas de especies por categoría de uso (es decir, medicinales, comestibles, fibras), a menudo con taxonomía conflictiva y cobertura geográfica incompleta. La lista de verificación anotada publicada en este volumen proporciona registros para más de 4,000 especies útiles no incluidas en ninguna de las listas anteriores, integrando todas las categorías de uso, con una taxonomía revisada, cubriendo la extensión geográfica del país y vinculando los registros con los perfiles expandidos de especies en ColPlantA. INTRODUCTION Colombia is one of the world’s most species-rich countries (Mittermeier et al., 1997; Myers et al., 2000; Raven et al., 2020), holding a broad range of continental ecosystems, from tropical rainforests to mangroves, dry forests, savannas, deserts, mountain forests, and páramos (Galeano, 2016; MADS et al., 2017). From sea level to perennial snow, the large range of elevations is reflected in the wide spectrum of climates and environments (Hooghiemstra & Flantua, 2019). Its tropical position grants high solar energy throughout the year, with reduced climate seasonality, mainly in terms of rainfall variation (Hurtado Montoya & Mesa Sánchez, 2015). The local biota relies on contrasting climate patterns with coasts on the Atlantic and Pacific oceans (Galeano, 2016). CATALOGUE OF USEFUL PLANTS OF COLOMBIA 135 CHAPTER 10 TABLE 1. Diversity of plant species across four datasets: Plants of the World (PW), Useful Plants of the World (WCUP), Checklist of Plants of Colombia (CPC) and Checklist of Useful Plants of Colombia (CUPC). Both the CPC and the CUPC were developed by the Useful Plants and Fungi of Colombia project. CATEGORY PW WCUP %WCUP/PW CPC CUPC %CUPC/CPC %CUPC/ WCUP Divisions 8 6 75.0 6 5 83.3 83.3 Classes 42 14 33.3 17 13 76.5 92.9 Orders 215 101 47.0 107 77 72.0 76.2 Families 952 433 45.5 394 258 65.5 59.6 Genera 14,717 6,737 45.8 3,583 2,140 59.7 31.8 Species 397,692 40,283 10.1 28,947 7,472 25.8 18.6 Colombia’s geographic position at the confluence of South and Central/North America through the Isthmus of Panama facilitated past intercontinental dispersal of terrestrial animals and plants (Bacon et al., 2015; O’Dea et al., 2016), generating complex biotas such as the hyperdiverse flora of Chocó (Pérez-Escobar et al., 2019). The above factors, together with its complex topography, including the three main Andean Cordilleras and numerous isolated mountain systems (Bernal, 2016), create a perfect scenario for speciation processes, resulting in very high biodiversity and endemism (Diazgranados & Barber, 2017; Flantua et al., 2019; Rangel-Ch, 2015; Raven et al., 2020). Colombia ranks second in the world in overall biodiversity and third in plant richness (Mittermeier et al., 1997; Raven et al., 2020), with at least 28,947 species based on ColPlantA (http:// colplanta.org/), of which 93.7% are native and over 32.7% are endemic. Colombia is also ethnically and culturally very rich, with mixed origins from the European colonisation, African descendants, immigrants from the Middle East, and the native indigenous inhabitants (DANE, 2019). At least 115 indigenous groups, with almost two million people, i.e., 4.4% of the country’s population, have survived colonisation in the country (DANE, 2019). All of this contributes to a diverse cultural heritage manifested in how Colombians interact with nature and use plants. The history of colonisation is probably one of the reasons to explain the very long social conflicts that this country has had for centuries. Currently, Colombia remains one of the world’s most unequal countries, where 42.5% of the population are under the line of monetary poverty, and 15.1% are under extreme monetary poverty, with inequality predominant in secondary cities and rural areas in conflict-affected regions (DANE, 2021). In 2019, the Royal Botanic Gardens, Kew, in collaboration with the Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, began the Useful Plants and Fungi of Colombia Project, which aims to empower that country to turn its plant and fungal diversity into an economic resource for sustainable development, by documenting and conducting 136 research into useful plants and fungi, broadly disseminating this knowledge, and promoting and supporting markets for high-value products based on their sustainable use (https:// www.kew.org/upfc; https://in-colombia.org/). Efforts to catalogue the useful plants of Colombia’s territory can be traced back to the Chronicles of the Indies in the sixteenth century (Pardo-Tomás & López Terrada, 1993). During the Royal Botanical Expedition to New Granada (1783–1816), headed by José Celestino Mutis, detailed annotations of plant uses were added to the illustrations of species (Piedrahita, 2008). Humboldt’s expeditions to the Americas (1799–1804) also included mentions of uses of plants (Humboldt & Bonpland, 1809 [1808]). In 1852, José Jerónimo Triana was appointed by the Colombian Government to create the first official inventory of useful plants for the country (Piedrahita & Lourteig, 1989). But it was Enrique Pérez Arbeláez, a century later, who cemented this path with his book “Useful Plants of Colombia”, containing almost 2,000 species (Pérez Arbeláez, 1956). Since then, several publications have emerged, most focused either on specific use categories such as medicinal (Bernal et al., 2011; García Barriga, 1975), edible (García, 2011; IAVH, 2014; López & García, 2021), fibres (Cadena et al., 2007; Linares et al., 2008) and timber (Cárdenas López & López Camacho, 2000; López Camacho et al., 2014), or geographic areas [e.g., Amazon (Cárdenas López & López Camacho, 2000), Orinoco (Acero, 2005; Diazgranados & Moreno, 2020), páramos (Duarte-Abadía & Parra-Ortega, 2015)]. However, these sources often lack taxonomic consistency, and there was no recent consolidated list. Thus, as part of this project, a multinational team of researchers compiled a comprehensive checklist of useful plants for this country, based on previous databases and available literature (see the detailed methodology for the assembly, consolidation, and data cleaning in chapter 12). This chapter aims to provide a summary of the taxonomic coverage of this checklist, including indications of the richest taxa and categories of use, phylogenetic coverage, notes on the origin of species, and brief comparisons with the Colombian flora. CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 10 MATERIALS AND METHODS This chapter analyses the taxonomic breadth of the Annotated Checklist of Useful Plants of Colombia (CUPC hereafter), available in chapter 12. The checklist resulted from the compilation of several data sources, taxonomic reconciliation, classification of reported uses, and data normalisation (for detailed methods, please refer to chapter 12). The CUPC was compared against the Checklist of Plants of Colombia (CPC hereafter), being updated as part of this project and available through ColPlantA (http://colplanta. org/). The CPC comprises 28,947 species, classified into 3,583 genera and 394 families (Table 1), ca. 3,000 more species than are currently reported by the Colombian Catalogue of Plants and Lichens, with 26,134 species (Bernal et al., 2019). Results were also compared with the World Checklist of Useful Plants (WCUP), produced by RBG Kew, the most comprehensive checklist that exists today (Diazgranados et al., 2020). It comprises 40,292 taxon names (including a few organisms frequently misidentified as plants, such as brown algae and nostoc), classified into three kingdoms (Plantae with 40,283 species, Chromista with eight species, and Bacteria with one species), six divisions/phyla, 14 classes, 101 orders, 433 families, and 6,737 genera (Diazgranados et al., 2020). The statistical analyses and figures were elaborated in R v.4.1.0, mainly using ggplot2 v.3.3.5 (Wickham et al., 2016) for graphical representations, and dplyr v.1.0.7 (Wickham et al., 2021) for efficiently manipulating datasets and summarising results. The categories of use were mapped onto a modified phylogenetic tree of vascular plants (Gastauer and Meira, 2017), using the packages ggtree v.3.0.2 (Yu et al., 2017), ggplot2, and treeio 3.13 (Wang et al., 2020) to visualise how useful taxa are distributed across the plant phylogeny and to compute the richness of useful species by clades. Considering that species often have multiple uses, a chord diagram was calculated using circlize v.0.4.13 (Gu et al., 2014) to reveal the network of inter-relationships among multiple use categories. Since some use categories could be correlated with each other, corrplot v.0.90 (Wei et al., 2017) was used to calculate the correlogram with the Pearson correlation coefficient (ρ), which measures the strength and direction of the linear relationship between two variables (Hauke & Kossowski, 2011). FIGURE 1. Treemap of the richness of useful species of Colombia by families. Red polygon: families comprising more than 50% of the useful species. Yellow polygon: families comprising more than 75% of the useful species. Green polygon: families (69) containing only one or two useful species. The numbers of species per family are indicated in chapter 12. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 137 CHAPTER 10 RESULTS Richness by taxa and species origin The CUPC comprises 7,472 species of useful plants, (7,472 species when including a few organisms frequently misidentified as plants, such as brown algae and nostoc) classified into 2,140 genera, 258 families, 77 orders, 13 classes, and five divisions (Table 1). When comparing these numbers to the CPC (28,947 spp.), almost one in four Colombian plant species has at least one reported use. Also, 59.7% of the genera and 65.5% of the families in the CPC contain useful plant species. More than 50% of the species are classified within 18 families, with 69 families containing only one or two useful species (Figure 1). Most of the useful species in Colombia are native (5,881 spp., 78.6%), and 481 species (6.4%) are endemic to this country. At least 1,077 species (14.4%) are commercially cultivated, and 643 (8.6%) are introduced and naturalised. These numbers are relevant considering that only 677 species (2.3% of the total) in the CPC have been identified as naturalised. In other words, 95% of the naturalised species in Colombia have reported uses. We could not report the origin for 12.8% of the species, because they are either cryptogenic species or presumed to be exotic. Still, we do not have confirmation of their naturalised status. Most families (155) comprise just native (incl. endemic) species (i.e., with no introduced species), with Annonaceae (102 spp.), Piperaceae (102), and Sapindaceae (73) being the richest ones. Three families include only introduced useful species: Balsaminaceae (5 spp.), Musaceae (7), and Sphenocleaceae (1). Overall, the most species-rich families in terms of useful species are Fabaceae (665 species), Asteraceae (418), Poaceae (338), Rubiaceae (285), and Malvaceae (221). At the same time, the richest genera are Solanum (88), Inga (84), Passiflora (78), Miconia (77), and Piper (67) (Table 2), which all have mainly native species, including several endemics to the country. TABLE 2. Top species-rich families and genera in the Checklist of Useful Plants of Colombia (CUPC) and the checklist of plants of Colombia (CPC). Top 10 most species-rich families CPC Top 10 most species-rich genera CUPC CPC CUPC Orchidaceae 4,436 Fabaceae 665 Epidendrum 558 Solanum 88 Asteraceae 1,405 Asteraceae 418 Stelis 450 Inga 84 Fabaceae 1,317 Poaceae 338 Piper 429 Passiflora 78 Rubiaceae 1,276 Rubiaceae 285 Miconia 414 Miconia 77 Melastomataceae 995 Malvaceae 221 Anthurium 392 Piper 67 Araceae 919 Arecaceae 214 Lepanthes 323 Maxillaria 62 Poaceae 919 Orchidaceae 195 Palicourea 313 Pouteria 54 Piperaceae 719 Solanaceae 171 Maxillaria 288 Ficus 46 Polypodiaceae 614 Melastomataceae 162 Peperomia 287 Ipomoea 43 Bromeliaceae 553 Euphorbiaceae 149 Philodendron 225 Peperomia 42 TABLE 3. Top endemic-species-rich families and genera in the Checklist of Useful Plants of Colombia (CUPC) and the Checklist of Plants of Colombia (CPC). Top 10 most endemic species-rich families CPC 138 Top 10 most endemic species-rich genera CUPC CPC CUPC Orchidaceae 2,097 Asteraceae 67 Stelis 323 Espeletia 29 Asteraceae 637 Melastomataceae 40 Epidendrum 238 Clidemia 17 Melastomataceae 474 Orchidaceae 27 Lepanthes 235 Passiflora 11 Araceae 403 Fabaceae 24 Anthurium 213 Oreopanax 9 Rubiaceae 295 Arecaceae 18 Piper 187 Leandra 8 Piperaceae 282 Araceae 17 Miconia 158 Magnolia 8 Lejeuneaceae 258 Malvaceae 16 Pleurothallis 114 Symplocos 7 Fabaceae 211 Araliaceae 14 Masdevallia 104 Matisia 7 Acanthaceae 208 Passifloraceae 11 Peperomia 95 Miconia 7 Bromeliaceae 186 Piperaceae 11 Maxillaria 94 Solanum 6 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 10 TABLE 4. Top naturalised-species-rich families and genera in the Checklist of Useful Plants of Colombia (CUPC) and the Checklist of Plants of Colombia (CPC). Top 10 most naturalised species-rich families CPC Top 10 most naturalised species-rich genera CUPC CPC CUPC Poaceae 160 Poaceae 153 Ipomoea 12 Eragrostis 12 Fabaceae 93 Fabaceae 86 Eragrostis 12 Ipomoea 10 Asteraceae 80 Asteraceae 77 Digitaria 10 Digitaria 10 Brassicaceae 24 Brassicaceae 24 Bambusa 9 Bambusa 9 Solanaceae 20 Solanaceae 20 Senna 8 Crotalaria 8 Lamiaceae 19 Lamiaceae 19 Crotalaria 8 Urochloa 8 Rubiaceae 16 Rubiaceae 15 Urochloa 8 Senna 7 Convolvulaceae 15 Convolvulaceae 13 Cenchrus 7 Cyperus 7 Cucurbitaceae 13 Polygonaceae 12 Cyperus 7 Setaria 7 Polygonaceae 12 Cucurbitaceae 12 Setaria 7 Trifolium 6 5108 Medicines 3806 Human Food 2363 Materials 2208 Environmental Uses 1038 Animal Food Poisons 734 Gene Sources 724 650 Social Uses 407 Fuels 182 Invertebrate Food 0 1000 2000 3000 4000 5000 Number of species FIGURE 2. Number of plant species per category of use in the CUPC. 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D C Saxifragales Saxifragales G r A S oss Tepha C axif ula Ite tra no ras ra ria ace g Pecar pet sula ac cea ae Ha nt pa ala ce ea e lor ho ea ce ae e Me Ge VaG garacecea ae e r Vivlian aniaitacrecoea ae e t FA s ce ae s r ia h cS Co TGapnchnia aC e aae u Ite m re o ce a x l On beret yiaacacneaareae ifr ar a e oe s a ia c L ag rtaacc a e aep s g c ea Me Vo Mythr rP acceeaaaer et ula ac ea e Cr lastochysyrta e p a e e yp m iaHceean ter ata ceaaee th ae lac ce ae A o al a Pelzateniacceeaeeora or ac ea e G a n acG G e eroae V ga ce ea e Streisso AphloaM esrsa i asb lomA iaaeceSaae ph ceCliaaex u nIt ta ce ae e S urg Taeta e Gu taphy eF ri t Vcaien araenifr lar i ac ce ae e leaarrcaec avoaipeassutahgaiaac c Cro Staamate Pceaare etn la acceaeae ae sso cC epa alaiace eae e ae ceeanan s hyuralaHG ea cec a ea ethc ceaalo Bie Picraomaota r oro e e e r e a c bers mnM m e a a c a gai ce ecaee ae areV y iaecGeeeb a ea inO r it c Nitte a a a e li ia n a c e e niata ra Vivceaa ce aeee e ar ceiaanetg KirkriFL aytaneiaharcaceacae e ae ia urs MeAnaBVca Gcr ena e r c ec a e a c coh oCrdoemraM e acaeaeeea e breeeyayria ce CSrimlSaaarosptuinoOdLcaianhce a tc eacee e ay g taeceaec aee ae bm ayce thrarasaeecia MeypVM a M el t a e aaeatcaeeacee e lasRoe chriayce rta GC A erryra yosn tout ac ae ae ce c eai e ia prd inmac telz et ataea cea Pen eacae eae roa Aelzae t Ge DipePTa c e ac n iaea naiateace aeaeeeae Ppi cae Ge ntA ensc eec od aea Str Sis Ap sissNe ata pon eace eacae eae hh olour astrTh lolac o caceae aso mad ym m aeeeea bulo ela Spha aea eae rg ataiatcceaceiea ac e e SGtauSbertauos epeala prhg acee ae eriaraac yBix cce ae le eea e Staapmhate CG a ia rou c e e e y c yMale la acee ae lva m cece ura so hCy Cro BStsPasaicoMu aeae ae ceacea atta etina ac ssieobecrsrahmymantin gia e eeeae cla cea e n c a ia Cistace teinu ee sSar oitm N aeaccea iarcce cola e aeae ra Bie PicDipt riatcena eeae eroc aeacaeceae irkiaaarpa beArn rBaursKm eae e ceaiacea nia eAkan sate ra Tropa e ce aee rdia ceolace NSaitaca ae ceaaee pinin ia Sim Moringa eeceae ceac roruabdraia Caricacea a ee KMirelkiaacecaeee aeae Setchellanthac iacecaee e utac erurarrd AnGaB sRKoeberliniaceae eae ae eLimnanthaceae inrac Pet aea cenrae cee a aceaBataceae SantTapa ia piscdia Dipe Salvadoraceae Sim eeaeeceae cecae odin ond ta a a Emblingia Ne ce cae ur ea ceae uadeaac Th roPentad iplandra acecea elab Sphaerym Mep eea eiaceae Tovar os ae eala ae liace Reseedaceeae ae cstem e Bixac Gerr RMau onac e tace eae Gyro lva ceaeae cea ard Cap aepara Cyin tina Pete a cea omaceae c Mu Cle e ntin e ae naCis ceaBra eaessicaceeae egia a T c tace a riac e ae p Dipen Sarcola aKraemeyllaceae iscena iaccea Dipt toderoc eZy aego onarpa e phillajacceeaaee taiacea N Tropa cceae ba ae eaeeQuFa raAkan deolace Thymeeu ae Surianaceea a c e Moringa a ac ceae e Sphae laeace e Polygalsaceaee roSetchellanthac ae Roeyaceaee sepaCaricacea la eae rb acea G ae Baea Limnanthaceae gn ceae r Bixacce Koeberliniaceae eae Elaha S oss mnaac eaAepe M alvaBataceae Te ha C axif ula Ite chm acea ceae DRira r a Salvadoraceae Cyt Ulmbaceatree a no as ra ria c inacea ceae Emblingia na raceaP ecar pet sulagac ceaeae Mun e n a tin ceae C Mo ce iplandra H aenet pa ala ce ea e Pentadgia ceiaceae ae ica ceaa e e h rt Tovar lo U Ci Rese ga are o a ce ae ae cedace aeeae ofa MaegGaeceeVaaegaracecea ae Sarcolaesta th o onac F licarca aitae c a e N stem G ceae Gyrona e e ceae Vyri a cne Dipteroca r Cappara M Frlaivniadnaathcaeiaaceceea ae rpa A S oss cea omace Cle e ae ganarinia cceaeae e ae uG ace J C Tepha C axif ula Ite e Akania ssic e c Bracea omasu re nodaralaceceaaeae eceeae tra no ras ra ria ace riac Krameylla Tropaeolace g OCnicbordeeyia ae tuce caee e ae Pecar pet sula ac cea ae T ag B taecyclleeaaaaeceaae eae Zygophilla L a jac e p H h y ri n e r Moringace e e p Qu ae a a Mae alo th ae lac cea ae e elaVoc MAynthisraocCcoeariaaerpaacceaaee Fabace ae ra ora ac ea e rta e oce th ce e M G Caricaceae naceC ryepstomhysia cryenadean ita ea e Suriaal er Vit gac ce eae e acea e c te a o a c a ae lyg a a li rb o Setchellanthaceae Po Rosace Ae ron taCc eAap e cu ela ce ae V an nia ac ea e lza ia ea eCu am ca ce eF ivia th c ea e ea ac P s r ey c te a Ge acAeaeen a ea e etr ati nia ce e an n ac ea e Limnanthaceae BaSrb T D go ya Coea Ge co ia ea e a iss pae gn c h e eacceaee r tram oce Koeberliniaceae Elaea loea ea ey a ea e Be botrstracOcm sna ia e m lo buac Rhach a nebareee iac cea e o e ce ae Sm rg ata Bataceae pid ela Hu rLayccaegar taeceeaee GuUtalm p aceea ri ece ae Dira Le MCe VonaM ambhayce athcreacaee ae leaaaaceeaae SalvadoraCceae ara rosCSata caeea e osnxcahlidnyia nnca tece rtaaaceceaaeee Cr la la C h o s y to y e o M a e c e ce o c s y ura O umn riap ec aeea e somrtic a eeaae pte Emblingiacea Bie Picra ata gaccceeaaeee A Croocaataellciaetac eea e berseU monfa thin Pentadiplandracea Pelzlaaetenruianccaeeloageiaec cea ae e te agcaeecaeeaee G Fia N En B h ae da ce a Noite Tovariacea ra iayric caae cea e Streisso AphloaeaeapcIervain KirekriMalaceneadaaeaceaee as lom iaC ce ean onalace ea e Burs ResedaAncea uiagcueaarine ceaae Staburge atacceaaeeP phoxy rac ceaeae e Jra ee ra a s e e ca cea d G p c ri a eane lac eae ua hyle ac eaneolothr phohnaiac ea ae ap rdCia tu Gyrostemona ce odaeB SimSce ae Sta mate ac eatee ry o c et iac ce ae e dTaicce ee eac Cero arouinae Cappara apehyllariacceaseesosochyuralaceeCae ERhiz Oonn lus lla ace ea bace ae y o ce M a ri P a a m aeCo rp Bciee ic eliace c e B C ph ric ac nisae CleoGma Ae ca a baeram atac eae utac erra lo pe em acRea eaoerynodanthitacceeearseeteinniaceeae Petenrdinac Brassic Ca Hy ost ea C po urb la eNaitra ceae ae e iac ae d er Taia eaeeA CuctramescaBccK eairakeriaceeaae pisciaacae Dipent Kram Po ti ce ia ce ia u e e e od la e a ae rs ontace T D gAonnayac eeara eceae phNeylurja ae caard cea c ia ae ZygoTh e ce a tr ad S ac e o B e llaelaeaceea ym eaee c ece a aeee S b asptrin SpQ haui erosep aro uadacce idoimla a ae acceae ala Fab ceeaaeee ae Lep Ce oMnHneluabliiardaace ace aea Bix ce c a ac n ea a lvacecae e e a acniaaae Ma G C Rxut ce ae SuriCy o rea p iaeceeaee in alaceaceeeaae PeretraenrdOae unac gtina aealle oly ntin PMu gia oc ea tac cea ae a e aeDipe TapisclaCeiaac scea ne e Cisotace R ru hae alo gia ace eae e ntodon E Bce Sarcolaena aaece e ae Ne in d c a eycea epae DiptB uradta IrvPan onalace eaeae eSp eThymela eroc arb Cce c arpaceae ac a ea n e g Akan eaceae phoxy rac ce eae e aiaceaae ceahaerosepala lo hr ho na c ea e Tropa Elaeeolace n o e ce ae a t ae p n Bix ea Moringa ce hamceae tee ry izo ch etiaiac cea ae e RCaricacea Malvaac hmeaaceae Cy ceCaeERh Oonn lusylla ace ea Setchellanthac ceae DiracUeae B C ph ric ac lm aceaMuentintina Limnanthaceae ceae lo pe em eCisgia b cSar eacola Koeberliniaceae ae Ca Hy ost annMaoraDipt e enatace ceae d a CBataceae e eroc ceae Po Salvadoraceaeticac Akan earpa a e r iacea e Emblingiaceae c U gaTropa e ae eolace a e Pentadiplandraceae fa c Moringa iaceae Tovar thoaeFaga cCaricacea ae ceaee e Nodace Rese a eae eae onaceaeeySetchellanthac ricLimnanthaceae Gyrostempara cea M dac eaee Cap aelanKoeberliniaceae Cleomace gae arinaccBataceae ace ea e Jueae Brassicriac dralaceaceae Krameylla asaeue enSalvadoraceae ce Emblingia C aee ph go d etuiplandra Zy eceae jacea Pentad Quillaba icoae iaceae Tovar eacdace FaT ce aeB yll ceaae naceea aee Rese ia h e r eeae Suriaal p ac o onac ia para rstem accea ae e ce Polyg iseaaeGyro o p sa r n e Cap Ro ceae a om A ac eaC e ocCle ea e ae thaace Barbey ssic gnaccery aceae aen Bra tace anme i riac Elaeamnao d e caeeeaae b ea Kra r o aea lce Rha C uphylla macA ejac eapee Zy c ce e go ac u a Dirach lm m illa a U bace C Qu ba aecea e ae etra Fa na tisnacnce ae ea e ceia ae Suaria Can Morace eaaec ea ace Te D galosaactce lyg Urticgaceeaae Poe c e ryeaae fa gac ae B Roo o treaeaceaae th ac b sea No Fa cace ae Baorbeyaac e l ri d u ce ae gn c y i e a ea M nd ceapEla e ceae la a e e C mna Heaaeera ce ae JugsuarinraLceaae Rhachmac neaaleida ce ae nace CaodendtulaceeaeDira Ulm oac nabra xceaaaoeeniaaceeaee Tic BeylleaacceaaeeCanC MoO ce neaae rp iac ea e ph ri e rticau iso ria ac ae gaccc An Coocarpthaceeaee Uofa C eae ll ac ea e c ageaoceanee ot c a ryn an ita eaaoeth Fa CoApodcurb elaccN e e Mlyricaarcueaaeeal gia ce ae e d B n Cu tramtisca ceaaeE cehae in da ce a e gla na p cerave n a ce ae Te Da gonia acceJauaeseuariCde ra y Ie n ea en lac aeea o la ce ae Be botrstra C acciceoadaeeBetulleaceeP o a eph xy a e ae r c pid ela HuarT a ce apehy riac eo a c l aae ro o a ce ae Le C o a ce aoeri rpaao nn lid nis ceetahe ph hn ia e ae n CoOxa oAniaaceCneaoeceae th e t c n rp oiaryc oedaat enrbitarcyceiazaeo e Oc e ia ce a e c E a la ll Cuoca C e u p C e n s c e e rune loAtaiaCcueceaaeameesR cahceaae on lu lla ac ea la tr a c ti g ia e e e E B ph in dTa cD e aa on ac B ae C hy ic c e Ce IrvPan onalaBceecegbaoatreytraceceaee op er ma a ocelaase eua ceaae l p e ph xy rid t a aera ce a olo hro heop naCece eH aee y s en ryt oLp ch tia concena liadeaeiacCeeaH o ae Ct E hiz O nne sia c ed lu CllaOxc oean pa eaoe R Bo C phy ricCauanccar lliacacPeeaaee e o t e m a c n lo e p e u lo ia e e Ca Hy osEt la Br pha ing dac cea ae e d Ce IrvPan onalacecea ae Po ph xy ra ce ae e olo hro ho na ce ea e en ryt op ch tia c ea e Ct E hiz O nne siallac cea ae lu e R Bo C phy rica ac lo pe em Ca Hy ost d Po Monocots Tecophilaeaceae Ixioliriaceae Hypoxidaceae Asteliaceae Lanariaceae Blandfordiaceae Boryacea Orchida e Liliaceaeceae Smilacace ae Philesiac eae Ripogon aceae Alstroem eriacea Colch Pete icaceae e Melanrmanniace th Ca iace ae Cormpynem ae Cyc siacea ataceae Pan lantha e Stemdanaceceae onac ae Triurid V a eae The highest number of useful endemic species is found within the families Asteraceae, Melastomataceae, and Orchidaceae (67, 40, and 27 spp., respectively) and within the genera Espeletia, Clidemia, and Passiflora (29, 17, and 11 spp., respectively) (Table 3). Most of the top 10 richest genera in useful endemic species are not within the 50 richest genera in the CPC. Indeed, some occupy discrete positions in the ranking: Matisia (52 spp., ranked in 81st position, with 21 useful spp. of which seven are endemic), Magnolia (42, 109th, 13, 8), Symplocos (34, 149th, 12, 7), and Leandra (28, 210th, 8, 8). As for the naturalised species, the families that contain more useful species are Poaceae, Fabaceae, and Asteraceae (153, 86, and 77 species, respectively), and the genera are Eragrostis, Ipomoea, and Digitaria (12, 10, and 10 species, respectively) (Table 4). Saxifragales Malvids Monocots Monoco ere ytaceae Didilo Ha ph ceae Montiaginaceae Mollu aceae Nyctagin riaceae Petivelac caceae Phyto ataceae Sarcob ceae Gisekia ae Aizoace ceae Barbeuia Kewaceae ceae Lophiocarpa Limeaceae Stegnospermataceae Amaranthaceae Achatocarpaceae Lo C p a Ce Pu hopryo nt tra yx car r M op nji ida ac B a Ela la va c ea Dic T ala lpig tin cac ce eae e Lo C Ch E ha rig no hia ace ea ae p a ry up pe on pa ce ae e Ce Pu hopryo so hr ta ia ce ae nt tra yx car H ba on lac ce ae r A um lan iac ea ae M E oplanjivida ace Gocha iria ace eae e alp la c a ce a B r ala ig tin a ce a e D La Pa V up iac ceaae e cis ssif io iac ea e Ch EichaTrig no hia acecea ae te lo lac ea e ry up pe on pa ce ae e so hr ta ia ce ae Samataraceeaee li Hubalaonialacecea ae EuRaff Percacceaaee Ac m na ce ae e le e PicPhyllphor siaaceaae ha iria ce ae G rod an bia cea e La Pa Voup riac ceaae en tha cea e cis ssif io iac ea e d c te lo lac ea e Be AIxona Lin raceeaee rbe ex nth ac ae Samataraceeaee li Eryrido toxic aceeae E Ra Pe ca ceaae Str thropsidaaceaae Ph uphoffles racceaee om pala ce e P icr ylla rb iaceeae bos ce ae od nth iac a ia C a e e e Xim oula cea e Ixo Lndraaceaae Apta enia cea e B erb Ae nan ina cea e ndra cea e Octo Ola e x th c e Eryrido toxic aceeae c ce e L knem ace ae Str thropsidaaceaae S oran ac ae om pala ce e Misochoepthaceeae bos ce ae a fi de ac e XimCoulaiaceaae Bala Opndraceeaae Apta e c e ili n ndnraiaceeaae Comophoraaceaee Octo Ola andr ceae c ce e acea CervaThesiac L knem ace ae e Schoranth aceaae esiaceeae Nannt a e M o c e is od ode pfiac eae ce ae Amph Santalea Bala Opndraceeaae orogyn aceaae ili n aceaee Comophoraaceaee Viscac Franke andr ceae e niaceea ac T Tamaric he ea Cerva si e Plumbag aceaae es aceae Nannt inaceaee Polygon od iaceae aceae ce Amph Santalea Dro ace orogyn aceaae Nepenser aceaee thaceaae Drosophyll e Viscac Franke aceae eae Ancistroclad nia Ta ric ceae aceae Dioncophyllac Plumbma ea eae aginaac Rhabdodendrac ceaee Polygon eae Simmondsiaceae aceae Dro ace Asteropeiaceae Nepenser thaceaae Physenaceae Drosophyll e aceae Macarthuriaceae Ancistroclad aceae Microteaceae Dioncophyllac eae Caryophyllaceae Rhabdodendrac eae Simmondsiaceae Asteropeiaceae Physenaceae Macarthuriaceae Microteaceae Caryophyllaceae Tecophilaeaceae Ixioliriaceae Hypoxidaceae Asteliaceae Lanariaceae Blandfordia Boryaceae ceae Orchida Liliaceaeceae Smilacace ae Philesiac eae Ripogon TecophilaeaceAls aetro aceae emeriac Ixioliriaceae Colch eae HypoxidaceaePe icaceae terman Asteliaceae M niaceae Lanariaceae Caelanthiace ae Blandfordia Cormpynem siacea atacea Boryacea ceae Cycla e Orchida e Pan nt e danahaceae LiliaceaeceaeSte Smilacace Triumonacceae ae Philesiac Ve ridaceaeae eae ziace e Ripogon D llo acea Alstroem Buio er score ae Doryanthaceae Iridaceae Xeronemataceae Asphodelaceae Xanthorrhoeaceae ae Amaryllidace Asparagaceae eae Dasypogonac Arecaceaee linacea Commeana ceae ngu Doryanthaceae HaPhilydraceae ae Iridaceae odorace em ceaee Xeronemataceae Ha ntederia Asphodelaceae PoHeliconiacea e aceaae Musia Xanthorrhoeaceae ce e ae Amaryllidace ea Strelitz Asparagaceae Lowiacceae eae Canna ceae Dasypogonac ta e an Arecaceaee Marib eraceaae linacea ce Commeana ZingC ceae osta ceae Hangu raceae ae Typeha Philyd ceae liace ora ceae ae Brom Haemod riaceea atea aee Rap yridace a Pontelicde iac ee X aulaceeae He on aceaae c c Musia ce e Erioayacaiaceaaee ea M urn ce e Strelitz Lowiacceae Th unca cea e J era ea Canna ceae antaac Cyprthriaacceaee eae Marib er ce a ae Anoalepidnaceeae ZingC osta ceae ntr tio ac ae ae Typeha Ce Resellarioaceeae liace g P ac ae ceae Brom e Fla ateaaceaee villeleac eaee Rap a Xyraidulaceeae Joinioco yllacceaae c c e ph a e e d Erioayacaiaceaaee Eceratoupteleeracceaae M urn ce e C E pavbalaace ae Th unca cea e J era ea Padiza ster aceeaee r m c a Cyprthriaacceaee La caeaper ridalaceeaee a Analepidnaceeae Cir enis rbe cu iac ea e M ntro tio ac ae Be nunSab nac ceaae Ce Resellarioaceeae Ra mbotanaaceeae e g P ac ae e Fla lu la te ac ea e villeleac eaee Ne P Prondr xac cea ae e Joinioco yllaccea e de Bu na ce a e de ph a ea e ho am eraiace ea ae Ec rato pteleeraccea e c h n oc e Ce Eu pavbalaaceaae Tr yrot un illeniscaiace ea ae Padiza ster aceeaee M G D rid on giac ce eae e r m c a La aeaper rida ce ae Pe aeltin lidallac ceaeae c e e la P A me hy lla c Cir nis rbe cu c ea e y ia Me Be nun abianac ceaae a S m idipiph or Ra mbotanaaceeae e Ha ercphn nom lu la te ac ea e C a y Ne P Prondr xac cea ae e D C de Bu na ce a e ho am eraiace ea ae c h n oc e Tr yrot un illeniscaiace ea ae M G D rid on giac ce eae e Pe aeltin lidallac ceaeae P A me hy lla c y ia a m idipiph or Ha ercphn nom C a y D C Categories of use Malvids The categories of use with most species are medicines Malvids (5,108 spp.), human food (3,806), materials (2,363), and Saxifragales environmental uses (2,208), the latter including ornamental species (Figure 2). Saxifragales Useful taxa are fairly distributed across the plant phylogeny, with most families having many types of use (Figure 3). Monilophytes (i.e., ferns and horsetails) contribute with the fewest useful taxa (178 spp., 2.4%), whereas Rosids [Vitales + Eurosids (Fabids + Malvids)] is the richest among the large clades (2,795 spp., 37.4%). Within Rosids, Fabids are the richest subclade (1,809 spp., 24.2%): Fabales (688 spp.), Malpighiales (647), Rosales (291), Cucurbitales (76), Malvids Oxalidales (58), Celastrales (31), Zygophyllales (9), and Fagales (9). Asterids [Cornales + Ericales + Eurasterids Malvids Fabids (Lamiids + Campanulids)] is the second richest large clade (2,204 spp., 29.5%). Within Asterids, Lamiids (1,299 spp., 17.4%) is the richest subclade: Lamiales (509 spp.), Gentianales (474), Solanales (250), Boraginales (58), Fabids Icacinales (6), and Metteniusales (2). Half of the species have more than one type of use (Figure 4). For instance, some species used as human food can also have social uses or are sources of materials and poisons. Uses in all ten categories are reported by 28 species [incl. 22 cultivated (e.g., Arachis hypogaea, Zea mays), 11 naturalised (e.g., Cocos nucifera, Persea americana), and eight natives (e.g., Anacardium occidentale and Spondias mombin)]. With at least four types of use, some endemic species are quite versatile, e.g., Hesperomeles goudotiana (5 categories of use), Astrocaryum malybo (4), and Passiflora antioquiensis Fabids (4). There are no significant correlations between categories Fabids of use (-0.13 ≤ ρ ≤ 0.36), with human food and medicinal plants potentially having any other type of use (Figure 5). The Fabids cladogram shows the highest correlations between animal food and fuel, social use and materials, and poisons and gene sources. Caryophy C sa chy Nys rostaae Hyd tace ceae eae Cactulaca erotac Por campse Analinacea Ta sellaceaeae Ba ereace eae ac Didilo yt Ha ph ceae Montiaginaceae Mollu aceae Nyctagin riaceae Petivelac caceae Phyto ataceae Sarcob ceae Gisekia ae Aizoace Barbeuiaceae Kewaceae ceae Lophiocarpa Limeaceae Stegnospermataceae Amaranthaceae Achatocarpaceae Santalales Santalales ae d Amph Santaelaaceae orogy ceae n Viscaaceae Franke ceae Tam niace Plumbaaricaceaaee naceae Polyggi on Droseraaceae ce Hydasace ae Montia aceae Lo rnace ae ceae Mollugin aceae ce ya Co Nyctagin ceae sa ch ria Nys rostaae eae Petivelac Hyd tace ceae eaPh e yto cac e Cac laca otac Sarcobatacea e Portucampseer Gisekiacea Analinacea Aizoaceae Ta sellaceaeae Barbeuiaceae ce Ba ceae ea Kewa e er Didilo ytacea Lophiocarpaceae Ha ph ceae imeaceae Montiaginaceae L tegnospermataceae Mollu ginaceae S Amaranthaceae Nycta riaceae Achatocarpaceae Petivelac caceae Phyto ataceae Sarcob ceae Gisekia ae Aizoace ceae Barbeuia Kewaceae ceae Lophiocarpa Limeaceae Stegnospermataceae Amaranthaceae Achatocarpaceae Lo C p a Ce Pu hopryo nt tra yx car r M op nji ida ac Ba alpElat lac vac ce eae Dic T la ig in ac e ae Ch E ha rig no hia ace ea ae ry up pe on pa ce ae e so hr ta ia ce Lae Hubalaonialac cea aeo C Lo C Ach mir nac ce eae e ph ary p a G a ia e aCe P o CLea PPuahsopryVooupiariac ceaae e u p oc cioa c ea e nt tra yx ar trasifylo nctiste la x r e e r ro n i a M oSp mnjia idracacceeaaee M B RalpElaPt alaliccvta E p ji da c a ce a affig ine aacccceeeaaaeee B a la la va ce ea D Euala eaaee e Ch EPichPaThryigphno lehia a lp t c c s araceceeD ceceaeaaeeie T la ig ina ac ea ae ry uicp ropellaonn prbaiaia so hr deta iathCceceaeae ch r n h i c e e i o ba on nladr caecheaeaeE g Ix a H o la Lia ca a a u o p ac e ae Be A Aum rbe chexnirannathinceaecaceeeareyes p pe n ac ea ae G rida toiax ceacaeeae o h ta iac e e r orpias cicea aeea b o a La Pa SEryVothupro l cis sstrifom io iapcacideaa acceeeaee Hu ala nia ac ea e e te lo la boc ela a ce ae A SamXatarCaoceuseia c m na ce a e a e ea li imcee alaececeaee G ha iria ce a e acn naia eae EuRaff PAerpcta ou ria ce a e le edarae ceP PicPhyllphOocr tosiaaOcla eaceaaa e pia ce aee eacL V s rod an Lbiakncem a c e e enSthaoracenaaeacceeaiest sif iola ce ae l dc th Lis rahoceea eac ae em ora ce ae Be AIxona M rbe ex nthinaocdceenadpeefiaceeaae S a c a e Bxaic ce Eryrido to laanaoceOaapeilira Raee Palicataceeaee ace cepaheoE Str thropsid eff e aom om paC ceaaea an edrrau r l aeP boCs lacceTehe ph es acceaae acea er iacvaP ae siac XimCoula eant criaohceyeaae llee orb iac ea e Apta eies e Nce an aeod ea d ant iac ea e ndniaceSa ae Octo OAm en ha ea e entalacce phcor lacra eaaog ea k e a e yn n em cea ViIxoaceae dra ce e L e scac cean S oranB aFr nea aeke Aeniace aceb aneLinacceaaee Misochoepthe ae de fiacreTa ma e ric xtea t ridinaoac oae ae mb Bala OpndraPlu xe hac eae ceE ceae arelyg noph iliace yagton Com ora SPo acp seidicaceeae hrace ea e ser andr ceaaDro t o r e acea o thacea Nepen paeae laace ae CervaThesiac e m Dro b sop nt ea hyll oae ace e Nanodesia Anc sia ce ae ce istro Cace oeae Dion ce ae Xclad coph Amph Santalea uaela ceaaee yllac im aceaae orogynRhab A dode e ndrac acSimm neae eae p taneceae iacceeae Viscac ondsia Franke ea d Asterop e Oeiaceae rac ae nia Ocecae Ta ric laceae Physenac eae Plumbma eato aginaac iaceaac e ceM LaeeaocMrkaaircntrhoeuterm Polygon acaeaeeae ac e Seacae Dro ace Caryophyllaceae ho ntha ceae Nepenser M thais cea Drosophyll ed epfia ceae o Ancistroclad aceae end aceae racceae Dioncoph Bayllac eae Opil laneae Rhabdodendrac op iac eae Simmondsia Coceae manhoraceeae Asteropeiaceae drac ae Physenaceae T MacaC rthe urrv iaceaehesia eae ceae Microteaca aetes Naenn Caryophyllaceae o iace Caryophyllale Santalales Caryophyllales Santalales Caryop Santalales 140 CATALOGUE OF USEFUL PLANTS OF COLOMBIA Sax aga es Magno ds CHAPTER 10 Monocots Monoco s Gymnospe ms Doryanthaceae Doryanthaceae Iridaceae Iridaceae Xeronemataceae Xeronemataceae Asphodelaceae Asphodelaceae Xanthorrhoeaceae rrhoeaceae ae Xantho Amaryllidace ae Amaryllidace Asparagaceae eae Asparagaceae eae Dasypogonac Dasypogonac Arecaceaee Doryanthaceae Arecaceaee linacea cea Commeana ae e c a d i r I lina ceae Commeana ceae Hangu raceae Xeronemataceae Hangu raceae Philyd ceae Asphodelaceae Philyd ceae Doryanthaceae emodora ae eae ora rrhoeac ce Ha Xantho ria Iridaceae ceaee Haemod Pontede iaceae Amaryllidaceae ntederia ea Xeronemataceae e Po iac Helicon aceaae e Asparagaceae Helicon Musia Asphodelaceae aceaae eae ce e Musia Dasypogonac ce e ea Xanthorrhoeaceae Strelitz Arecaceaee ae ea Strelitz Lowiacceae Amaryllidace linacea e Lowiacceae Doryanthaceae Commeana Canna cea Asparagaceae ceaee Canna Iridaceae antaac Hangu raceae ea taceaee ypogonaceae Marib er ce Philyd ceae ae Xeronemataceae Das Arecaceae Maran acea ng er ora ae ta ib Zi ce ae Asphodelaceae linaceaee Cosha ceae Haemod ZingC riaceea osta ceae Commeana cea Typeliaceaaee Xanthorrhoeaceae Pontelicde iac ee ae Hangu raceae ae Typeha ea He on ce Amaryllidace liace ae Philyd ceae Brom Doryanthaceae Musacceae atea aee ora Asparagaceae Brom teace ae Rap yridace Iridaceae eae Haemod riaceae relitzia eae apa ace e cea St R a X e la Dasypogonac Pontelicde Lowiacceae Xeronemataceae Xyraidulaceeae iaceaee cau cea Arecaceaee c c ae He on Erioayacaiaceaaee aceaae Canna linacea e ceaee Asphodelace Musia Erioayacaiaceaaee M urn ce e Commeana antaac ce e cea ea Xanthorrhoeaceae M urn ce e Marib Th unca cea e er ce Hangu raceae ea ae Strelitz llidaceae Th unca cea e J era ea ZingC Philyd ceae Lowiacceae osta ceae Amary J era ea ragaceae ora Cyprthriaacceaee ae Canna ae Aspa eae Haemod riaceae Cyprthriacceaee Typha antace e e ypogonac liace Das na id ea ea a a e a p eae A Pontelicde Marib cac ce iaceaee le Are nac eae eracce ae Anoalepidnaceeae Brom e atea aee He on ntro tio ac ae aceaae ZingC melinacea e osta ceae Musia Rap yridace a ntr tio ac ae Ce Resellarioaceeae ce e X aulaceeaCom eHanguanacea Ce Resellarioaceeae ea Typha ceae eae lag P ac ae Strelitz c c g rac e c e F P lia ilyd a e a ille ce e Lowiacceae ae ea Erioayacaiaceaaee Phod Fla inv olea cea e Bromateace ae ville ac ae M rn e eem orace a o e a ae Canna le e J ceaee Rap yridace ae ioc yll ce e a ntederiaceea Joinioco yllacceaae ThuuncacceHa antaac e ea X aulaceeae de ph a ea e Marib J era ePo aee liconiac eae er ce ae de ph a e e Eceratoupteleeracceaae c c ac ae Eceratoupteleeracceaae ZingC CyprthriaacceaHe Erioayacaiaceaaee osta ceae Musia C E pavbalaace ae ce e aee re M urn ce e C E pavbalaace ae litz ac ae Padiza ster aceeaee ea AnoalepidnaceeaSt Typeha Th unca cea e liace ae Padiza ster aceeaee r m c a J era ea ntr tio ac ae Lowi ceae r m c a La caeaper ridalaceeaee Bromateace ae Ce Resellari ace aeCanna ceae yp ac ae La ea er a ce e a C id o a p Rap yridace ae r g c P e e la rthri ace ae Cir eniserbencubiacceaae ace aeearantaaceae X lace e Fla Cir enis rbe cu iac ea e M B u a a e e ille ace M Ana pid ce e a iber ceae e n v c cau cea le a b n e n c a ng M o u le S e a e B ta ona cea e e Erioayacaiaceaaee boanaaceeae e Ra Joinioco yllaccZieaaC an Sa on ac ea e entr esti aria cea e e os ceae M C b e c R R n a rn e e ph a e Teypha a a d e ceae lum lat te ac ea e gell Poaaceaae Thuuncacceaee Ec rato ptele rac eaam lum lat te ac ea e e e elia aee Fla Ne P Prondr xac cea ae e J era ea a ville ac ae eaapeateaace Ne P Prondr xac cea ae e Ce Eu pavebalacBcro de Bu na ce a e Cyp riac eae oin ole ace ae e u a e ae e id ce Padiza steraaRceeXayr ho am eraiace ea ae od B mn rac cea ae e e lace e c h n oc e a e ia ce a ch ar ea erm ac oecaauecaceaae L n h o e c a a Tr yrot un illeniscaiace ea ae a E e iaceeae lariM Tr yrot un illeniscaiace ea ae Circ nisprberid caeyeuarn cu ia M G D id n iac e ae e c e c e e G c e o r D e g M achJceuanacea ceaae rid on gia ce ea e M Be nun ab nT PePaeltin lidallac ceaeae e e PePaeltin lidallac ceaeae Ra S boanaCaycepeeara A me hy lla c riaaecceaee A me hy lla c y ia ae ea lum lat te acrtchpeid a y ia a m idipiph or a ea aec ae Ne P PronAdnroxle m idipiph or e ntru aticoneaacee ae Ha ercphn nom oCde BRmenserallcacrieoaaaceeeeae Ha ercphn nom C a y a ge P e c e ch C ia y a la D C n h o ailclecaeaacaeeaeae D C Tr yrot Fun illenin iscvncia oleceacae eae o M G D Jrid Doryanthaceae ioo giayll caecceeaeaaeee Doryanthaceae aeetoinphlitedale idcaecE PeeceaPde a acce eeaaee iIdra Irm cc ea e eAEltm eea taca era upe hayvllellraala e aC a ata ornoenm XeXreA c ecclae eaemaPidaippizipahby oteriaraaceeaae ecae doedla Aspshpohrrhoeac eae a rcardn ams rm ac ae Xantho rrhoeaceae Hae Xantho eL pchaneo pe rid ce ae Amaryllidace ae llidace C y Amary ir a eae ragac nis be ula ce ae Aspa eae ragac Aspa eaeDCMCe Ber uncabiaaceeaee onac eae ypog n S n c a Das onac ypog eae Das cac Are eae Ra mbotanaaceeae e cac Are cea lina e ee me cea lu la te ac ea e lina Com me cea e Com ana cea ngu Ne P Prondr xac cea ae e ana eae ngu rac HaHa eae ilyd rac de Bu na ce a e ae ilyd ce PhPh ae ora ce ho am eraiace ea ae ora em odod em aeae oc deria cece HaHa th n n ac e ae r e ria n nte de o T e nte eaea PoPo yr u ille c c e ae iaciac ea onon ee liclic ac M G D ridisoniagiac ce eae e ea HeHe acia usus ce MM aeaee Pe aeltin lidallac ceaeae iace litzlitz e P rere eaea StSt A me hy lla c acac wiwi aeae LoLo y ia cece a nana Can aeaee m idipiph or Can cece tata ea e anan ac ea arar Ha ercphn nom er MM ac ib aeae iber C a y cece ae ngng os tata ZiZi C ae D C e Cos cece haha Typ aeaae liace Typ ce e aeae elia ace mm Bro ace ate Bro aeae aapte ace yraiduce caee RaRp yr X Xoaid alaclae eeae ric cauycaaccaecacaeeaee a rn EriEoa iae u iac aecaeeaae MMTyuhrn Th ucnacc caee e e a ra c e e e JuJnyepra caecacaeeaee CyCnprtahrtrihariaid cae caeea AnAa pleidp ncae caeeaee o a o le tr n ti e netrno setisoeallriaaricoacaecaeeaeae e CeC ReRla gegll PoPailalecaecacaeeaee FlaF vle cae ea ovinilocleoaleyallcaecaceaeeaee JoJin yhll alecaeecaeeaae ioecio ptohp le dcedto te cra ee EcE raera putep evrae claeca eae CeC EuEpaapva ablaa craecaeaeeaee a b c a PaPdiz daiz srte caeaecaeeaeae rar esatemram acclaecaeaeeaee LaLciraceaapisepr erid erid lau ec ea CirCeMneisneBrbeernbucnucbaiabciacaecaeaeeaee M B naun aS noan caec aeea e S bob aan caec eea e RaR ant ate aeca eea e lumtla a luem la P otreodnradcar xcaececaeeaeae NeN P PrP ne xu ca ae e doedBuB nmanacraececaeeaeeae e ma r e cia a ea hcoh ocro otrhoathnunnennleianaccaececaeaeeae e Tr T il c c ea y ia le u is r G il y is id ian ciaec ae eae MM G D D rider oenogiag dceaceace eae e PePPaPealtAinltinlidealillaycllaceaaceeaceea A maemhyh llayll cia a ip y h iar mamidrcipidipnhip omr o HaHeCrcepahpnhnoym no C aD yC D C aga es laaecaecaeeae TeITxceioocploirhpia ilhaie IxHypoxida ioliriacecaeeae ceae Hypoxida Ate stliealciacceae AsLanar eaeeae iaceae Lanar iaceae Bland fordiaceae Bland fordia Bory aceaeceae Bory e Orcacea hida cea Orc e hida cea Lilia cea Lilia e e Smcea ilaceaceae Sm Philac ilesace Ph iacae iles eae iac Rip og Rip oneae og Als onem acac tro Als eaea e eea tro eri Co em ac lch eri Tecophilaeaceae Co ac ica lch ea ce Pe ee ica ae te Pe rmce anae Ixioliriaceae Mte niaceae elrm anan MCa nia elan thia ce mth aeae Ca Hypoxidaceae pyia cece Cm ae orpy nene siac at ea CC oryc mm siac at ea Asteliaceae e acac CP ea eae e lant ycan e la ha nt ce PS antedadaha Lanariacea nace aeae ST mna teriu cece mrid aeae onon Blandford e Triu acac Verid ac eaea ea lloac VD ee ello Boryacea iaceae iosc ceeaee ziaziaea Dio usc reaace BuB rmo oce e Orchidaceeae artahnarenance ae NPaNrtrm iace a n e e Lil e ia c h iac Petr troescia iacce aeae ym os avc ea CyCm Smilaeaece upopdoadvoiacciaeecaeeaee RR ae PoPusopspid Phile ca ia iaoceaeecaeaceeae oid aceae PP taomnocneia aec ae Ripogsion ZoZosotasm ea te goiagceeto ate MJaM eeto aneae unradora A eae c ls u u tr c JA n a nn iaeca eac cea oemac er ASpunpcoancgdaia eeae eae e gin C ia c ol ce e ch o S chn oin g caaceea Pete icaceae ae B Bcuheueougcehtoeato enae e mc a H Huyto to Melanrmanniace roa hazceznearicaecea A Aylidsrodm thiace ae T Tliosm mcahcacheaarieriaaececaaeeaee Cam ta AArAorfiaecfildealdtaiata ri e a c p ta e c ynem A acc ea cceeaeacee eae Tecophilaeaceae Co Ixioliriaceae ec ae a LaLcaouraroeraceia Cy rsiacea ataceae ue ac e eae e e Hypoxidaceae HH M Meor rancaenceeaaeae Asteliaceae Pa clantha e A Aothennaim ndiaadeia e Lanariaceae G Gtho nim Stenmdanaceceae Blandfordia S Soip emroosroiascpeiacceeaceae Boryaceae ceae rptee ear aee T e o CAaCipaalymaorrute c o p h i l T n ri Ixriiod e aaaeceaeceaaaee A n ru n gr m Orchida liriace E Enunpolyncoancannacagcaecamcaetaatac Ve uHypoxida e aceceae Liliaceaeceae H Huim o a natch theaa aeea aece ea Smilacace Ao stzeia liaceaeae D Dim DiollsLanar e po nm ea ce e ae e ae ceae Philesiac M Me ga amn caetia iaceae aecace aeae c eae B MAMaygraggenenntaeatatia o Rip n Bland e u re og fordia e on A yr r Bory ceae o nria aece Alstroemaceae d dcrea a TecophilaeaN acea L Lra risis nisot li eliria a ceaae m n eriacea a Colchica aacacceercaecaeaceeae n Orc ASsAascatiscttoototicloaic r IxioliriaceaP e ia cea c a lo S ceae e echida Pete ce e Lilia etrothSm Hypoxidaceae cea P Pa aau orar ridc ccheiaaeeaaeeaee eae e iaace Mel rmannia ce eaeaeae AsteliaceaC H Hipyedipureurrarucreidceaahaia s ilac e Ca anthiace ceae yd rn raa aac ce cecaeceea Tecophilaeaceae Pha via Lanariaceae iacc RuypmoRip eae e Cormpynem ae no aocr ccee e a ae eae e Ixioliriaceae d iles Blandfordia ogonac aee Cyc siacea ataceae ra eaac aaee e Hypoxidaceae pia Also Peoceae Boryacea eaericeea trocem ce ea Pan lantha e Asteliaceae s Coclch ac OrchidaP e id ica cea ae e Stemdanaceceae Lanariaceae eceniaaeeaeaee Liliaceae otea oPe ntermaan Triu onac ae Blandfordia SmilacZ anc ce oae Ve ridaceaeae ace the oMgelmia stemCCa Boryaceae ceae iaa ceeae ae PhilesM ziace e py Diollo e iac ne eae Orc r hida Ripogon au aCcycoresiactoeanm at Bu score ae Liliaceaeceae AlstroJ uac ea aceae Tecophilaeaceae N 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ADVerlloazicaac Pan lantha e e P ae Philesiac eae Ripogon H uto uch ton ae aia ZPotaa Stemdanaceceae eae Ripogon LNBaauiocrmoscaronareece aceae Alstroem A ydroma ze ace e temeoongiae caeaeaee ae aaeea c o onac ae Triurid aceae Alstroem eriacea Tolisma chaceaeriaceaae u eae rthre nciaece eMausa Colchica H e ra to V P ac ce eri e e Colch acea A fi e ta ri ceae Pete D llozia eae MCeertrosaacciace aAaJeunc ndiaceae nace A rac eld ce tac Pete icaceae e Mel rmanniace Buioscoreceae L co ea iace ae eae Mel rmanniace A RouypmnoiaadoncvdeiaeacaeaBaeSeeecphoenoaggeinacceeae ae th ce ae Ca an N rman ace H au rac e ae th ce ae ae py iam Ca an cHeauto uc ton ae Corm GoPtPhooetasnidpim eaeia ae Pearthe niaceae py iam M er rac ea oncia siacne e m h Corm a Cycla eae ataceae siacne A o na ea e C trosacviaceaeae c A cydro a zeri ce os rmoogia Cycla eae ataceae Si MZm nthace G th nim nd e iace Panda Ruypmodo p nthace au te ora spetoceaeTareocalifiseeemaatacehacrietaae aceaaee C Panda ae Sipomeros iaceiace a c J S na p P e a e te u ia ce a mon o p C Stem nace ae acealdia ce c An aSlAyponrcuanngdriniatecceegaaLeeAnArcm Triurid Pootasidonceaeaceae aceaae ae An alyarunrteg ermae e on T o riu ia ac V n Z e ac c m Eu nBoucheauogeaacceHaaeeacuraraaecateee ceaaee eae c e zi eae cea E a Velloridaceaeae Maosteraogeto Diollo e Himuponon anthaceaceaatac acea to a n H a zi e e to e M c sc r D c H u n p a e a J nmaa htzheriA aocane nanaeeaeece D Bu ore e n ce n e eae ydro Buioscoreceae D im Ao M e a m ce ac e A unc dia ae aceae N rman ace c aathce e im ae dia a M a ge nt at ae eae N rman ace e Ta lism m chc eaSG S pon agin ceae Pearthe niaceae A y g n an iac rita Pearthe niaceaeM g Ar onfieldaatat cae aeipceeoacmeeoraraoteespiaeceaceeae e B che oge ace C tros ciace ae L r ris no er d ea iaciaeAC C tros ciace aMe ag Aecoaceta H uto uch ton ae anealy A a ist tic lia iac ra e R ymo aviac ae aruen eg rm A ydroma ze ace reacaend R ymo aviac aAe y HLnaun S s ct olo a c e ce auc P uppiadoce eae ep neoancan aceace ata P a a or c c ea ae ae r eHeEim Tolisma chaceaeriaceaae PousppiadcoceaLceaeris ris r Pootasidonceaeaceae tha a ae ce Mtoeo rnlariaaceieaaD H ipeururaceida hiaeae e o a fi a A ri e id e a e ta e P e e m a a c ae ia ta Z yd r ra a ce ce m e ce toAthic A rac ld ce c Z otamonia A G enim M oste oge ceae no ac ce e ae ae a iandciaMaeMecaeegaeacntea atiaeaecea L co ea iace ae eae M oste ogeSceasea toS olo J aun race ton ra ea ae H au rac e ae J aun raceP toanu r ACaripidamr corrhotesAcpLeecaArearaiscyeaterisaegteniconlieaerianacderacceae e A unc dia ae aceae ce e M er rac ea e aceaa A unc diaH aip un Sia o S pon agin ceae gsaa cmtoe A o na ea e ae nnelyca Euec u artaloc acecea eae eae Scponoaginacyeae e rH a a c Bucheuogetoacea c Gothenimiandia e a c o c P r n r p an t eip eu eea idc h a e B he ge ced r D im ro c H to ch n e S H uto uch ton ane Ma A ydroma ze ace C ip mo sp ce ea ce aacotmaaetiacHeyhadaenaecreeaaraeucraaacceeeaeaceeaiaaeceae A ydroma ze aceo A a aru rte er ae e T lism ch cea riac ae aery aggeeanne aa e e o e e T lism ch cea riac AM E n lyc n ga m A ofi ata ari e eae ris no eer nde cea ra a ae A ofi ata ari e Leare a H u no an ac ce ata A rac eld ce tac D im po na th ea ae cea A rac eld ce tac A ac isto ctice lia iac rac e ce e L co ea iace ae eae e M e a m ce ac e La coraeaeiaceaaePSeaasear tor loc aacecea eae eae ae Heauraracee ae M aggennta atia ae eae u id H c e u a a M r c ip H e ru ce a hiaeae e n c A y na ea e Moernraceeae Athonim yd r ra a ce ce La ris ristnoli eria dra eae nd e a a A a n G n a t ic c A no c ce e ae ae G th im d e S o ero ia iac S s ct olo a c e ce ra ea ae S o ero ia iac P a a or c c ea ae ae C ip mo sp ce ea ce e C ip mo sp ce ea H ipeururaceida hiaeae e A a aru rte er ae e yd r ra a ce ce ae A a aru rte er ae e E n lyc n ga m E n lyc n ga m no ac ce e ae ae H u no an ac ce ata H u no an ac ce ata D im po na th ea ae cea ra ea ae De im pomnac tha eae ae cea e M eg an ma ceaace e ce e a c e e M t a e a M t e a ia g e n 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Brassicriac e la a eae Limnanthaceae Ca Hy ost Juguarinraceaae Krameylla ceae d Koeberliniaceae Caosdendtulaceeae Zygophillajaceae Po Bataceae Qu baceae Tic Beylleaacceaaee Salvadoraceae Fa ceae ph riari ace ae o na Emblingiaceae ria is Su alaceae An Coocarpthaceeaee lyg saceae Pentadiplandraceae c Po n n a ry a ita e Tovariaceae Ro aceae ae CoApodcurb elacceaee Resedaceeae eae Barbey Cu tramtisca ceaae gnac ae onac e Gyrostempara Elaeamnaceeae cea ce e Te Da gonia Cap ya ea e Rha mac eae ae Cleomace Be botrstracceaae Dirach aceae a e e o Ulmbac ceaaee Brassicriac a eae pid ela Huaracceaae a n Krameyllaceae Le C ae Can Morace n lida ce ae ce n ph e a e go o ea Zy jac ae C Oxa oniaaceeaee Urticgaceeaae Quillaba n rp c a ofa ac e Fa ce ae Cuoca elliataceeaee naceea Noth Fagcaceaae e Suriaal e e ri ac ae un lo c ea e My ndacceae Ela Br pha ingiadac cea ae ce Polyg e la a e Rosaaceae Juguarinraceaae Ce IrvPan onalacecea ae eae Barbey h y a e e Caosdendtulaceeae gnac ae lop rox or ac ea ae Elaeamnaceeae ic Be lleacceaee T ha no th ph n c e e y a a R mac eae te Ery izo Ochnetiasiac ceaeae e ophoriari aceeae ac e C is Dirach lm a h U bace e n lu lla c ea An C ocarpthac eaee R a c na Bo C phy rica ac ryn an ita ea ae Can Morace lo pe em CoApodcurb elacceaee ace e Urticgaceeaae Ca Hy ost Cu tramtisca ceaae ofaagac ae ia d e e e a th n c T a o D go ya ce e F cace ae N Po Be botrstra ceaae Myrindacceeae a e e o e la a pid Cela Huaracceaae Jugsuarinraceaae e L nnalidaiaceeaee Caodendtulaceeae o C Ox on ac ea e Tic Beylleaacceaaee n rp c a ph ri e Cuoca elliataceeaee iso ria ac ae e un lo c ea e An Coocarpthaceeaee Ela Br pha ingiadac cea ae c ryn an ita ea e CoApodcurb elacceaee Ce IrvPan onalacecea ae Cu tramtisca ceaae ph xy ra ce ae e ce e Te Da gonia olo hro ho na ce ea e ya ea e en ryt op ch tia c ea e Be botrstracceaae Ct E hiz O nne siallac cea ae a o e e lu e R pid ela Huaracceaae Bo C phy rica ac Le C nn lida ce ae lo pe em CoOxa oniaaceeaee Ca Hy ost n rp c a d Cuoca elliataceeaee Po e un lo c ea e Ela Br pha ingiadac cea ae e Ce IrvPan onalacecea ae ph xy ra ce ae e olo hro ho na ce ea e en ryt op ch tia c ea e Ct E hiz O nne siallac cea ae lu e R o y B C ph rica ac lo pe em Ca Hy ost d Po Ca egory o use Santalales Caryophyllales Caryophyllales Anima Enviro Fuels Gene Huma Inverte Mater Medic Social Poison EricalesCampanulids Campanulids Caryophyllales Campanulids Campanulids Caryophyllales Ericales Ericales Campanulids Ericales Ericales Ericales Caryophyllales F GURE 3 Ca ego es o use n he CUPC mapped on a phy ogeny o vascu a p an am es Caryophyllales CATALOGUE OF USEFUL PLANTS OF COLOMB A 141 CHAPTER 10 Medicines 6 ls ria te Ma 2 0 4 0 Po is 2 on 4 s 4 2 6 4 rate 0 6 0 2 Inve 4 es rteb l Us ia Soc 2 Foo d 0 0 4 4 d Ani mal Foo d 2 n Foo 2 Huma 6 0 4 0 en G e es rc u So 2 2 0 6 4 4 Fuels 2 0 FIGURE 4. Chord diagram of categories of use for species in the CUPC. 142 CATALOGUE OF USEFUL PLANTS OF COLOMBIA En e nm o vir l nta es Us CHAPTER 10 A B FIGURE 5. Correlogram of categories of use for species in the CUPC. A cladogram; B Pearson correlation coefficients. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 143 CHAPTER 10 DISCUSSION Before this project, Colombia did not have an updated national list of useful plants. The available lists were not recent, and were focused on specific use categories or geographic areas. The largest species lists always included fewer than 2,600 species [e.g., 2,567 species in IAVH (2014), 2,404 in Bernal et al. (2011), fewer than 2,000 in Pérez Arbeláez (1956), 703 in López & García (2021)], but most of the species were repeated across the publications. By counting species in the CUPC that are not found in any of those important datasets, we estimate that the annotated checklist published in this volume (7,472 spp.) provides records for more than 4,000 useful species that were not included in any of the previous lists. However, the exact number is difficult to estimate. For the first time, the country now has a species list that integrates all use categories. When comparing the number of useful species to the CPC, one in four Colombian plant species has uses reported. Other biodiverse, well-studied countries have a similar richness of useful plants. For instance, for Perú, there are at least 5,000 wild species with known uses (Brack, 1999), and for Ecuador, 5,172 species, which is three of every ten species reported for that country (De la Torre et al., 2008). Unfortunately, only a few countries have comprehensive checklists of useful plants, and vast gaps in ethnobotanical information persist in the region (CámaraLeret et al., 2014), precluding us from making conclusions about these numbers. Most useful species are native, but a significant proportion of them were introduced and are currently naturalised (8.6%), often transforming the natural ecosystems. Several of these species are considered invasive (e.g., various species of Acacia, Eucalyptus, and Pinus), which should be a major concern. Previous studies have reported at least 597 introduced species in Colombia, 42 of which are considered high risk of invasion (Cárdenas-López et al., 2017). The CUPC cites uses for 95% of the naturalised species in Colombia and for 90% of the species with a high risk of invasion. Stunningly, one-third of the species with a high risk of invasion are considered species of high economic interest as well (Cárdenas-López et al., 2017). A review on the possible pathways of introduction reveals both accidental and intentional introductions, such as deliberate release (e.g., ecological restoration), escape from captivity (e.g., ornamental garden plants), contaminants of commodities (e.g., weed seeds), and stowaways on transport vectors (Cárdenas-López et al., 2017). Regardless of their use, biological invasions are deemed to be one of the greatest causes of biodiversity loss (Pyšek et al., 2020), and the use, propagation, and transport of invasive species are prohibited in Colombia (e.g., Law 13 of 1990; Decree 2256 of 1991; Law 599 of 2000). Sixty-five percent of the species (4,844) had uses reported in the WCUP, of which 1,571 species had no reported uses in the country but are used elsewhere. For instance, there are 1,022 species with medicinal uses in other parts of the world (MPNS, 2021), 101 useful species 144 reported in tropical Africa (PROTA, 2020), 97 in Asia (Jansen et al., 1991), 71 in arid and semi-arid lands of the world (RBG Kew, 1999), and 35 in New Guinea (Cámara-Leret & Dennehy, 2019). This significant proportion of neglected or underutilised useful species in Colombia represents an opportunity to further develop the bioeconomy and local practices of sustainable use. Only 18 families contain more than 50% of the useful species in Colombia. This scenario is not surprising, considering that those families are in general speciesrich, abundant, and sources of major crops [e.g., Fabaceae (beans), Asteraceae (sunflowers), Poaceae (cereals), Arecaceae (palms), Rubiaceae (coffee), and Solanaceae (tomatoes, potatoes)]. Fabaceae are the most used plants in Colombia, with hundreds of species used as a source of food and materials distributed across the elevation gradient in all regions. It is also one of the most important families of the lowlands in the neotropics (Gentry, 1988). Asteraceae ranks second in richness of useful species, with its medicinal use being the most important use category by far. The family becomes dominant towards the high-elevation ecosystems (e.g., in páramos) (Rangel-Ch., 2000). Perhaps that is the reason for the common belief that high mountains are the natural pharmacies of Colombians. Poaceae, in the third position, is particularly important as animal food, with most of the current pastures in the country belonging to species introduced in the 20th century, dramatically transforming the native ecosystems. However, many native species of Poaceae still maintain their traditional importance in producing materials (mainly fibres) for handicrafts and construction (Cadena et al., 2007; Linares et al., 2008). Some of the richest genera also include major crops [e.g., Solanum (tomatoes, potatoes), Passiflora (passionfruit)], but many others, with hundreds or tens of useful species, are rarely commercialised. The percentage of endemic useful species is relatively low compared to the overall endemics in the Colombian flora (6.4% vs 32.7%, respectively). This scenario is probably because most of the endemic species are still poorly known and often recently described. They are considered rare or grow in remote areas from which ethnobotanical information is still scanty. Ranks of the most diverse families and genera in the CUPC (Table 2) are relatively consistent with the Colombian flora, listing most of these taxa in the top 25 positions of the CPC, except for Ipomea and Pouteria (in positions 49 and 53 in the CPC, respectively). Some remarkably species-rich genera with several potential ornamental species do not make it to the top positions in the CUPC: three genera of orchids, i.e., Epidendrum (558 spp.), Stelis (450), and Lepanthes (323); and two of aroids, i.e., Anthurium (392) and Philodendron (225) (Table 2). This scenario could be for various reasons, including information gaps on the local use of plants; lack of knowledge on the taxonomic diversity and potential of the flora; the natural history of species (rare species, difficult to find or harvest species, or difficult to propagate species); and current regulations restricting commercialisation of species in CITES lists, including most wild orchids in Colombia (Betancur et al., 2015). CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 10 Results highlight some groups with comparatively low percentages of useful species in the Colombian flora. For instance, Orchidaceae, by far the most diverse family in the country (4,436 spp.) and with the largest genera, ranks in 7th position, with only 195 reported useful species. However, according to CITES, between 1975 and 2012 Colombia exported at least 1,352 species (excluding unidentified or exotic species), mainly to eight countries: Australia, Canada, France, Germany, Japan, Switzerland, United Kingdom and United States of America (Betancur et al., 2015). At the same time, Colombia’s national conservation plan for orchids promotes the artificial propagation and commercialisation of native species and their reintroduction into the natural ecosystems (Betancur et al., 2015). Understanding that most of the Colombian species of orchids are under threat because of landuse change and wild harvesting (Calderón Sáenz, 2007), some recent initiatives in the country are promoting the artificial propagation and sustainable commercialisation of native species of orchids as a way to increase household income and improve the livelihoods of local communities (Castellanos-Castro & Torres-Morales, 2018). The categories of use with more species are medicines (5,108 spp.), human food (3,806), materials (2,363), and environmental uses (2,208) (Figure 2). This scenario is consistent with the top four categories in the World Checklist of Useful Plants (Diazgranados et al., 2020): medicines (26,662 species), materials (13,663), environmental uses (8,983), and human food (7,039). Also, it is not surprising to see medicinal use as the most frequent use for species in Colombia. Numerous studies have focused on this topic (see Chapter 3), and there are guidelines for the sustainable use of prioritised species (Bernal et al., 2011). The report of 5,108 species with medicinal use doubled the 2,404 species previously reported (Bernal et al., 2011). When matching the CPC list with the latest available version of the Medicinal Plant Names Services list (MPNS, 2021), we obtained 3,898 species with medicinal use, with only 75 species not found in any of the other sources we used to compile the WCUP. It is of particular interest to draw attention to the relatively large diversity of plants reported as sources of human food (3,806 spp.), representing 50.1% of the world’s total number of food plants identified to date (7,039) by Ulian et al. (2020), considering that the latter did not include the CUPC database. Also, this represented a significant leap from previous reports, e.g., 703 wild species with edible fruits (López & García, 2021), or 399 species used for human feeding in Colombia (García, 2011). As described in chapter 4, this striking diversity of edible plants evokes calling the country the “hidden food basket of Latin America”. A visual inspection of the distribution of useful taxa across the phylogeny (Figure 3) does not allow a clear identification of patterns, with most families having many types of use (Figure 3). Rosids and Asterids are the clades that concentrate most of the diversity of useful plants (66.9%). Detailed analyses, considering morphological traits, categories of use, ecosystem dominance and richness, are needed to understand these results. Data limitations and potential taxonomic gaps Despite the enormous effort of the team to compile and categorise the available information on plant uses, some caveats are worth highlighting: • There are still numerous sources and records that have not been added to the CUPC because of access limitations, permits, taxonomic problems or limited time. • Ethnobotanical research is still needed in many biodiverse areas of the country. • There is a strong research bias towards certain geographic areas and ecosystems (e.g., Andean Forest) and certain taxonomic groups (e.g., vascular plants). • Interpretation of the categories is not always straightforward. Reports of use often had to be discarded because of misinterpretations (e.g., a wild plant pollinated by bees should not be categorised as “invertebrate food” if it has not been intentionally planted with that purpose). These data limitations can explain some potentially missing taxa. For instance, there are 30 plant families in Colombia (CPC) that have uses reported in the WCUP, but not in the CUPC (e.g., Cardiopteridaceae: 17 spp. in the WCUP / 4 spp. in the CPC; Elatinaceae: 6 / 3; Lentibulariaceae: 17 / 38; Sabiaceae: 17 / 27; and Velloziaceae: 10 / 1). Also, circa 250 genera found in Colombia are reported to include useful species in the WCUP but not in the CUPC (e.g., Dichapetalum: 24 spp. in the WCUP / 10 spp. in the CPC; Homalomena: 16 / 9; Meliosma: 13 / 23; Merremia: 10 / 9; Utricularia: 13 / 32; and Vriesea: 7 / 31). It is possible that with further work, uses within these clades could be added to updated versions of the CUPC. Another limitation is that the database does not include frequency or intensity of use, which is only available for a small proportion of the species. Therefore, we cannot describe how much or how often taxa are used. However, it is quite likely that most species are rarely used, especially by people in urban areas. Last, the database does not reflect the geographic or temporal scales of use. Plant uses vary over time and geographically (Cámara-Leret et al., 2014), depending on cultural changes, society needs and opportunities, local regulations, availability, market behaviour and industrial developments. In the 18th century, frailejones (Espeletia spp.) were largely used to extract turpentine, to construct shelters in the páramos, and for medicinal and social purposes (Cuatrecasas, 2013). Today, those uses of frailejones are not permitted, and people risk jail or a significant fine if frailejones, now with a recognised and protected environmental use, are harvested as a source of materials. The African oil palm (Elaeis guineensis Jacq.) was introduced to Colombia in 1945 by the United Fruit Company (Aguilera-Díaz, 2002), but it only became an economically important species in recent decades. Today, CATALOGUE OF USEFUL PLANTS OF COLOMBIA 145 CHAPTER 10 Colombia is the world’s fourth oil palm producer, first in the Americas, with more than 500,000 ha of oil palm crops in 161 municipalities and 21 departments (Lain & OcampoPeñuela, 2019). Therefore, changes of plant use through time and space are relevant to fully understand the dynamic interaction of societies with nature. CONCLUSIONS This chapter provides an overview of the taxonomic coverage of the CUPC, with notes on the species origin, and mentions of the most diverse taxa, species richness by category of use, phylogenetic coverage, and data limitations. With 7,472 species of useful plants, classified in 2,140 genera and 258 families, Colombia’s rich diversity stands out compared to that of other countries and the WCUP as a natural reservoir of useful species. Also, this reflects, at least in part, this project’s efforts to compile, digitise, and categorise the available information published in this volume and accessible to the community through the ColPlantA portal. Most of the reported species in the CUPC are native (78.6%), but an important proportion is introduced and naturalised (8.6%), some of them with high risk of becoming invasive. The country must pay special attention to those species, as they can be harmful to the native ecosystems and species, yielding negative outcomes in the long run. Circa 21% of useful species had no reports of plant use in the country but are used elsewhere. This significant proportion of neglected or underutilised useful species in Colombia represents an opportunity to further develop the bioeconomy and local practices of sustainable use. The clades Rosids and Asterids are the most useful species-rich large clades. Within them, Fabids and Lamiids concentrate most of the diversity. At the family level, Fabaceae (665 spp.), Asteraceae (418) and Poaceae (339) are the richest, while Solanum (88), Inga (84) and Passiflora (78) are the richest genera. The categories of use with most species are medicines (5,108 spp.), human food (3,806), materials (2,363), and environmental uses (2,208). Using plants to develop this country’s local and national bioeconomy may contribute to ending poverty and hunger, achieving food security, improving nutrition and health, and combating climate change, among other benefits. Therefore, research on economic botany and ethnobotany should continue to complete the inventory of the useful plants of Colombia, filling taxonomic and geographic gaps of information. Also, more studies are needed to investigate temporal and geographic patterns of plant uses and to continue identifying priority species to promote the country’s sustainable development. References Acero L (2005) Plantas útiles de la cuenca del Orinoco. 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CATALOGUE OF USEFUL PLANTS OF COLOMBIA 147 CHAPTER 11 Moon Island in the medium-zone of the Guaviare river. 148 Mateo Fernandez Lucero CATALOGUE OF USEFUL PLANTS OF COLOMBIA Chapter 11 Notes on the geographic distribution of the useful plants of Colombia Mauricio Diazgranados1* & Laura Kor 1 1 Royal Botanic Gardens, Kew. *Corresponding author: m.diazgranados@kew.org Keywords: biogeography, georeferenced records, beta diversity, plant diversity, weighted endemism. ABSTRACT Colombia ranks second in the world for species richness and has high cultural diversity, reflected in the range of ethnic groups and native languages it supports. This rich biocultural diversity has resulted in thousands of documented plant uses, with ecological and social importance. In this chapter, we characterise the geographic distribution of the species listed in the checklist of Useful Plants of Colombia (UPC) by describing distribution patterns at varying biological and geographical scales. In total, the 7,472 plant species with documented uses are represented by 435,230 publicly available georeferenced records. Of the 13 bioregions assessed, the Andean humid forests bioregion has by far the largest number of records, species, genera, and families. The humid forests of the Amazonia and Caribe followed in terms of species richness. Antioquia Department has the largest number of useful species, followed by Cundinamarca and Valle del Cauca, all of which have a remarkable proportion of non-native species. At the municipality level, Solano (Caquetá), Manizales (Caldas), and Leticia (Amazonas) report the highest species richness. However, records are unevenly distributed across both taxonomic groups and geographic areas. Just ten species account for 12.6% of all occurrences, while large and ecologically diverse bioregions such as the Llanos Orientales support very few records concentrated around urban centres. Within all use categories, species richness decreased with elevation. We discuss the potential drivers and implications of these data and knowledge gaps. This characterisation of our current understanding of the distribution of useful plants can be used as a baseline to target future research efforts, supporting plant conservation planning and efforts across Colombia. RESUMEN Colombia ocupa el segundo lugar en el mundo en el número de especies que contiene y posee una alta diversidad cultural, reflejada en la variedad de grupos étnicos y lenguas nativas que preserva. Esta rica diversidad biocultural ha dado lugar a miles de usos de plantas documentados, con importancia tanto ecológica como social. En este capítulo, caracterizamos la distribución geográfica de las especies enumeradas en la lista de verificación de Plantas Útiles de Colombia (UPC) mediante la descripción de patrones de ocurrencia en diversas escalas biológicas y geográficas. En total, las 7.472 especies de plantas con usos documentados están representadas por 435.230 registros georreferenciados disponibles públicamente. De las 13 biorregiones evaluadas, el bosque húmedo de los Andes fue, por mucho, la biorregión con el mayor número de registros, especies, géneros y familias. Le siguieron los bosques húmedos de la Amazonia y el Caribe en términos de riqueza de especies. Antioquia tiene la mayor cantidad de especies útiles, seguido de Cundinamarca y Valle del Cauca, aunque incluyen una proporción notable de especies no nativas. A nivel municipal, Solano (Caquetá), Manizales (Caldas) y Leticia (Amazonas) reportan la mayor riqueza de especies. Sin embargo, los registros están distribuidos de manera desigual entre grupos taxonómicos y áreas geográficas. Solo 10 especies representan el 12,6% de todas las ocurrencias, mientras que las bioregiones grandes y ecológicamente diversas como los Llanos Orientales mantienen muy pocos registros, que en cambio se concentran alrededor de los centros urbanos. Dentro de todas las categorías de uso, la riqueza de especies disminuyó con la elevación. Discutimos los predictores potenciales y las implicaciones de estos datos, así como los vacíos de conocimiento. Esta caracterización de nuestra comprensión actual de la distribución de plantas útiles se puede utilizar como línea de base para orientar los esfuerzos de investigación futuros, apoyando así la planificación y los esfuerzos de conservación de plantas en Colombia. INTRODUCTION The world is facing unprecedented environmental challenges, with consequences for the natural environment and the livelihoods of millions of people (Cardinale et al., 2012). These changes are particularly striking in megadiverse countries such as Colombia. After decades of armed conflict, Colombia is experiencing rapid economic growth but concurrently has one of the world’s highest deforestation rates (Clerici et al., 2020). Large areas of forest, many still poorly explored by scientists, are being transformed into pastures for livestock or severely damaged by mining and other land-use changes. With this loss of biodiversity, nature’s potential contribution to people is also being lost. Therefore, scientists are in a race against time to document Colombia’s biological diversity CATALOGUE OF USEFUL PLANTS OF COLOMBIA 149 CHAPTER 11 A B C FIGURE 1. Biological regionalisation of Colombia. A Bioregions used in this chapter (13 classes); B macro-biomes (12 classes); C general terrestrial ecosystems (93 classes). and its uses before they disappear. One of the fundamental questions in this race is understanding “what grows where”. Colombia is one of the most biodiverse countries on earth (Corzo et al., 2010; Raven et al., 2020), hosting at least 28,947 plant species (24% endemic; >1,000 threatened), belonging to 3,583 genera and 395 families. The country ranks second in the world for its diversity of plants and is the centre of origin of tomatoes, peppers, potatoes, chillies, and many other crops and their wild relatives (Diazgranados et al., 2020a; Khoury et al., 2016). The country’s fungal diversity is also impressive, with at least 7,140 species belonging to 1,763 genera and 448 families. Given that the observed ratio between fungi and plant species in well-studied areas is 9.8:1, Colombia could harbour up to 300,000 species of fungi, representing 9% of the global diversity (Gaya et al., 2021). Colombia’s ecosystem diversity is also very rich, with five continental ecological regions, 13 bioregions, at least 93 general ecosystems (MADS et al., 2017), and 23 climate provinces, from hot-arid to extremely cold and super humid (Figure 1). As part of the Useful Plants and Fungi of Colombia project (UPFC), a multinational team of researchers led by the Royal Botanic Gardens, Kew, with the collaboration of the Instituto de Investigaciones en Recursos Biológicos Alexander von Humboldt, has been compiling, digitising, and georeferencing reported plant uses, as a first step in evaluating the status of knowledge on useful plants in the country. This effort has resulted in this publication, which presents a comprehensive annotated checklist of the plants with reported uses in Colombia, including 7,472 plant species (481 endemics; 150 250 threatened with extinction), belonging to 2,140 genera and 258 families. Our life and well-being rely on the services provided by plants and fungi (Raven, 2021). This fact is true not just for regulating services at the landscape and ecosystem levels, such as soil enrichment, water regulation, and carbon storage, but also through more direct uses such as food, medicines, natural fibres, fuel, building materials, cosmetics, cultural resources, and more (Millennium Ecosystem Assessment, 2005). The preservation and sustainable use of this biodiversity are therefore essential to human wellbeing and economic prosperity. However, a crucial first step in protecting this diversity in situ is understanding its geographical distribution. Useful plant species across bioregions A first comprehensive assessment of the importance of different Colombian bioregions for useful plant species and the quality of available distribution data was undertaken by Bystriakova et al. (2021) during the first phase of the UPFC. The authors assembled a dataset of georeferenced collection localities from online databases of all vascular plants of Colombia (522,257 unique observations of 23,961 species), including useful plant species (197,166 unique distribution records of 3,870 species). The distribution of records per bioregion was investigated using a map with 14 generalised ecosystem types (bioregions). Gaps in the completeness and coverage of records in geographic and environmental space were identified to assess the reliability of species number estimates. CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 11 It was found that the largest numbers of useful plant species occurred in the Andean moist forests bioregion (13,909 species of vascular plants; 2,781 species of useful plants), followed by the Amazon (6,650; 1,612), Pacific (5,168; 1,336), Llanos (2,938; 1,151), and Caribbean moist forests (3,055; 1,166) (Bystriakova et al., 2021). In addition, results showed that medicinal use was the most common useful attribute across all bioregions, followed by materials, environmental uses, and human food (Figure 2). The proportion of well-surveyed 10×10 km grid cells (i.e., with ≥ 25 observation records of useful plants) was less than 50% of the total number of surveyed cells in all bioregions, except for the Andean páramo. The lowest survey coverage was observed in the three dry bioregions: Caribbean deserts and xeric shrublands, Llanos, and Caribbean dry forests. Bystriakova et al. (2021) highlighted that the sampling effort for useful plants of Colombia has been inadequate in the majority of the bioregions. The authors suggested prioritising future research on useful plants in the Amazon and Llanos (Orinoquia) regions, where survey completeness and coverage appeared to be less adequate than that in other regions. Chapter aims and background Building on the biogeographical analyses undertaken by Bystriakova et al. (2021), this chapter aims to describe the geographic distribution of the useful plants of Colombia (UPC) using a significantly larger dataset of compiled species (7,472 versus 3,870) published in this book. Results presented here are primarily descriptive, giving context to other chapters and offering a baseline upon which the checklist can be further improved. For detailed analyses on completeness and coverage of records in geographic and environmental space, see Bystriakova et al. (2021). In this chapter, we aimed to describe the distribution, richness, species density, and other relevant characteristics of useful plant species at a range of biological and geographical scales. Biologically, we assessed unique georeferenced records of herbarium specimens (hereafter occurrences) of all plants sensu lato (Viridiplantae or “green plants”) at the taxonomic level of species, genera, and families. Our geographic analyses considered bioregions (13), departments (32), municipalities (1,122), and grid cells (10×10 km) in recognition of the importance of both ecological and political factors in understanding and managing biodiversity. FIGURE 2. Number of records of useful plants (log-transformed) of Colombia per category of use (x-axis) and bioregion (y-axis). Dendrograms show clusters based on the number of records. From Bystriakova et al. (2021). CATALOGUE OF USEFUL PLANTS OF COLOMBIA 151 CHAPTER 11 Species richness and endemism are two fundamental measures in conservation planning and studies on community ecology and biogeography (Gotelli et al., 2009; Gotelli & Colwell, 2011). Despite this, they remain difficult variables to determine accurately, with various definitions, indices, and models developed for their calculation. Considering the descriptive scope of this chapter, we focused on absolute values of species richness (and the richness of higher taxonomic ranks) based on occurrence data rather than the application of predictive models. Even within this scope, many considerations are involved in measures of species richness, most simply defined as the number of species in an assemblage. While the species-area relationship has long been used to explain plant richness at various scales, other factors such as vegetation types, elevational belts, and their interaction have also been shown to be important determinants (Kreft & Jetz, 2007). Meanwhile, the absolute value of a species’ range is often used as an important determinant in the definition of endemism, an important indicator of concentrations of unique biodiversity. However, this has its limitations, with numerous alternative methods developed based on the relative range sizes of different taxa. For instance, ‘weighted endemism’ calculates the level of endemism of a particular grid cell based on its species richness inversely weighted by the range of each species. This value can be corrected to account for the often observed correlation between species richness and endemism, resulting in the ‘corrected weighted endemism’ index (Crisp et al., 2001). Determining these variables for useful plant species in Colombia, and recognising both their importance and limitations, would significantly contribute to characterising our current understanding of their distribution, thereby supporting conservation efforts and the identification of further research required to fill knowledge and data gaps. Another interesting biogeographical question is how species richness of UPC is distributed along elevation gradients. In Colombian folklore, while páramos are considered “the natural pharmacies” of the Andes, the middle elevations (e.g., Andean and Sub-Andean forest) are referred to as the “food pantries of the cities”. Additionally, Colombia’s highest human population density occurs in the Andean region, at middle and high elevations (DANE, 2015). However, studying the relationship between richness and elevation is a complex subject due to the strong correlation between area and elevation, data limitations (e.g., sampling effort) and various environmental and ecological factors affecting the species distributions. Considering these caveats and acknowledging the need for further detailed studies, we undertook preliminary descriptive analyses on the number of species by use category along elevation gradients. MATERIALS AND METHODS Analyses were based on the full checklist of UPC, published in this book and complied from relevant databases, reports, and scientific literature. The UPC comprises 7,472 species of green plants, classified within 2,140 genera and 258 families. 152 Uses in the UPC were grouped into the ten categories applied in the World Checklist of Useful Plant Species (Diazgranados et al., 2020b), based on the ‘Level 1’ categories of the Economic Botany Data Collection Standard (EBDCS) (Cook, 1995). Native, endemic, cultivated, and naturalised species were identified using the Colombian Catalogue of Plants and Lichens (CCPL) (Bernal et al., 2020), the Plants of the World Online (POWO, 2019), and other reviewed sources. Georeferenced occurrence records were downloaded from GBIF on 15 July 2021 (GBIF Occurrence Download, 2021), using the package rgbif v3.5.2 (Chamberlain & Boettiger, 2017) in R v.4.1.1 (R Core Team, 2020). We cleaned these data with the shinyCCleaner tool (Ondo et al., unpub.) by removing records that were: with coordinates outside Colombia; duplicated; assigned to country’s centroids or institutions; non-terrestrial; rounded to less than one decimal place; with equal or zero longitudes and latitudes; and within 1 km of the capital. Because of the importance of uses associated with some naturalised and cultivated species, which have been incorporated into the cultural practices of communities, we carried out all the analyses on all useful plants regardless of their origin. Datasets for analyses and spatial mapping were prepared using the package dplyr v.1.0.6 (Wickham et al., 2021). Spatial analyses were performed in R v.4.1.1 and ArcGIS Pro 2.8.0 (Esri Inc., 2021). Initially, we used the Colombian Ecosystem map (MADS et al., 2017), containing five continental ecological regions, 12 general biomes, 30 synthesised ecosystems, and 93 general ecosystems. On a scale of 1:100,000, the map was too detailed with too many categories to use in our biome and ecosystem analyses and too coarse with too few categories to be used at the continental region levels. Therefore, we used a map of bioregions produced by the spatial project team, combining the map of Terrestrial Ecoregions of the World (Olson et al., 2001) with the five continental ecological regions in Colombia (Bystriakova et al., 2021). This map resulted in 13 discrete bioregions: deserts and xeric shrublands – Caribe, dry forests – Andes, dry forests – Caribe, dry forests – Llanos, humid forests – Amazonas, humid forests – Andes, humid forests – Caribe, humid forests – Llanos, humid forests – Pacific, mangroves, páramos – Andes, páramos – Caribe, and savannas – Llanos (Figure 1). Bystriakova et al. (2021) created a fourteenth ‘bioregion’ by merging the cells with transformed natural land. However, we did not include this category here for three reasons: 1) cells (10×10 km) of transformed vegetation generally include patches of natural vegetation, ecological corridors, and even nature reserves, where the native useful plants can be still found; 2) even in transformed areas, several local useful plants are cultivated, in orchards, gardens, riverbanks, and along roads; and 3) flora of transformed areas may differ considerably among bioregions because of contrasting environmental conditions; for example, introduced pastures for grazing cattle in the Caribbean region are floristically completely different to those in the high plains of the Andes. In addition, we did not carry out analyses of survey completeness and coverage, which were comprehensively undertaken by Bystriakova et al. (2021) and beyond the descriptive scope of this chapter. CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 11 We used the shapefiles of political divisions for Colombia from the GADM database v. 3.6 (https://gadm.org/). Grid analyses (10×10 km) were used to measure species density. The numbers of occurrences, species, genera, and families by bioregions, departments, and municipalities were computed using the R packages rgdal v1.5–23 (Bivand et al., 2021), raster (Hijmans, 2021), and sp (Bivand et al., 2013). Complete datasets with the results are accessible in Figshare (https:// doi.org/10.6084/m9.figshare.19123157.v1). To analyse broad-scale patterns of useful plant diversity, we calculated estimates of species richness (SR), weighted endemism (WE), and corrected weighted endemism (CWE) using the SDMtoolbox 2.4 (Brown 2014; Brown et al. 2017), a python-based ArcGIS toolbox for spatial analyses. The SR is the sum of unique species per cell, the WE corresponds to the sum of the reciprocal of the total number of cells each species in a grid cell is found in, and the CWE is simply the WE divided by the total species number in a cell. The WE and the CWE emphasise areas with a high proportion of species with restricted ranges, but in the latter, the areas are not necessarily species-rich. The SR, WE, and CWE were estimated using grid cells of 10×10 km. The resulting raster layers were classified using Natural Breaks (Jenks) with ten classes. Studying the relationship between richness and elevation is a complex subject due to the strong correlation between area and elevation. At higher elevations, geographical area is restricted, therefore resulting in lower species richness. This relationship is not usually linear but often unimodal, with a mid-elevation peak (Rahbek, 1995). Species with moderate to large elevational ranges are more likely to overlap at midelevation than at the extremes, resulting in a phenomenon termed the ‘‘mid-domain effect’’ (MDE). MDE is attributed to geometrically constrained stochastic processes affecting the underlying range-size frequency distribution (Colwell, 2008). However, there may be multiple additional factors, such as area availability and environmental variables, that affect the species richness-elevation relationship and result in concentrations around midpoint attractors (Colwell et al., 2016). Recognising the methodological challenges and data limitations, we undertook a simple descriptive analysis of useful species richness using category along elevation to provide an initial picture of the potential patterns. To explore useful plant richness along elevation gradients, we based our analyses on the elevation ranges of each species reported in the Colombian Catalogue of Plants and Lichens, which were classified by experts of each group (Bernal et al., 2020). The total elevation range (i.e., range between the minimum and maximum elevations reported for all species) was divided every 100 m. Each 100 m band was transformed into a bin using the R package linbin (Welty et al., 2015). The number of species occurring in each bin was counted with the R package data.table 1.14.0 (Dowle et al., 2021). Results were visualised with the R package ggplot2 (Wickham et al., 2016). We also compared the empirical curve of species number along the elevation with the computed curve assuming a ‘mid-domain effect’’ (MDE). The empirical richness means of computed richness, standard deviation, and Veech’s statistic (D) (Veech, 2000) were obtained using 1,000 randomisations for 6,581 species with elevation ranges, at 50 sampling sites (from 0 to 5,000 m a.s.l.), using Range Model5Win (Colwell, 2008; Colwell et al., 2016). The null distribution was estimated using the empirical range frequency distribution with random midpoints (box 5 in Range Model5Win). Veech’s statistic measures the mean displacement of the simulated curve from the average random curve, treating MDE as a falsifiable hypothesis (Colwell, 2008). Using a Wilcoxon Test in R, we tested for differences between the empirical richness and the computed richness means. RESULTS Taxonomic coverage and species origin The cleaned database of georeferenced records (occurrences) included 435,230 occurrences, belonging to 6,320 (85%) species, 1,936 (91%) genera, and 248 (96%) families. Occurrences were unevenly distributed across species, with the top ten species accounting for 12.6% of the total number of records (Acacia decurrens with 10,962 occurrences; Eucalyptus globulus 9,578; Pinus radiata 9,353; Gliricidia sepium 6,596; Cedrela angustifolia 4,588; Guazuma ulmifolia 4,484; Quercus humboldtii 3,187; Pinus patula 3,094; and Tapirira guianensis with 2,895 records). In total, 1,161 useful species did not have any adequate georeferenced records, 442 species had only one occurrence, 339 only two occurrences, and 2,172 had ten or fewer occurrences. The list used for the analyses included 6,897 (92%) species with confirmed presence in at least one Colombian department, and 6,581 (89%) species showed elevation range information. The majority of useful plants of Colombia are native (at least 5,881 species, 78.6%), including 481 endemic species to Colombia (8.2% of the native species). There are at least 643 naturalised species and 1,077 cultivated species reported. However, information about species origin is still unclear for 442 species. Bioregions Of the 13 bioregions assessed, the Andes region as a whole (including dry forests, humid forests, and páramos) supported the highest numbers of individual occurrences, species, genera, and families (Figure 3 A–D). The Orinoquia (including the dry forests and savannas in the Llanos Orientales) supported the fewest useful plant records, followed by the Caribbean dry forests and páramos, and the mangroves (Figure 3 A). The humid forests in the Andes held 52.4% of all the occurrences for the country, five times more than the Amazon region, which is the largest by area (Table 1). The Caribbean páramos, located in the Sierra Nevada de Santa Marta, had only 177 occurrences. The humid forests in the Andes supported occurrence records for 82.3% of the useful species in Colombia, representing almost double the species richness of the Amazon region (5,236 compared to 2,848 species) (Table 1). It was also the richest in terms of genera (90.4%) and families (95.2%). The dry forests in the Andes had more genera and families than the Amazon region (1,028 and 186 versus 1,012 and 179, respectively). CATALOGUE OF USEFUL PLANTS OF COLOMBIA 153 CHAPTER 11 A B C D E F G H FIGURE 3. Occurrences, species, genera and family richness of useful plants of Colombia by bioregions A–D and department E–H. 154 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 11 TABLE 1. Occurrences, species, genera, and family richness of useful plants of Colombia per bioregion. Pale green: biomes with the highest overall species richness. Biome region Area (100 km2) Occurrences (%) Species (%) Genera (%) Families (%) Deserts & xeric shrublands – Caribe 273.24 27,006 (6.2) 1,368 (21.6) 746 (38.5) 151 (60.9) Dry forests – Andes 291.34 31,189 (7.2) 2,204 (34.9) 1,028 (53.1) 186 (75) Dry forests – Caribe 248.42 3,660 (0.8) 942 (14.9) 575 (29.7) 141 (56.9) Dry forests – Llanos 245.33 7,539 (1.7) 1,668 (26.4) 815 (42.1) 159 (64.1) Humid forests – Amazonas 3215.99 41,260 (9.5) 2,848 (45.1) 1,012 (52.3) 179 (72.2) Humid forests – Andes 2559.94 228,047 (52.4) 5,236 (82.8) 1,750 (90.4) 236 (95.2) Humid forests – Caribe 862.46 20,494 (4.7) 2,398 (37.9) 1,060 (54.8) 188 (75.8) Humid forests – Llanos 1324.20 16,136 (3.7) 2,026 (32.1) 839 (43.3) 159 (64.1) Humid forests – Pacifico 582.37 16,737 (3.8) 2,145 (33.9) 938 (48.5) 183 (73.8) Mangroves 85.81 7,980 (1.8) 1,175 (18.6) 673 (34.8) 142 (57.3) Paramos – Andes 142.30 17,658 (4.1) 1,326 (21.0) 634 (32.7) 171 (69.0) Paramos – Caribe 12.39 177 (0.1) 93 (1.5) 77 (4.0) 46 (18.5) Savannas – Llanos 1525.94 15,936 (3.7) 1,716 (27.1) 802 (41.4) 164 (66.1) 3500 611 Native Endemic Non-Native 183 3000 9 33 608 297 14 44 11 89 571 264 1 105 10 114 560 350 5 50 10 41 639 12 68 12 50 639 555 29 94 643 21 75 806 53 215 59 126 752 738 65 184 16 145 1000 856 67 131 983 22 42 1132 41 132 63 160 1311 1049 735 78 702 40 72 1426 80 247 98 197 1457 121 284 69 209 1474 1619 1352 35 108 1775 61 206 67 146 1705 125 223 124 82 146 1928 1614 1225 500 1783 1000 172 1500 1913 2000 2673 Species number 398 698 2500 0 FIGURE 4. Species richness of useful plants of Colombia by departments. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 155 CHAPTER 11 A B C D E F G H FIGURE 5. Occurrences, species, genera and family richness of useful plants of Colombia by municipality A–D and grid (10×10 km) E–H. 156 CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 11 TABLE 2. Top 10 municipalities by overall species richness of useful plants (pale green). Name Area (100 km2) Occurrences (%) Species (%) Genera (%) Families (%) Solano (Caquetá) 496.8 9,425 (2.2) 1,455 (23.0) 589 (30.4) 138 (55.6) Manizales (Caldas) 4.4 4,057 (0.9) 1,321 (20.9) 745 (38.5) 162 (65.3) Leticia (Amazonas) 41.7 3,692 (0.8) 1,236 (19.6) 558 (28.8) 134 (54.0) San José del Guaviare (Guav.) 242.1 6,582 (1.5) 1,212 (19.2) 596 (30.8) 131 (52.8) Medellín (Antioquia) 3.6 4,887 (1.1) 1,131 (17.9) 667 (34.5) 157 (63.3) Santafé de Bogotá (Cund.) 16.5 13,400 (3.1) 1,107 (17.5) 608 (31.4) 153 (61.7) Puerto Santander (Amazonas) 90.4 3,867 (0.9) 1,083 (17.1) 476 (24.6) 124 (50.0) Florencia (Caquetá) 20.2 2,390 (0.5) 990 (15.7) 539 (27.8) 135 (54.4) Urrao (Antioquia) 26.4 3,125 (0.7) 978 (15.5) 527 (27.2) 146 (58.9) Santa Marta (D. Esp.) (Mag.) 23.9 2,316 (0.5) 960 (15.2) 584 (30.2) 151 (60.9) Departments Antioquia and Cundinamarca (including Bogotá DC) were the richest departments in terms of individual occurrences, species, genera, and families (Figure 3 E–H), whilst the poorest departments were in the Eastern side of the country. In terms of individual occurrences, Antioquia and Cundinamarca each had over 80,000 records, four times more than Boyacá, in the third position. Eight of the 32 departments had fewer than 5,000 records each. This scenario translated to species richness of 3,616, 2,783, and 2,587 species, respectively, across the top three richest departments. Meanwhile, Sucre, San Andrés y Providencia, and Arauca supported the lowest levels of species richness, with 405, 370, and 355 species, respectively (Figure 4). Antioquia, Cundinamarca, and Bogotá DC had the largest numbers of non-native species, all with more than 600 (Figure 4). A similar pattern was observed for the richness of genera (Figure 3 G), with Antioquia, Cundinamarca, and Santander containing occurrence records for the greatest diversity of genera (1,405, 1,221, and 1,167, respectively), while Guanía, Atlántico, and San Andrés y Providencia had the lowest genus richness (426, 413, and 118 genera). Results at the family level showed the same departments in the top and bottom positions (Figure 3 H). Municipalities Most municipalities displayed relatively low numbers of occurrences compared to Guatavita (Cundinamarca department; 23,883), Jericó (Antioquia; 19,459), Sesquilé (Cundinamarca; 14,856), Santafé de Bogotá (Cundinamarca; 13,400), and Solano (Caquetá; 9,425 occurrences) (Figure 5 A). Excluding Solano, these are relatively small municipalities (the first three being less than 200 km2 in size). Solano, however, is the largest municipality in the country, at over 43,000 km2, making it larger than 23 of the departments. Twenty municipalities had no occurrences, 27 had one or two, and 137 (13%) had fewer than 10. The top 10 richest municipalities in species number are shown in Table 2 and Figure 5 B. Solano, Manizales (Caldas), and Leticia (Amazonas) are the richest in terms of species (1,455, 1,321, and 1,236 species, respectively). Three large cities, Manizales (Caldas), Bogotá DC, and Medellín (Antioquia), had the highest genus (745, 667 and 608, respectively) and family richness (162, 153 and 157, respectively). Grid analyses and endemicity Analyses by grid cells (10×10 km) showed gaps in occurrences in the Amazon and Orinoco (Llanos) regions (south-east of Colombia). Meanwhile, a few cells with particularly high occurrence numbers corresponded to the areas around the cities of Bogotá and Medellín (Figure 5 E). Five speciesrich areas were clearly identified, all associated with the areas of influence of large cities: Bogotá (Cundinamarca), Medellín (Antioquia), the coffee belt (Manizales-PereiraArmenia), Bucaramanga (Santander), and Cali (Valle del Cauca) (Figure 5 F). The same pattern was repeated for the richness of genera (Figure 5 G). The map showing family richness displayed some additional hotspots in Santa Marta (Magdalena), Leticia (Amazonas), Pasto (Nariño), and Florencia (Caquetá) (Figure 5 H). Bogotá and Medellín’s areas supported the highest diversity of species, followed by the coffee belt (CaldasRisaralda-Quindío), Santander, and the Valle del Cauca (Figure 6 A). The weighted endemism map also displayed the same hotspots (Figure 6 B). When correcting by the species number (i.e., corrected weighted endemism), several hotspots of high endemic value emerged in areas such as northern Huila, Caquetá, Vaupés, Vichada, and Casanare (Figure 6 C). CATALOGUE OF USEFUL PLANTS OF COLOMBIA 157 CHAPTER 11 A B C FIGURE 6. Biological regionalisation of Colombia. A Bioregions used in this chapter (13 classes); B macro-biomes (12 classes); C general terrestrial ecosystems (93 classes). Elevation Species richness along elevation gradients showed a strong inverse relationship between the number of useful species and increasing elevation in all categories of use (Figure 7 A). A linear regression of all useful plants along elevation was highly significant (R2 = 0.6; p-value < 0.0001). Analyses by use types showed that plant species with medicinal uses were the most numerous at any elevation and reached the highest elevations. The diversity of plants with environmental use and gene sources was relatively constant between 100 m and 1,500 m before gradually decreasing as elevation increased beyond this point. Very few species with uses other than medicinal occurred above 4,000 m, with only medicinal plants reaching the highest reported elevations of almost 5,000 m (Cerastium arvense, Hypochaeris sessiliflora and Stevia lucida). We observed a weak mid-domain effect (Figure 7 B): for the observed curves, D = 523.44; for 1,000 null curves, mean D = 18.4, range of D = 8.92–40.31. However, we cannot reject the null hypothesis of no differences between the means of the observed distribution and the computed distribution based on 1,000 replications (p-value = 0.65). DISCUSSION Based on publicly available species occurrence data, this chapter presents a geographic description of the current state of knowledge on the useful plants of Colombia. As part of this process, we have characterised useful plant richness at varying biological and geographical scales and highlighted areas of high endemism. These analyses complement and add to that of Bystriakova et al. (2021) and can be used 158 as a baseline upon which an improved understanding of the useful plants of Colombia is built over time. We used a larger dataset of georeferenced occurrences of useful plants (435,230 occurrences, belonging to 6,320 species and 248 families, compared to 197,166 occurrences of 3,870 species and 228 families in Bystriakova et al. (2021)), made possible through the additional work undertaken to increase the comprehensiveness of the UPC. In contrast to Bystriakova et al. (2021), we did not compare the distribution of all vascular plants with that of useful plants or estimate survey completeness and coverage. However, we present counts of species by department and municipality and a first descriptive view on the number of species along elevation. However, there are limitations involved in the data used and analyses undertaken. In this section, we provide context for the patterns observed, discuss the data limitations involved, and outline potential next steps to further our understanding of the useful plants of Colombia. Origin of useful plants in Colombia The checklist of UPC drew on numerous information sources to classify the origin of useful species in Colombia. The proportion of naturalised (8.6%) to native useful species (78.6%) was relatively high, compared to 2.3% and 93.6%, respectively, for all plant species in Colombia. In fact, 643 of the 677 naturalised species have uses reported, suggesting that their utility may be a potential cause for their introduction to natural habitats. A number of these naturalised useful species have become highly invasive both in Colombia and globally (Cárdenas-López et al., 2017; Turbelin et al., 2017), reflected in the higher number of occurrences (e.g., CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 11 A B 2500 Computed Lower 95%CI: Computed Upper 95%CI: Number of species 2000 Empirical Richness: Computed Mean Richness: 1500 1000 0 0 100 200 300 400 500 600 700 800 900 1000 1100 1200 1300 1400 1500 1600 1700 1800 1900 2000 2100 2200 2300 2400 2500 2600 2700 2800 2900 3000 3100 3200 3300 3400 3500 3600 3700 3800 3900 4000 4100 4200 4300 4400 4500 4600 4700 4800 4900 500 Elevation (m) FIGURE 7. Richness of useful plants of Colombia along elevation. A Richness of all plants, all useful plants and species by level 1 category of use. B Empirical richness of useful plants compared to the mid-domain effect (MDE) null-model predictions. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 159 CHAPTER 11 Acacia decurrens, Eucalyptus globulus, Pinus radiata, and P. patula). This result is an important consideration as invasive species are a major driver of biodiversity loss globally, with huge associated livelihood and economic costs (Millennium Ecosystem Assessment, 2005). This scenario poses questions about how best to manage useful plant species, especially introduced and naturalised species, in a way that sustainably maximises their benefits while causing minimal impacts on the wider environment. Distribution of occurrence records Occurrence records were unevenly distributed across species. This result is a common pattern in distribution data of this kind, where the number of singletons (species represented by one individual) and doubletons (species represented by two individuals) often represent a high proportion of species (Gotelli & Colwell, 2011), in our case accounting for 10.4% of UPC. Occurrences were also unevenly distributed across bioregions. Despite the large land area of the llanos Orientales (Orinoquia region), this area’s dry forests and savanna bioregions supported a relatively low number of occurrence records for useful plants. This result is consistent with previous species richness estimates for Colombia (Rangel-Ch, 2015). Still, it is likely to be partly due to under-sampling, as reported by Bystriakova et al. (2021). The estuary nature of large parts of the llanos causes it to be flooded for several months a year, affecting road connectivity. This is an important factor, with a global study on the patterns of plant diversity and floristic knowledge highlighting flooded grasslands and flooded savannas as especially lacking adequate data (Kier et al., 2005). Historical and security factors are also likely to have affected sampling efforts. Traditionally, there has been more focus on sampling in tropical rainforests, such as remote areas of the Amazon region (Redford et al., 1990), than in the Orinoquia region. Meanwhile, some of the areas in the Orinoquia region are well-known corridors of illegal activities (e.g., “Corredor del Sarare-Cubará”, “Corredor de SácamaTame-Saravena”). They have had a long history of violence hampering scientific research. Similar circumstances occur in other poorly studied bioregions, such as the Pacific mangroves and the Caribbean dry forests and páramos. The areas with the highest records remain near large cities, which are likely to have been most intensively surveyed as they are more easily accessible with better supporting infrastructure and safety conditions. This pattern of uneven occurrence distribution is also visible at the department level (Figure 2). Departments such as Guanía and Vichada in the Orinoco region show relatively low numbers of occurrences and species. Their capital cities, Puerto Inírida and Puerto Carreño, are only accessible by plane, with infrequent, relatively costly flights. These areas therefore remain hard to access for biology students and researchers. Meanwhile, despite supporting a wide range of habitats and a large elevation range from sea level to over 5,000 m, the Arauca department shows very poor species diversity based on occurrence records. 160 This result is likely to be because conflict has impeded research in the area for decades. A similar situation is seen in the departments of Norte de Santander, Cesar, Cauca, and Nariño. The latter, bordering Ecuador, has many characteristics which would be expected to make it one of the most biodiverse departments in Colombia. Geographically, it supports a range of biomes, including humid forests of the Amazon and the Pacific, Andean humid and dry forests, isolated areas of páramos, and mangroves, with large areas continuing to be covered by natural vegetation and protected as part of indigenous reserves (MADS et al., 2017). The cultural and ethnic diversity of the department is also notable, with the 2018 census identifying 15.7% of its population as belonging to indigenous groups and 17.8% as Afro-Colombians (DANE, 2018). However, the combination of years of conflict and its distance from the main cities of Colombia has limited the scientific exploration of its rich flora and its uses. Results at the municipality level can be explained by considering their unequal land areas and the influence of urban centres. Very large municipalities or small municipalities near large urban centres tend to have a higher diversity of useful plants. In the second case, several of the richest municipalities by area (e.g., Tenjo, Sasaima, Tabio) are within city regions or urban ecological footprints (Ortega-Montoya & Johari, 2020). Last, human population density is also an important factor. While 29 municipalities in the Amazon and Orinoco regions support fewer than one inhabitant per km2, 25 municipalities, mainly in the Andes, have at least 1 million inhabitants per km2 (DANE, 2018). This scenario produces disproportionate reports of plant uses across the territory. The uneven distribution of sampling effort and the factors contributing to it are common limitations associated with the use of occurrence data, and they have knock-on effects for subsequent analyses and for conservation and management decisions (Gotelli & Colwell, 2011). While accounting for these data limitations through modelling approaches was beyond this chapter’s scope, they should be recognised when interpreting the presented data and statistics on species richness and endemism. Biogeographic characteristics of useful plant richness In terms of overall species, genera, and family richness, the humid forests in the Andes were by far the richest bioregion, followed by the humid forests in the Amazon and Caribe (Table 1). These results coincide with the findings reported by Bystriakova et al. (2021). The regional importance of the Andes in terms of biodiversity, ahead of the Amazon region, has been indicated numerous times. It is possibly due to the higher variety of niches resulting from the area’s diverse climates and soils (Rangel-Ch & Aguilar, 1995). Another reason is the population size of the region. All departments within the Andean region’s combined population are over 26 million (55% of the country’s population) (DANE, 2018). This scenario is important because useful plants can only be recognised where there is interaction with human communities, i.e., when they are used and reported. CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 11 The mangrove and páramos bioregions have a noteworthy diversity of useful plants, despite their very small size compared to other bioregions (Table 1). Mangroves are known to support the production of many goods of economic interest, including wood for firewood and charcoal, timber for construction of dwellings, furniture, boats and fishing gear, tannins for the textile industry, food, and medicinal extracts (Bandaranayake, 1998, García & Polanía, 2007). In the páramos, numerous studies have reported plants’ uses and the ecosystem services they provide (De La Torre et al., 2008, Diazgranados et al., 2021, Duarte & Parra, 2015). Páramos are often referred to as “the pharmacies of the Andes” due to the abundance of a few important medicinal species of Asteraceae, a family supporting many species known to have local medicinal uses (Simpson, 2009). Additionally, páramo grasses are used to thatch roofs of shelters, while Espeletia species were traditionally used as a source of turpentine, for construction, as well as medicinally (Cuatrecasas, 2013). Perhaps the concentration of these useful species in smaller areas and the greater number of studies undertaken due to relatively good accessibility make these bioregions significant for the presence of useful plant species. Based on the analyses of survey completeness and coverage, Bystriakova et al. (2021) suggested prioritising future research on useful plants in the Amazon and Llanos (Orinoquia) regions, for which sampling appeared to be less adequate compared to that in other regions. Our results support this suggestion, with the humid forests in the Amazon, Savannas and Llanos being the least surveyed areas (Figures 5 and 6). These regions have the lowest human population density, with large proportions of their area remaining understudied (Kier et al., 2005). These two factors may explain the results, and further ethnobotanical studies targeting such habitats are likely to unveil a much higher diversity of useful plants in them. The analysis of corrected weighted endemism reveals several hotspots (seen as dark brown cells in Figure 6 C) of high endemic value in areas such as northern Huila, Amazonia (Amazonas, Caquetá, and Vaupés), and Orinoquia (Vichada and Casanare). Those hotspots might coincide with well-surveyed small areas, such as localised studies in nature reserves or around some urban centres. Also, they may reflect the importance of the unique diversity of useful plants of these regions. None of those hotspots can be seen in departments such as Cundinamarca and Antioquia. We undertook a simple descriptive analysis of useful species richness using category along elevation, using equal elevation range size bands (every 100 m). This analysis suggested a general trend of decreasing useful plant species richness with increasing elevation across all categories of use, except for environmental uses and gene sources, where species numbers remain relatively constant between 100 and 1,500 m (Figure 7 A). This result contradicts the common belief that more useful species of certain use types can be found at higher elevations, with the highest species richness found to occur below 1,000 m elevation. In addition, the distribution of the number of useful species along elevation supported a weak MDE. The available data did not provide explanatory variables (climate, ecological conditions, and sampling effort) to address this relationship in detail, which was beyond the scope of this chapter. However, we suggest further exploring the relationship between useful plant richness and elevation, considering sampling across elevation bands of equal area, and including predictors such as climate and human population density. CONCLUSIONS The geographic analyses of the checklist of useful plants presented here reveal both important distributional patterns and gaps in data, which can be used to prioritise further work. The uneven distribution of sampling effort is evident. This scenario is expected in a country like Colombia, where several areas are only now becoming accessible to researchers while others remain difficult to study. At the bioregion level, the humid forests of the Andes were found to support the richest diversity of useful plants. More studies are required in areas such as the Orinoco savannas and forests, for which the information is still limited compared to that for other areas. At the department and municipality levels, size, population density, geolocation, conflict situation, and knowledge availability could explain the results. At the grid level, the observed hotspots of richness can often be explained by the vicinity of large urban centres, highlighting the potential bias in the data available to date. Colombia’s inequality is evident not just in social indicators but also in terms of the distribution of scientific knowledge on the diversity of useful plants. We hope that future research can focus on filling in the identified gaps to decrease current biases in data and knowledge, thereby revealing more representative geographic patterns of useful plants in Colombia. Acknowledgements This work directly contributes to the ‘Useful Plants and Fungi of Colombia’ project, supported by a Professional Development and Engagement grant under the Newton-Caldas Fund partnership. The grant is funded by the UK Department for Business, Energy and Industrial Strategy (BEIS) and Minciencias and delivered by the British Council. LK is supported by a studentship awarded by the Natural Environment Research Council [Grant Number NE/ S007229/1]. References Bandaranayake WM (1998) Traditional and medicinal uses of mangroves. Mangroves Salt Marshes, 2:133-148. Bernal R, Gradstein SR. Celis M (2020) Catálogo de Plantas y Líquenes de Colombia. Bogotá, Universidad Nacional de Colombia. doi: 10.15472/7avdhn Bivand R, Keitt T, Rowlingson B (2021) rgdal: Bindings for the ‘Geospatial’ Data Abstraction Library. R package version 1.5-23. 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BioScience, 51, 933-938. Ondo I, Zizka A, Pironon S, Antonelli A (unpub.) ShinyCCleaner: an interactive app for cleaning species occurrence records. CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 11 Ortega-Montoya CY, Johari A (2020) Urban Ecological Footprints. Sustainable Cities and Communities, 812-824. POWO (2019). Plants of the World Online. Facilitated by the Royal Botanic Gardens, Kew. R Core Team (2020) R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing. Available: https://www.R-project.org/. Rahbek C (1995) The elevational gradient of species richness: a uniform pattern? Ecography, 200-205. Rangel-Ch JO (2015) La biodiversidad de Colombia: significado y distribución regional. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 39, 176–200. Rangel-Ch JO, Aguilar M (1995) Una aproximación sobre la diversidad climática en las regiones naturales de Colombia. Diversidad Biótica I. Instituto de Ciencias Naturales-Universidad Nacional de ColombiaInderena, Bogotá, 25-77. Raven PH (2021) Plants make our existence possible. PLANTS, PEOPLE, PLANET, 2–6. Raven PH, Gereau RE, Phillipson PB, Chatelain C, Jenkins CN, Ulloa Ulloa C (2020) The distribution of biodiversity richness in the tropics. Science Advances, 6, eabc6228. Redford KH, Taber A, Simonetti JA (1990) There is more to biodiversity than the tropical rain forests. Conservation Biology, 4, 328–330. Simpson B (2009) Economic importance of Compositae. In: Funk VA, Susanna A, Stuessy TF, Bayer RJ (eds) Systematics, evolution, and biogeography of Compositae. Washington, D.C., USA: Smithsonian Institution. Turbelin AJ, Malamud BD, Francis RA (2017) Mapping the global state of invasive alien species: patterns of invasion and policy responses. Global Ecology and Biogeography, 26, 78–92. Veech JA (2000) A null model for detecting non-random patterns of species richness along spatial gradients. Ecology, 81, 1143-1149. Welty EZ, Torgersen CE, Brenkman SJ, Duda JJ, Armstrong JB (2015) Multiscale analysis of river networks using the R package linbin. North American Journal of Fisheries Management, 35, 802-809. Wickham H, Chang W, Wickham MH (2016) Package ‘ggplot2’. Create Elegant Data Visualisations Using the Grammar of Graphics. Version, 2, 1-189. Wickham H, François R, Henry L, Müller K (2021) dplyr: A Grammar of Data Manipulation. Available: https://CRAN.R-project.org/package=dplyr. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 163 CHAPTER 12 Specimen of Cattleya trianae Linden & Rchb.f., collected by F. C. Lehmann, in around 1901. It is Colombia’s national flower (also known as Flor de Mayo, or the “Christmas orchid”) and is from one of the identified 29 endemic, threatened species digitised by the UPFC digitisation project officers Priscila dos Reis and Ellie Graves. 164 Digital Collection - RBG Kew CATALOGUE OF USEFUL PLANTS OF COLOMBIA Chapter 12 Annotated checklist of useful plants of Colombia Mauricio Diazgranados1*, Tiziana Cossu1, Laura Kor 1, Benedetta Gori 1, Germán Torres-Morales2, Alejandra Aguilar-Giraldo2, Daniel JiménezPastrana2, Lina Isabel Guevara-Ruiz2, Julia Carretero1, Joaquim de Souza 1, Henry Agudelo3, Fabio Ávila4 & Raquel Negrao1 1 2 3 4 Royal Botanic Gardens, Kew. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt. Instituto de Ciencias Naturales de Colombia, Universidad Nacional de Colombia. Jardín Botánico de Bogotá “José Celestino Mutis”. *Corresponding author: m.diazgranados@kew.org Keywords: edible plants, ethnobotany, food plants, fuel plants, medicinal plants, plant uses ABSTRACT A comprehensive checklist of the useful plants of Colombia is presented in this chapter. This checklist is one of the core outputs of the Useful Plants and Fungi of Colombia (UPFC) project, produced by a multinational team of researchers. The list includes supraspecific taxa, accepted species and authors, common names, species origin (when available), geographic information (habitats, regions, departments, elevation range), conservation status (assessments of the accepted name), and level 1 of the category of use. The taxonomic coverage and statistics are provided in Chapter 10. Notes on the geographic distribution are provided in Chapter 11. Analyses on the conservation status are included in Chapter 2. At the end of the catalogue, indexes of synonyms, families and genera, and common names are also provided. For full details on the species, readers should scan the QR codes or visit the species profiles in http://colplanta.org/, which include morphological descriptions, geographic distribution and maps, synonymy, notes on uses, links to herbarium specimens, images of live plants, illustrations, economic botany items and anatomical laminae, bibliography and additional sources. RESUMEN En este capítulo se presenta una lista de verificación completa de plantas útiles de Colombia. Éste es uno de los resultados centrales del proyecto Plantas y Hongos Útiles de Colombia (UPFC), desarrollado por un equipo multinacional de investigadores. La lista incluye taxones supraespecíficos, especies aceptadas y autores, nombres comunes, origen de las especies (si estaba disponible), información geográfica (hábitats, regiones, departamentos, rango de elevación), estado de conservación (evaluaciones de riesgo de extinction para nombres aceptados), y nivel 1 de la categoría de uso. La cobertura taxonómica y las estadísticas de diversidad son presentadas en el capítulo 10. En el capítulo 11 se presenta una descripción general de la distribución geográfica. Los análisis sobre el estado de conservación de plantas útiles se incluyen en el capítulo 2. Al final del catálogo, también se proporcionan índices de sinónimos, familias y géneros, y nombres comunes. Para obtener detalles completos sobre la especie, los lectores pueden escanear los códigos QR o visitar los perfiles de especies en http://colplanta.org/, que incluyen descripciones morfológicas, distribución geográfica y mapas, sinonimia, notas sobre usos, enlaces a especímenes de herbario, imágenes de plantas vivas, ilustraciones, elementos de botánica económica y láminas anatómicas, bibliografía y fuentes adicionales. INTRODUCTION The elaboration of the checklist of useful plants of Colombia followed five main steps: 1. identification of useful species, digitisation, and compilation of key datasets and sources; 2. taxonomic reconciliation; 3. classification of uses; 4. annotation of species descriptors; and 5. database normalisation. All plant species with a reported current or historical human use in published literature, in Colombia or elsewhere, were included regardless of origin or domestication status. Plants with no reported uses (even if they are informally known to be used or if they could be used) were excluded. Additionally, the checklist did not include infraspecific categories (e.g., subspecies, varieties, forms, and cultivars). 1. Identification, digitisation, and compilation of key datasets and sources A thorough identification process of datasets and sources was carried out over this project (Nov. 2019–Feb. 2022). Initially, we matched the species from the World Checklist CATALOGUE OF USEFUL PLANTS OF COLOMBIA 165 CHAPTER 12 of Useful Plants (WCUP; Diazgranados et al., 2020) with the updated Checklist of Plants of Colombia (CPC), available from ColPlantA (http://colplanta.org/). The 13 large datasets used to prepare the WCUP contributed a significant amount of information (in parentheses: the source and the number of Colombian species with reported uses): 1. Kew’s Economic Botany Collection (EcBot; https://ecbot. science.kew.org/; 1,591 species); 2. European Red List of Medicinal Plants (Allen et al., 2014; 83 species); 3. Germplasm Resources Information Network from the United States Department of Agriculture (GRIN; https:// www.ars-grin.gov; 2,109 species); 4. Medicinal Plant Names Services v.10 (MPNS; http:// mpns.kew.org/; 3,898 species); 5. Useful Plants of New Guinea (Cámara-Leret & Dennehy, 2019; 455 species); 6. Palms of the World Online (http://www.palmweb.org; 121 species); 7. Plant Resources of South-East Asia – PROSEA (Jansen et al., 1991; 812 species); 8. Plant Resources for Tropical Africa (https://www.prota4u. org/database; 922 species); 9. Survey of Economic Plants for Arid and Semi-Arid Lands (SEPASAL; http://apps.kew.org/sepasalweb/sepaweb; 417 species); 10. Useful Plant Project (UPP; Ulian et al., 2017; 253 species); 11. Useful Plants of West Tropical Africa (UPWTA; Burkill, 1995; 688 species); 12. Malaria and Fever in Latin America dataset (Malaria; Milliken, 1997; 626 species); 13. Crop Wild Relatives (CWR; https://www.cwrdiversity.org; Dempewolf et al., 2014; 291 species). We also identified and digitised data from over 1,500 sources reporting plant uses, most from Colombia, such as scientific journal publications, books, technical reports, theses, and databases (the full list is available in the bibliography of ColPlantA). We recognise that some of the species from the checklist do not have reported uses in the country but have uses reported elsewhere, as indicated in Chapter 10. However, we decided that it was important to include them to call attention to the potential and breadth of uses of the Colombian flora. Those taxa can be considered neglected and underutilised species (NUS), and the incorporation of their uses may generate new venues for the improvement of livelihoods of local communities, supporting the further development of the country’s bioeconomy (Hunter et al., 2019; Padulosi et al., 2013; Ulian et al., 2020). 166 2. Taxonomic reconciliation The nomenclature primarily follows the taxonomic backbone of the International Plant Name Index (IPNI, 2021) and the World Checklist of Vascular Plants v.5.0 (WCVP, 2021), also displayed in Plants of the World Online (POWO, 2021). For a few unresolved taxa (<0.1%) such as algae, which are not available in IPNI nor WCVP, we used Tropicos 3.0.2 (TROPICOS, 2021) and AlgaeBase (Guiry & Guiry, 2021). For the higher taxonomy (order level and above), we used the taxonomic backbone of the GBIF species matching tool (GBIF, 2021). The reconciliation of scientific names was done primarily in R version 4.1.0, using the packages plyr version 1.8.5 (Wickham & Wickham, 2020), rgdal version 1.4-8 (Bivand et al., 2015), dplyr version 1.0.6 (Wickham et al., 2021), and doBy version 4.6-3 (Højsgaard et al., 2019). Misspelt species names from the sources were corrected whenever possible, and mismatched names were discarded. Synonymy and homonymy were resolved, and illegitimate and invalid names were excluded. IPNI Life Sciences Identifier (LSID or IPNI ID) were assigned to most names (99.9%), except for 22 accepted names that were not in the current version of IPNI (15 in Tropicos, 3 in AlgaeBase, 1 in USDA, 1 in The Plant List). 3. Classification of uses The categorisation of plant uses followed a simplified version of the level 1 states proposed by Cook (1995) in the Economic Botany Data Collections Standard, as defined in the WCUP (Diazgranados et al., 2020): • Animal Food (AF): Forage and fodder for vertebrate animals only; • Environmental Uses (EU): Examples include intercrops and nurse crops, ornamentals, barrier hedges, shade plants, windbreaks, soil improvers, plants for revegetation and erosion control, wastewater purifiers, indicators of the presence of metals, pollution, or underground water; • Fuels (FU): Wood, charcoal, petroleum substitutes, fuel alcohols, etc. – separated from MATERIALS because of their importance; • Gene Sources (GS): Wild relatives of major crops which may possess traits associated with biotic or abiotic resistance and may be valuable for breeding programs; • Human Food (HF): Food, including beverages, for humans only; • Invertebrate Food (IF): Only plants eaten by invertebrates useful to humans, such as silkworms, lac insects and edible grubs, are covered here. This group does not include bee-pollinated wild species; • Materials (MA): Woods, fibres, cork, cane, tannins, latex, resins, gums, waxes, oils, lipids, etc. and their derived products; • Medicines (ME): Both human and veterinary; • Poisons (PO): Plants that are poisonous to vertebrates and invertebrates, both accidentally and usefully, e.g., for hunting and fishing; CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 12 • Social Uses (SU): Plants used for social purposes, which are not definable as food or medicines, for instance, masticatories, smoking materials, narcotics, hallucinogens and psychoactive drugs, contraceptives and abortifacients, and plants with ritual or religious significance. 4. Annotation of species-related information Common names from the original sources and from the database of common names of Colombian plants (Bernal et al., 2017) were added to the species list. Indications of origin (i.e., endemic, native, introduced, naturalised) were obtained from POWO’s database, from the Catalogue of Plants and Lichens of Colombia (CPLC; Bernal et al., 2019), and from a literature review. We combined the geographic information in the CPLC with the cleaned geographic data in GBIF (used in Chapter 11) to determine the presence in regions and departments. The geographic dataset was also used to extract IUCN habitats. Global conservation status was retrieved from the IUCN Red List portal (IUCN, 2020) and matched to the CPC using the R package dplyr version 1.0.6 (Wickham et al., 2021), taking into account only the assessments related to the accepted name. National assessments were based on data from the IUCN SSC Colombia Plant Specialist Group. Habitat information was extracted from the global map of terrestrial habitat types (Jung et al., 2020), using the R packages sp (Bivand et al., 2013), raster (Hijmans et al., 2013), dplyr (Wickham et al., 2021), and the available georeferenced records (Chapter 11). Last, the CITES status was obtained from the checklist of the CITES Species portal (https://checklist.cites.org/). QR codes linking the ColPlantA species profiles were created for each species from its Permanent URLs using the R package QRcode version 0.1.3 (Teh, 2015). 5. Database normalisation The data were structured, normalised, and formatted according to the standards agreed for this catalogue and space limitations. Please see the list of abbreviations used below. Example of plant record 6174 Miconia formicoheterophylla Michelang. (Melastomataceae) IPNI: 77207843-1 Habit: Shrub, Subshrub Use: HF Origin: Endemic Region: Amazonia, Andean Dept.: AMA, ANT, CAQ, CAU, MET, PUT Elev.: 150–1420 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE Abbreviations and codes used IPNI number: IPNI Life Sciences Identifier (LSID) code. Each name is given a unique LSID code, which can solve misspelt names or homonymy. Therefore, we very much encourage the use of LSID codes, which can be seen at the end of the ColPlantA species profile URLs (e.g., http://powo.science.kew. org/taxon/urn:lsid:ipni.org:names:665697-1?site=colplanta), and can be retrieved using Kew’s Names Reconciliation Service (http://namematch.science.kew.org/). Level 1 categories of use: Animal Food (AF), Environmental Uses (EU), Fuels (FU), Gene Sources (GS), Human Food (HF), Invertebrate Food (IF), Materials (MA), Medicines (ME), Poisons (PO), Social Uses (SU). IUCN Habitat codes: Forest (1), Savanna (2), Shrubland (3), Grassland (4), Wetlands (inland) (5), Rocky Areas (e.g., inland cliffs, mountain peaks) (6), Desert (8), Artificial – Terrestrial (14). Numbers follow the classification of habitats according to the IUCN global map of terrestrial habitat types (Jung et al., 2020). Missing numbers are habitats not found for useful plants of Colombia. Detailed information on the species habitats may be available in the online species profiles and the respective references. Colombian departments: Amazonas (AMA), Antioquia (ANT), Arauca (ARA), Atlántico (ATL), Bogotá D.C. (BOG), Bolívar (BOL), Boyacá (BOY), Caldas (CAL), Caquetá (CAQ), Casanare (CAS), Cauca (CAU), Cesar (CES), Chocó (CHO), Córdoba (COR), Cundinamarca (CUN), Guainía (GUA), Guaviare (GUV), Huila (HUI), La Guajira (LAG), Magdalena (MAG), Meta (MET), Nariño (NAR), Norte de Santander (NSA), Putumayo (PUT), Quindío (QUI), Risaralda (RIS), San Andrés y Providencia (SAP), Santander (SAN), Sucre (SUC), Tolima (TOL), Valle del Cauca (VAC), Vaupés (VAU), and Vichada (VID). Conservation status: International and national conservation status were based on the IUCN Red List (IUCN, 2021) and the “Colombian Plants Specialist Group” CPSG (i.e., the list of national extinction risk assessments carried out by the IUCN Colombian Plant Specialist Group). The assessments were considered only when related to the currently accepted names or when the name was unambiguous (for more detailed information, see Chapter 2). CITES status was presented when available. TAXONOMIC CLASSIFICATION Kingdoms: Plantae (7,471 species) and Chromista (1). Divisions/Phyla: Bryophyta (5 species), Marchantiophyta (2), Ochrophyta (1), Rhodophyta (2), and Tracheophyta (7,462). Classes: Bryopsida (1 species), Cycadopsida (10), Florideophyceae (2), Ginkgoopsida (1), Gnetopsida (5), Lycopodiopsida (32), Magnoliophyta/Angiospermae (7,210), Marchantiopsida (2), Phaeophyceae (1), Pinopsida (26), Polypodiopsida (178), and Sphagnopsida (4). CATALOGUE OF USEFUL PLANTS OF COLOMBIA 167 CHAPTER 12 Orders: Alismatales (160 species), Apiales (82), Aquifoliales (11), Arecales (214), Asparagales (298), Asterales (449), Austrobaileyales (1), Boraginales (58), Brassicales (100), Buxales (3), Canellales (2), Caryophyllales (254), Celastrales (31), Ceratophyllales (1), Chloranthales (15), Commelinales (36), Cornales (6), Crossosomatales (1), Cucurbitales (76), Cyatheales (15), Cycadales (10), Dilleniales (14), Dioscoreales (29), Dipsacales (32), Equisetales (5), Ericales (320), Escalloniales (5), Fabales (688), Fagales (9), Fucales (1), Gentianales (474), Geraniales (15), Ginkgoales (1), Gleicheniales (5), Gnetales (5), Gracilariales (1), Gunnerales (4), Hymenophyllales (6), Icacinales (6), Lamiales (509), Laurales (101), Liliales (18), Lunulariales (1), Lycopodiales (12), Magnoliales (156), Malpighiales (647), Malvales (232), Marchantiales (1), Metteniusales (2), Myrtales (368), Nymphaeales (10), Ophioglossales (4), Osmundales (2), Oxalidales (58), Pandanales (18), Picramniales (8), Pinales (26), Piperales (133), Plocamiales (1), Poales (485), Polypodiales (129), Polytrichales (1), Proteales (16), Psilotales (2), Ranunculales (70), Rosales (291), Salviniales (5), Santalales (42), Sapindales (251), Saxifragales (39), Schizaeales (5), Selaginellales (20), Solanales (250), Sphagnales (4), Vitales (11), Zingiberales (101), and Zygophyllales (9). Families: Acanthaceae (73 species), Achariaceae (17), Achatocarpaceae (1), Actinidiaceae (16), Aizoaceae (11), Alismataceae (14), Alstroemeriaceae (7), Altingiaceae (1), Amaranthaceae (56), Amaryllidaceae (36), Anacardiaceae (31), Annonaceae (105), Apiaceae (34), Apocynaceae (128), Aquifoliaceae (10), Araceae (137), Araliaceae (46), Araucariaceae (4), Arecaceae (214), Aristolochiaceae (24), Asparagaceae (32), Asphodelaceae (9), Aspleniaceae (25), Asteraceae (418), Balanophoraceae (3), Balsaminaceae (5), Basellaceae (4), Bataceae (1), Begoniaceae (27), Berberidaceae (9), Betulaceae (1), Bignoniaceae (82), Bixaceae (6), Bonnetiaceae (1), Boraginaceae (58), Brassicaceae (46), Bromeliaceae (47), Brunelliaceae (5), Burseraceae (58), Buxaceae (3), Cabombaceae (1), Cactaceae (47), Calceolariaceae (3), Calophyllaceae (11), Campanulaceae (29), Cannabaceae (6), Cannaceae (5), Capparaceae (19), Caprifoliaceae (21), Caricaceae (14), Caryocaraceae (10), Caryophyllaceae (30), Casuarinaceae (2), Celastraceae (31), Ceratophyllaceae (1), Chloranthaceae (15), Chrysobalanaceae (50), Cleomaceae (12), Clethraceae (3), Clusiaceae (55), Colchicaceae (1), Combretaceae (18), Commelinaceae (26), Connaraceae (6), Convolvulaceae (77), Coriariaceae (1), Costaceae (20), Crassulaceae (31), Cucurbitaceae (48), Cunoniaceae (9), Cupressaceae (9), Cyatheaceae (15), Cycadaceae (2), Cyclanthaceae (16), Cyperaceae (81), Cyrillaceae (1), Cystopteridaceae (1), Dennstaedtiaceae (6), Dichapetalaceae (5), Dilleniaceae (14), Dioscoreaceae (29), Droseraceae (1), Ebenaceae (8), Elaeocarpaceae (12), Equisetaceae (5), Ericaceae (61), Eriocaulaceae (3), Erythroxylaceae (26), Escalloniaceae (5), Euphorbiaceae (149), Fabaceae (665), Fagaceae (1), Fucaceae (1), Gentianaceae (30), Geraniaceae (15), 168 Gesneriaceae (58), Ginkgoaceae (1), Gleicheniaceae (5), Gnetaceae (5), Goupiaceae (1), Gracilariaceae (1), Grossulariaceae (1), Gunneraceae (4), Haemodoraceae (4), Haloragaceae (1), Hamamelidaceae (1), Heliconiaceae (18), Hernandiaceae (5), Humiriaceae (12), Hydrangeaceae (1), Hydrocharitaceae (4), Hydroleaceae (1), Hymenophyllaceae (6), Hypericaceae (21), Hypoxidaceae (3), Icacinaceae (2), Iridaceae (23), Juglandaceae (2), Juncaceae (9), Krameriaceae (2), Lacistemataceae (2), Lamiaceae (145), Lauraceae (81), Lecythidaceae (45), Lentibulariaceae (1), Liliaceae (1), Linaceae (2), Linderniaceae (7), Loasaceae (5), Loganiaceae (30), Loranthaceae (10), Lunulariaceae (1), Lycopodiaceae (12), Lythraceae (21), Magnoliaceae (13), Malpighiaceae (51), Malvaceae (221), Marantaceae (29), Marcgraviaceae (5), Marchantiaceae (1), Marsileaceae (1), Martyniaceae (2), Melastomataceae (162), Meliaceae (38), Menispermaceae (37), Menyanthaceae (2), Metteniusaceae (7), Microteaceae (1), Molluginaceae (2), Monimiaceae (2), Montiaceae (3), Moraceae (113), Moringaceae (1), Muntingiaceae (2), Musaceae (7), Myricaceae (3), Myristicaceae (38), Myrtaceae (119), Nelumbonaceae (1), Nyctaginaceae (19), Nymphaeaceae (9), Ochnaceae (22), Olacaceae (11), Oleaceae (19), Onagraceae (34), Ophioglossaceae (4), Opiliaceae (2), Orchidaceae (195), Orobanchaceae (10), Osmundaceae (2), Oxalidaceae (26), Pandanaceae (1), Papaveraceae (9), Passifloraceae (88), Paulowniaceae (1), Pedaliaceae (1), Pentaphylacaceae (4), Peraceae (3), Petiveriaceae (8), Phrymaceae (1), Phyllanthaceae (27), Phytolaccaceae (9), Picramniaceae (8), Pinaceae (8), Piperaceae (109), Pittosporaceae (2), Plantaginaceae (43), Platanaceae (1), Plocamiaceae (1), Plumbaginaceae (5), Poaceae (339), Podocarpaceae (5), Podostemaceae (1), Polemoniaceae (3), Polygalaceae (22), Polygonaceae (51), Polypodiaceae (52), Polytrichaceae (1), Pontederiaceae (6), Portulacaceae (4), Potamogetonaceae (4), Primulaceae (47), Proteaceae (14), Psilotaceae (2), Pteridaceae (45), Putranjivaceae (1), Ranunculaceae (15), Rapateaceae (1), Resedaceae (2), Rhamnaceae (18), Rhizophoraceae (4), Rosaceae (79), Rubiaceae (286), Ruppiaceae (1), Rutaceae (34), Sabiaceae (1), Salicaceae (56), Salviniaceae (4), Santalaceae (15), Sapindaceae (78), Sapotaceae (96), Saxifragaceae (4), Schisandraceae (1), Schizaeaceae (5), Schlegeliaceae (4), Schoepfiaceae (1), Scrophulariaceae (9), Selaginellaceae (20), Simaroubaceae (12), Siparunaceae (13), Smilacaceae (9), Solanaceae (171), Sphagnaceae (4), Sphenocleaceae (1), Staphyleaceae (1), Strelitziaceae (4), Styracaceae (10), Surianaceae (1), Symplocaceae (12), Talinaceae (2), Tetrameristaceae (1), Theaceae (3), Thymelaeaceae (3), Triuridaceae (1), Tropaeolaceae (5), Typhaceae (3), Ulmaceae (6), Urticaceae (69), Verbenaceae (50), Viburnaceae (11), Violaceae (31), Vitaceae (11), Vochysiaceae (14), Winteraceae (2), Xyridaceae (2), Zamiaceae (8), Zingiberaceae (18), and Zygophyllaceae (7). CATALOGUE OF USEFUL PLANTS OF COLOMBIA CHAPTER 12 References Allen D, Bilz M, Leaman DJ, Miller RM, Timoshyna A, Window J (2014) European Red List of medicinal plants. Publications Office of the European Union, Luxembourg, 10, 907382. Bernal R, Galeano G, Rodríguez A, Sarmiento H, Gutiérrez M (2017) Nombres Comunes de las Plantas de Colombia. http://www.biovirtual. unal.edu.co/nombrescomunes/. Bernal R, Gradstein S, Celis M (eds) (2019) Catálogo de plantas y líquenes de Colombia. http://catalogoplantasdecolombia.unal.edu.co/. Instituto de Ciencias Naturales, Universidad Nacional de Colombia. Bivand R, Keitt T, Rowlingson B, Pebesma E, Sumner M, Hijmans R, Rouault E, Bivand MR (2015) Package ‘rgdal’. Bindings for the Geospatial Data Abstraction Library. https://cran.r-project.org/web/packages/rgdal/ index.html. Bivand R, Pebesma E, Gómez-Rubio V (2013) Applied spatial data: analysis with R. New York, Springer. Burkill HM (1995) The useful plants of west tropical Africa, Vols. 1-3. 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POWO (2021) Plants of the World Online. Facilitated by the Royal Botanic Gardens, Kew. http://www.plantsoftheworldonline.org/. Teh V (2015) QRcode Generator for R (0.1.3). https://doi.org/10.5281/ zenodo.5040088. TROPICOS (2021) Tropicos.org. Facilitated by the Missouri Botanical Garden. https://www.tropicos.org/. Ulian T, Diazgranados M, Pironon S, Padulosi S, Liu U, Davies L, Howes MJR, Borrell JS, Ondo I, Pérez‐Escobar OA (2020) Unlocking plant resources to support food security and promote sustainable agriculture. Plants, People, Planet, 2, 421-445. Ulian T, Sacandé M, Hudson A, Mattana E (2017) Conservation of indigenous plants to support community livelihoods: the MGU–Useful Plants Project. Journal of Environmental Planning and Management, 60, 668-683. WCVP (2021) World Checklist of Vascular Plants, version 2.0. Facilitated by the Royal Botanic Gardens, Kew (ed.). http://wcvp.science.kew.org/. Wickham H, François R, Henry L, Müller K (2021) dplyr: a Grammar of Data Manipulation. https://CRAN.R-project.org/package=dplyr. Wickham H, Wickham MH (2020) Package ‘plyr’. https://cran.r-project.org/. CATALOGUE OF USEFUL PLANTS OF COLOMBIA 169 Pacific forest. Mar Palanca and Mateo Fernandez Lucer CHECKLIST OF USEFUL PLANTS OF COLOMBIA Marchantiophyta Marchantiopsida Lunulariales 1 Lunularia cruciata (L.) Dumort. ex Lindb. (Lunulariaceae) IPNI: 622149 Use: EU Origin: Endemic, Cultivated Region: Andean Dept.: CUN Elev.: 2650–2650 m a.s.l. Conservation status: (IUCN) NE Marchantiales 2 Marchantia polymorpha L. (Marchantiaceae) IPNI: 622330 Common name: paragüitas Use: ME Origin: Endemic Region: Andean Dept.: BOY, CAU, CES, CUN, MAG, NAR, NSA, SAN Elev.: 1600–4700 m a.s.l. IUCN Habitat code: 1,3,4,6,14 Conservation status: (IUCN) NE Bryophyta Bryopsida Polytrichales 3 Polytrichum juniperinum Hedw. (Polytrichaceae) IPNI: 579099 Use: ME Origin: Endemic Region: Andean Dept.: ANT, ARA, BOY, CAL, CAQ, CAU, CHO, CUN, MAG, MET, NAR, NSA, PUT, RIS, SAN, TOL, VAC Elev.: 950–4680 m a.s.l. IUCN Habitat code: 1,2,3,4,5,6,8,14 Conservation status: (IUCN) NE Sphagnopsida Sphagnales 4 Sphagnum cuspidatum Ehrh. ex Hoffm. (Sphagnaceae) IPNI: 604247 Use: EU, MA, ME Origin: Endemic Region: Andean Dept.: ANT, BOY, CAL, CAU, CUN, HUI, MET, NAR, NSA, SAN, VAC Elev.: 2340-4030 m.a.s.l. IUCN Habitat code: 1,3,4,8,14 Conservation status: (IUCN) NE (CPSG) LC 5 Sphagnum magellanicum Brid. (Sphagnaceae) IPNI: 611262 Use: ME Origin: Endemic Region: Andean Dept.: ANT, ARA, BOY, CAL, CAS, CAU, CES, CHO, CUN, HUI, MAG, MET, NAR, NSA, PUT, QUI, RIS, SAN, TOL, VAC Elev.: 1450-3850 m.a.s.l. IUCN Habitat code: 1,3,4,6,8,14 Conservation status: (IUCN) NE (CPSG) LC 6 Sphagnum meridense (Hampe) Müll. Hal. (Sphagnaceae) IPNI: 611352 Use: MA, ME Origin: Endemic Region: Andean Dept.: ANT, BOY, CAQ, CAU, CUN, HUI, MAG, NAR, PUT, RIS, SAN, TOL Elev.: 1220-3650 m.a.s.l. IUCN Habitat code: 1,3,4,6,8,14 Conservation status: (IUCN) NE (CPSG) LC 7 Sphagnum subsecundum Nees (Sphagnaceae) IPNI: 612658 Use: ME Origin: Endemic Region: Andean Dept.: ANT, CUN, SAN Elev.: 2310-3500 m.a.s.l. IUCN Habitat code: 1,4,14 Conservation status: (IUCN) NE (CPSG) NT Tracheophyta Lycopodiopsida Lycopodiales 8 Huperzia attenuata (Spring) Trevis. (Lycopodiaceae) IPNI: 124842-2 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: BOY, CUN, NAR, RIS Elev.: 2800–3800 m a.s.l. IUCN Habitat code: 1,4,14 Conservation status: (IUCN) NE 9 Huperzia crassa (Humb. & Bonpl. ex Willd.) Rothm. (Lycopodiaceae) IPNI: 124844-2 Common name: cacho de venao, colchón de pobre, piligan Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ARA, BOY, CAL, CHO, MAG, MET, NAR, PUT, RIS, SAN, TOL Elev.: 3350–4700 m a.s.l. IUCN Habitat code: 1,3,4,6,8,14 Conservation status: (IUCN) NE 10 Huperzia dichotoma (Jacq.) Trevis. (Lycopodiaceae) IPNI: 1090207-2 Habit: Herb Use: ME Origin: Native Region: Andean, Pacific Dept.: ANT, CHO Elev.: 0–700 m a.s.l. IUCN Habitat code: 1,3 Conservation status: (IUCN) NE 11 Huperzia saururus (Lam.) Trevis. (Lycopodiaceae) IPNI: 60447820-2 Habit: Herb Use: MA, ME Origin: Native Region: Andean Dept.: CHO, MAG Elev.: 3800–3850 m a.s.l. IUCN Habitat code: 4 Conservation status: (IUCN) NE 12 Lycopodiella alopecuroides (L.) Cranfill (Lycopodiaceae) IPNI: 146929-2 Use: ME IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) NE (CPSG) Potential LC 13 Lycopodiella caroliniana (L.) Pic.Serm. (Lycopodiaceae) IPNI: 146931-2 Habit: Herb Use: SU Origin: Native Region: Amazonia, Andean, Pacific Dept.: AMA, ANT, CAQ, CHO, GUA, NAR, SAN, VAU Elev.: 50–2500 m a.s.l. IUCN Habitat code: 1,4,14 Conservation status: (IUCN) NE 14 Lycopodiella cernua (L.) Pic.Serm. (Lycopodiaceae) IPNI: 1054587-2 Common name: abrecaminos, cacho de venado, caminadera, colchon de pobre, colchón de pobre, gateadera, uña de gato Habit: Herb Use: EU, HF, MA, ME, SU Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, ARA, BOY, CAL, CAQ, CAU, CES, CHO, CUN, GUA, HUI, LAG, MAG, MET, NAR, NSA, QUI, RIS, SAP, SAN, TOL, VAC, VAU, VID Elev.: 50-3000 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC QUI, RIS, SAN, TOL, VAC Elev.: 1000-3700 m.a.s.l. IUCN Habitat code: 1,3,4,8,14 Conservation status: (IUCN) NE (CPSG) Potential LC 19 Lycopodium thyoides Willd. (Lycopodiaceae) IPNI: 147665-2 Common name: cacho de venado, caminadera, gateadera, helecho gateadero Habit: Herb Use: ME Origin: Native Region: Andean, Caribbean Dept.: ANT, BOY, CAL, CAU, CES, CHO, CUN, HUI, MAG, MET, NAR, NSA, PUT, QUI, RIS, SAN, TOL, VAC Elev.: 1700-4300 m.a.s.l. Conservation status: (IUCN) NE (CPSG) Potential LC Selaginellales 20 Selaginella amazonica Spring (Selaginellaceae) IPNI: 231969-2 Habit: Herb Use: ME Origin: Native Region: Amazonia, Pacific Dept.: AMA, CAQ, CAU, GUA, VAU Elev.: 150–250 m a.s.l. IUCN Habitat code: 1,2,3,5,14 Conservation status: (IUCN) NE 21 Selaginella articulata (Kunze) Spring (Selaginellaceae) IPNI: 77186609-1 Habit: Herb Use: ME Origin: Native Region: Amazonia, Pacific Dept.: ANT, CHO, PUT, VAC Elev.: 0–1400 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 22 Selaginella asperula Mart. ex Spring (Selaginellaceae) IPNI: 77186584-1 Common name: busukobenaeji, little tree mano de sapo Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Orinoquia Dept.: AMA, BOG, CAQ, CAU, GUA, GUV, MET, NAR, VAU, VID Elev.: 0–2000 m a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE 15 Lycopodiella pendulina (Hook.) B.Øllg. (Lycopodiaceae) IPNI: 279987-2 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, BOY, CAU, CUN, NAR, PUT Elev.: 2100– 3600 m a.s.l. IUCN Habitat code: 1,3,4,8,14 Conservation status: (IUCN) NE 23 Selaginella convoluta (Arn.) Spring (Selaginellaceae) IPNI: 77142531-1 Habit: Herb Use: ME Origin: Native Region: Andean, Caribbean Dept.: CES, MAG, NSA Elev.: 450–1900 m a.s.l. IUCN Habitat code: 1,2,14 Conservation status: (IUCN) NE 16 Lycopodium clavatum L. (Lycopodiaceae) IPNI: 304307-2 Common name: cacho de venado, cacho de venao, caminadera, colchón de pobre, gateadera, lycopodio Habit: Herb Use: AF, EU, FU, HF, MA, ME, SU Origin: Native Region: Andean, Caribbean Dept.: ANT, BOG, BOY, CAL, CAS, CAU, CES, CUN, HUI, LAG, MAG, NAR, NSA, RIS, SAN, TOL, VAC Elev.: 1050-4200 m.a.s.l. IUCN Habitat code: 1,2,3,4,6,8,14 Conservation status: (IUCN) NE (CPSG) Potential LC 24 Selaginella diffusa (C.Presl) Spring (Selaginellaceae) IPNI: 77186646-1 Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Orinoquia, Pacific Dept.: ANT, BOY, CAL, CAQ, CAU, CES, CHO, CUN, HUI, MAG, MET, NAR, NSA, PUT, QUI, SAN, TOL, VAC Elev.: 200-2800 m.a.s.l. IUCN Habitat code: 1,2,3,14 Conservation status: (IUCN) NE (CPSG) Potential LC 17 Lycopodium complanatum L. (Lycopodiaceae) IPNI: 147262-2 Common name: caminadera Use: ME, PO, SU Dept.: BOG IUCN Habitat code: 1,4,14 Conservation status: 25 Selaginella erythropus (Mart.) Spring (Selaginellaceae) IPNI: 30243959-2 Habit: Herb Use: EU Origin: Native Region: Pacific Dept.: ANT, CAL, CHO, CUN, HUI, NAR, NSA, TOL, VAC Elev.: 150–2000 m a.s.l. IUCN Habitat code: 1,2,3,4,14 Conservation status: (IUCN) NE (IUCN) NE 18 Lycopodium jussiaei Desv. (Lycopodiaceae) IPNI: 147420-2 Common name: cacho de venado Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, BOY, CAL, CAU, CHO, CUN, HUI, MAG, MET, NAR, NSA, PUT, CATALOGUE OF USEFUL PLANTS OF COLOMBIA 26 Selaginella exaltata (Kunze) Spring (Selaginellaceae) IPNI: 77186589-1 Common name: yerba de loro Habit: Herb Use: ME Origin: Native Region: Amazonia, Pacific Dept.: AMA, ANT, CAQ, CAU, CHO, GUV, NAR, PUT, VAC Elev.: 0–400 m 171 CHECKLIST OF USEFUL PLANTS OF COLOMBIA a.s.l. IUCN Habitat code: 1,3,5,14 Conservation status: (IUCN) NE 27 Selaginella flabellata (L.) Spring (Selaginellaceae) IPNI: 30226649-2 Habit: Herb Use: ME Origin: Native Region: Amazonia, Pacific Dept.: AMA, ANT, CAQ, CHO, VAC Elev.: 200–900 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) NE 28 Selaginella flagellata Spring (Selaginellaceae) IPNI: 232097-2 Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Pacific Dept.: AMA, ANT, BOY, CAQ, MAG, MET, QUI, SAN Elev.: 200–1800 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 29 Selaginella geniculata (C.Presl) Spring (Selaginellaceae) IPNI: 77186554-1 Habit: Herb Use: ME Origin: Native Region: Andean, Pacific Dept.: ANT, BOY, CAL, CAQ, CAU, CHO, CUN, HUI, NAR, PUT, QUI, RIS, TOL, VAC Elev.: 0-3400 m.a.s.l. IUCN Habitat code: 1,3,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 30 Selaginella lingulata Spring (Selaginellaceae) IPNI: 232147-2 Common name: doradilla Habit: Herb Use: ME Origin: Native Region: Andean, Pacific Dept.: ANT, BOY, CAU, CHO, CUN, NAR, PUT, RIS, SAN, TOL, VAC Elev.: 50–3500 m a.s.l. IUCN Habitat code: 1,3,5,14 Conservation status: (IUCN) NE 31 Selaginella pallescens (C.Presl) Spring (Selaginellaceae) IPNI: 30016320-2 Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Caribbean Dept.: CES, GUA, LAG, MAG Elev.: 700–1200 m a.s.l. IUCN Habitat code: 1 Conservation status: (IUCN) NE 32 Selaginella parkeri (Hook. & Grev.) Spring (Selaginellaceae) IPNI: 77186636-1 Habit: Herb Use: ME Origin: Native Region: Amazonia Dept.: AMA, CAQ, PUT, VAU Elev.: 100–400 m a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE 33 Selaginella plana (Desv.) Hieron. (Selaginellaceae) IPNI: 77186707-1 Habit: Herb Use: AF, HF, MA, ME Origin: Naturalised Region: Pacific Dept.: NAR, TOL, VAC Elev.: 0–450 m a.s.l. IUCN Habitat code: 14 Conservation status: (IUCN) NE 34 Selaginella rosea Alston (Selaginellaceae) IPNI: 232255-2 Common name: yerba del carpintero Habit: Herb Use: ME Origin: Endemic Region: Andean Dept.: ANT, TOL Elev.: 1600–2500 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 35 Selaginella schultesii Alston ex Crabbe & Jermy (Selaginellaceae) IPNI: 232288-2 Habit: Herb Use: ME Origin: Native Region: Amazonia Dept.: AMA, VAU Elev.: 200–700 m a.s.l. IUCN Habitat code: 1,5 Conservation status: (IUCN) NE 172 36 Selaginella sellowii Hieron. (Selaginellaceae) IPNI: 232292-2 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: BOG, BOY, CUN, HUI, NAR, SAN Elev.: 700– 2700 m a.s.l. Conservation status: (IUCN) NE 37 Selaginella speciosa A.Br. (Selaginellaceae) IPNI: 77186694-1 Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Pacific Dept.: AMA, ANT, CAL, CAQ, CAU, CHO, CUN, PUT, TOL, VAC Elev.: 100–1800 m a.s.l. Conservation status: (IUCN) NE 38 Selaginella stellata Spring (Selaginellaceae) IPNI: 77186626-1 Habit: Herb Use: ME Origin: Native Region: Amazonia Dept.: VAU Elev.: 300–350 m a.s.l. IUCN Habitat code: 1,5 Conservation status: (IUCN) NE 39 Selaginella willdenowii (Desv.) Baker (Selaginellaceae) IPNI: 60471496-2 Habit: Herb Use: EU, HF, ME Origin: Naturalised Region: Andean Dept.: VAC Elev.: 1350–1350 m a.s.l. IUCN Habitat code: 14 Conservation status: (IUCN) NE Polypodiopsida Cyatheales 40 Alsophila cuspidata (Kunze) D.S.Conant (Cyatheaceae) IPNI: 9954-2 Use: HF Origin: Native Region: Amazonia, Andean, Guiana Shield, Pacific Dept.: ANT, CAU, CHO, MET, NSA, PUT, RIS, SAN, VAC Elev.: 50-1600 m.a.s.l. IUCN Habitat code: 1,3,5,14 Conservation status: (IUCN) NE (CPSG) LC 41 Alsophila erinacea (H.Karst.) D.S.Conant (Cyatheaceae) IPNI: 9971-2 Common name: helecho estacoso Use: ME Origin: Native Region: Andean, Pacific Dept.: ANT, BOY, CAU, CHO, CUN, NAR, PUT, SAN, VAC Elev.: 500-2600 m.a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE (CPSG) LC 42 Alsophila firma (Baker) D.S.Conant (Cyatheaceae) IPNI: 9976-2 Use: ME Origin: Native Region: Andean Dept.: NAR Elev.: 13001300 m.a.s.l. IUCN Habitat code: 1 Conservation status: (IUCN) NE (CPSG) LC 43 Cyathea arborea (L.) Sm. (Cyatheaceae) IPNI: 17071820-1 Use: EU Origin: Native Region: Caribbean Dept.: LAG Elev.: 600650 m.a.s.l. IUCN Habitat code: 3 Conservation status: (IUCN) NE (CPSG) VU 44 Cyathea aterrima Domin (Cyatheaceae) IPNI: 73117-2 Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Pacific Dept.: CAQ, CHO, GUV, MAG, MET, SAN, VAC, VAU Elev.: 250-1800 m.a.s.l. IUCN Habitat code: 1,4,14 Conservation status: (IUCN) NE (CPSG) LC CATALOGUE OF USEFUL PLANTS OF COLOMBIA 45 Cyathea bicrenata Liebm. (Cyatheaceae) IPNI: 73125-2 Common name: Palo de la vida, Pelma Use: HF Origin: Native Region: Amazonia, Andean, Pacific Dept.: AMA, ANT, CAL, CHO, HUI, NAR, TOL, VAC Elev.: 0-2500 m.a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE (CPSG) LC 46 Cyathea caracasana (Klotzsch) Domin (Cyatheaceae) IPNI: 73146-2 Common name: helecho, helecho macho, palma boba Use: EU, MA, ME, SU Origin: Native Region: Andean, Guiana Shield, Caribbean, Pacific Dept.: ANT, BOG, BOY, CAL, CAU, CHO, CUN, HUI, LAG, MAG, MET, NAR, NSA, PUT, RIS, SAN, TOL, VAC Elev.: 1200-4200 m.a.s.l. IUCN Habitat code: 1,2,3,4,14 Conservation status: (IUCN) NE (CPSG) Potential LC 47 Cyathea fulva (M.Martens & Galeotti) Fée (Cyatheaceae) IPNI: 17072730-1 Use: MA Origin: Native Region: Andean Dept.: ANT, BOY, CAL, CAU, CUN, NSA, QUI Elev.: 1800–3300 m a.s.l. IUCN Habitat code: 1,2,3,14 Conservation status: (IUCN) NE 48 Cyathea lasiosora Domin (Cyatheaceae) IPNI: 73267-2 Common name: helecho arborescente, tacis, ùjùmedù Use: ME Origin: Native Region: Amazonia, Andean, Orinoquia Dept.: AMA, ANT, CAQ, CAU, MET, NAR, PUT, VAC, VAU Elev.: 100-1800 m.a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE (CPSG) LC 49 Cyathea microdonta (Desv.) Domin (Cyatheaceae) IPNI: 73288-2 Common name: lecho, palma boba, trichi o tricho Use: HF, ME Origin: Native Region: Andean, Orinoquia, Pacific Dept.: ANT, CAQ, CAS, CAU, CHO, CUN, MET, NAR, VAC, VID Elev.: 0-2000 m.a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE (CPSG) LC 50 Cyathea tryonorum (Riba) Lellinger (Cyatheaceae) IPNI: 275500-2 Use: ME Origin: Native Region: Andean Dept.: ANT, BOY, CAU, CUN, SAN, VAC Elev.: 1650-2900 m.a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE (CPSG) LC 51 Dicksonia karsteniana (Klotzsch) T.Moore (Cyatheaceae) IPNI: 1190416-2 Use: ME Origin: Native Conservation status: (IUCN) NE (CPSG) LC 52 Dicksonia sellowiana (C.Presl) Hook. (Cyatheaceae) IPNI: 275823-2 Use: MA, ME Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CAL, CAU, CES, CUN, HUI, MAG, NAR, NSA, PUT, QUI, RIS, SAN, TOL, VAC Elev.: 1500-3800 m.a.s.l. IUCN Habitat code: 1,3,4,5,8,14 Conservation status: (IUCN) NE (CPSG) Potential LC CHECKLIST OF USEFUL PLANTS OF COLOMBIA 53 Lophosoria quadripinnata (J.F.Gmel.) C.Chr. (Cyatheaceae) IPNI: 143624-2 Common name: boba, helecho de árbol, lecho Use: HF, ME Origin: Native Region: Andean, Guiana Shield, Pacific Dept.: ANT, BOG, BOY, CAL, CAQ, CAU, CHO, CUN, HUI, MAG, MET, NAR, NSA, PUT, RIS, SAN, TOL, VAC Elev.: 100-3510 m.a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE (CPSG) LC 61 Gleichenella pectinata (Willd.) Ching (Gleicheniaceae) IPNI: 110733-2 Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, BOY, CAQ, CAU, CES, CHO, CUN, GUV, HUI, MAG, MET, NAR, QUI, SAN, TOL, VAC, VID Elev.: 0-1950 m.a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) NE (CPSG) Potential LC 69 Trichomanes elegans Rich. (Hymenophyllaceae) IPNI: 17229030-1 Common name: loro, navegadora Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Pacific Dept.: AMA, ANT, BOY, CAQ, CAU, CHO, CUN, GUA, MET, NAR, NSA, PUT, RIS, SAN, VAC, VAU Elev.: 0–1500 m a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE 54 Sphaeropteris quindiuensis (H.Karst.) R.M.Tryon (Cyatheaceae) IPNI: 241965-2 Use: ME Origin: Native Region: Andean Dept.: ANT, BOY, CAL, CUN, HUI, NAR, QUI, RIS, SAN, TOL Elev.: 750-3000 m.a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) NE (CPSG) LC 62 Sticherus bifidus (Willd.) Ching (Gleicheniaceae) IPNI: 245596-2 Common name: helecho pata de gallina, helecho pategallina Habit: Herb Use: MA Origin: Native Region: Andean, Pacific Dept.: ANT, BOG, BOY, CAL, CAQ, CAU, CHO, CUN, HUI, MAG, MET, NAR, NSA, PUT, SAN, VAC Elev.: 50-2450 m.a.s.l. IUCN Habitat code: 1,2,3,4,14 Conservation status: (IUCN) NE (CPSG) Potential LC 70 Trichomanes pinnatum Hedw. (Hymenophyllaceae) IPNI: 320804-2 Common name: helecho de pescado, rabo de chucha Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, BOY, CAQ, CAU, CES, CHO, COR, CUN, MET, NAR, NSA, PUT, SAN, TOL, VAC, VAU, VID Elev.: 0-1000 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC Equisetales 55 Equisetum arvense L. (Equisetaceae) IPNI: 300073-2 Use: HF, MA, ME, PO Conservation status: (IUCN) LC 56 Equisetum bogotense Kunth (Equisetaceae) IPNI: 329489-2 Common name: canutillo, cola de caballo, equisetum, herb de conejo, juntalli, tembladera, tembladera pequeña, yerba de chivo Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, ARA, BOG, BOY, CAQ, CAU, CES, CHO, CUN, HUI, MAG, MET, NAR, NSA, PUT, QUI, RIS, SAN, TOL, VAC Elev.: 1500-3800 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 57 Equisetum giganteum L. (Equisetaceae) IPNI: 320302-2 Common name: canutillo, cola de caballo, rabo de mula Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Caribbean Dept.: ANT, BOG, BOY, CES, CHO, CUN, GUA, HUI, NAR, PUT, QUI, TOL, VAC Elev.: 600–3000 m a.s.l. IUCN Habitat code: 1,2,3,14 Conservation status: (IUCN) LC 58 Equisetum hyemale L. (Equisetaceae) IPNI: 92939-2 Use: EU, MA, ME Region: Andean Dept.: BOG Conservation status: (IUCN) LC 59 Equisetum myriochaetum Schltdl. & Cham. (Equisetaceae) IPNI: 77060052-1 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, CHO, CUN, TOL, VAC Elev.: 1400–2200 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) NE Gleicheniales 60 Dicranopteris flexuosa (Schrad.) Underw. (Gleicheniaceae) IPNI: 79839-2 Common name: helecho pategallina Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Pacific Dept.: ANT, BOY, CAQ, CAU, CES, CHO, CUN, GUA, MET, SAN, VAC, VAU Elev.: 100-2600 m.a.s.l. IUCN Habitat code: 1,2,3,14 Conservation status: (IUCN) NE (CPSG) Potential LC 63 Sticherus nudus (Moritz ex Reichardt) Nakai (Gleicheniaceae) IPNI: 245632-2 Habit: Herb Use: MA Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CAU, CUN, HUI, PUT, VAC Elev.: 1700–2500 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 64 Sticherus rubiginosus (Mett.) Nakai (Gleicheniaceae) IPNI: 245647-2 Common name: helecho gallinero, pata de gallina Habit: Herb Use: MA Origin: Native Region: Andean, Pacific Dept.: ANT, BOG, BOY, CAQ, CAU, CHO, CUN, HUI, MAG, NAR, NSA, SAN, TOL, VAC Elev.: 1250-2800 m.a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE (CPSG) Potential LC Hymenophyllales 65 Hymenophyllum fragile (Hedw.) C.V.Morton (Hymenophyllaceae) IPNI: 125766-2 Habit: Herb Use: ME Origin: Native Region: Andean, Pacific Dept.: ANT, BOY, CHO, CUN, MAG, RIS, VAC Elev.: 1450–2800 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 66 Hymenophyllum fucoides (Sw.) Sw. (Hymenophyllaceae) IPNI: 1009149-2 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CAU, CES, CHO, CUN, HUI, MAG, NSA, PUT, RIS, SAN, TOL, VAC Elev.: 400-4300 m.a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) NE (CPSG) Potential LC 67 Trichomanes crispum L. (Hymenophyllaceae) IPNI: 257306-2 Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, BOY, CAU, CHO, CUN, MET, NAR, NSA, SAN, TOL, VAC Elev.: 0–2050 m a.s.l. Conservation status: (IUCN) NE 68 Trichomanes diversifrons (Bory) Mett. (Hymenophyllaceae) IPNI: 17228970-1 Common name: churrusco Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean, Pacific Dept.: AMA, ANT, CAU, CHO, MET, NAR, SAN, TOL Elev.: 0–1000 m a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE CATALOGUE OF USEFUL PLANTS OF COLOMBIA Ophioglossales 71 Botrychium schaffneri Underw. (Ophioglossaceae) IPNI: 35345-2 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: BOY, MAG, MET, SAN Elev.: 3250–4000 m a.s.l. IUCN Habitat code: 4,6 Conservation status: (IUCN) NE 72 Botrychium underwoodianum Maxon (Ophioglossaceae) IPNI: 35373-2 Habit: Herb Use: ME Origin: Native Conservation status: (IUCN) NE 73 Botrychium virginianum (L.) Sw. (Ophioglossaceae) IPNI: 1106719-2 Habit: Herb Use: HF, ME Origin: Native Region: Andean Dept.: BOY, CUN, MAG, SAN Elev.: 1800–3300 m a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) NE 74 Ophioglossum reticulatum L. (Ophioglossaceae) IPNI: 174612-2 Habit: Herb Use: AF, GS, HF, MA, ME, SU Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CAU, CUN, SAN, TOL, VAC Elev.: 1500–3850 m a.s.l. IUCN Habitat code: 14 Conservation status: (IUCN) LC Osmundales 75 Osmunda regalis L. (Osmundaceae) IPNI: 305721-2 Common name: helecho de espiga Habit: Herb Use: EU, GS, HF, MA, ME, PO Origin: Native Region: Andean Dept.: ANT, CAU, CUN, HUI, SAN Elev.: 1600– 2700 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) LC 76 Osmundastrum cinnamomeum (L.) C.Presl (Osmundaceae) IPNI: 60447721-2 Habit: Herb Use: EU, ME Origin: Native Region: Andean Dept.: ANT, BOY, HUI, SAN Elev.: 1800–2200 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) NE Polypodiales 173 CHECKLIST OF USEFUL PLANTS OF COLOMBIA 77 Asplenium auritum Sw. (Aspleniaceae) IPNI: 22222-2 Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Orinoquia, Pacific Dept.: ANT, BOG, BOY, CAL, CAQ, CAU, CHO, CUN, HUI, MAG, MET, NAR, NSA, PUT, QUI, RIS, SAN, TOL, VAU, VID Elev.: 02820 m.a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 78 Asplenium bipartitum Bory ex Willd. (Aspleniaceae) IPNI: 17039970-1 Use: ME Conservation status: (IUCN) NE 79 Asplenium cuneatum Lam. (Aspleniaceae) IPNI: 328699-2 Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean Dept.: AMA, ANT, NSA Elev.: 200–600 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 80 Asplenium formosum Willd. (Aspleniaceae) IPNI: 22417-2 Habit: Herb Use: EU Origin: Native Region: Andean, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: ANT, BOY, CAL, CAU, CES, CHO, CUN, LAG, MAG, MET, NAR, TOL Elev.: 50–1300 m a.s.l. IUCN Habitat code: 1,2,3,14 Conservation status: (IUCN) LC 81 Asplenium monanthes L. (Aspleniaceae) IPNI: 22558-2 Habit: Herb Use: ME Origin: Native Region: Andean, Guiana Shield Dept.: ANT, ARA, BOG, BOY, CAL, CAU, CUN, MAG, MET, NAR, NSA, RIS, SAN, TOL Elev.: 1050-4400 m.a.s.l. IUCN Habitat code: 1,3,4,8,14 Conservation status: (IUCN) NE (CPSG) Potential LC 82 Asplenium praemorsum Sw. (Aspleniaceae) IPNI: 328509-2 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CAU, CES, CUN, HUI, MAG, NAR, NSA, QUI, SAN Elev.: 1200–3050 m a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) NE 83 Asplenium radicans L. (Aspleniaceae) IPNI: 22662-2 Habit: Herb Use: EU, ME Origin: Native Region: Andean, Guiana Shield, Pacific Dept.: ANT, BOG, BOY, CAL, CAQ, CAS, CAU, CES, CHO, CUN, HUI, MAG, MET, NAR, NSA, PUT, QUI, RIS, SAN, TOL, VAC Elev.: 1003500 m.a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 84 Asplenium serratum L. (Aspleniaceae) IPNI: 22751-2 Common name: doradilla, pencón Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, CAQ, CAU, CHO, GUA, LAG, MAG, MET, NSA, PUT, RIS, SAN, VAC, VAU, VID Elev.: 40-900 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 174 85 Athyrium asplenioides Desv. (Aspleniaceae) IPNI: 17054310-1 Use: ME Region: Andean Dept.: BOG Conservation status: (IUCN) NE 86 Athyrium filix-femina (L.) Roth (Aspleniaceae) IPNI: 30055316-2 Common name: helecho macho Habit: Herb Use: EU, HF, ME Origin: Native Region: Andean Dept.: ANT, BOG, CUN Elev.: 2300–3100 m a.s.l. IUCN Habitat code: 1,4 Conservation status: (IUCN) NE 87 Blechnum brasiliense Desv. (Aspleniaceae) IPNI: 17058320-1 Habit: Herb Use: EU Origin: Native, Cultivated Region: Andean Dept.: ANT, HUI Elev.: 850–1100 m a.s.l. IUCN Habitat code: 3 Conservation status: (IUCN) NE 88 Blechnum cordatum (Desv.) Hieron. (Aspleniaceae) IPNI: 33533-2 Habit: Herb Use: HF, ME Origin: Native Region: Andean, Caribbean, Pacific Dept.: ANT, BOL, BOY, CAL, CAQ, CAU, CHO, CUN, HUI, LAG, NAR, PUT, RIS, SAN, TOL, VAC Elev.: 200-3100 m.a.s.l. IUCN Habitat code: 1,3,4,8,14 Conservation status: (IUCN) NE (CPSG) Potential LC 89 Blechnum loxense (Kunth) Hook. ex Salomon (Aspleniaceae) IPNI: 77186436-1 Habit: Herb Use: EU Origin: Native Region: Andean Dept.: ANT, BOY, CAL, CAU, CES, CHO, CUN, HUI, MAG, MET, NAR, PUT, RIS, SAN, TOL Elev.: 2600-3900 m.a.s.l. IUCN Habitat code: 1,3,4,5,8,14 Conservation status: (IUCN) NE (CPSG) Potential LC 90 Blechnum occidentale L. (Aspleniaceae) IPNI: 33632-2 Habit: Herb Use: HF, ME Origin: Native Region: Andean, Guiana Shield, Caribbean, Pacific Dept.: ANT, BOG, BOY, CAL, CAU, CES, CHO, CUN, HUI, LAG, MAG, MET, NAR, NSA, QUI, RIS, SAN, TOL, VAC Elev.: 50-2850 m.a.s.l. IUCN Habitat code: 1,2,3,4,14 Conservation status: (IUCN) NE (CPSG) Potential LC 91 Diplazium expansum Willd. (Aspleniaceae) IPNI: 17086820-1 Habit: Herb Use: HF Origin: Native Region: Andean Dept.: ANT, BOY, CUN Elev.: 2000–2800 m a.s.l. IUCN Habitat code: 1,3 Conservation status: (IUCN) NE 94 Macrothelypteris torresiana (Gaudich.) Ching (Aspleniaceae) IPNI: 1019960-2 Habit: Herb Use: EU, ME Origin: Naturalised Region: Andean, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: ANT, BOY, CAL, CAU, CHO, CUN, HUI, LAG, MAG, MET, NAR, NSA, PUT, QUI, RIS, SAN, TOL, VAC Elev.: 0–2500 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 95 Telmatoblechnum serrulatum (Rich.) Perrie, D.J.Ohlsen & Brownsey (Aspleniaceae) IPNI: 77142412-1 Habit: Herb Use: HF Origin: Native Region: Amazonia, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: ANT, CAQ, CHO, COR, GUA, MET, NAR, SAN, VAU, VID Elev.: 0–550 m a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE 96 Thelypteris dentata (Forssk.) E.P.St.John (Aspleniaceae) IPNI: 251308-2 Habit: Herb Use: MA, ME, PO Origin: Naturalised Region: Amazonia, Andean, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, CAQ, CHO, CUN, HUI, MAG, MET, TOL, VAC Elev.: 50–1700 m a.s.l. IUCN Habitat code: 1,5,14 Conservation status: (IUCN) NE 97 Thelypteris gongylodes (Schkuhr) Small (Aspleniaceae) IPNI: 251353-2 Use: GS, HF, ME Origin: Native IUCN Habitat code: 1,3 Conservation status: (IUCN) NE 98 Thelypteris interrupta (Willd.) K.Iwats. (Aspleniaceae) IPNI: 17458930-1 Habit: Herb Use: HF, MA, ME Origin: Native Region: Andean, Caribbean, Pacific Dept.: ANT, BOL, CUN, VAC Elev.: 100–2400 m a.s.l. Conservation status: (IUCN) NE 99 Thelypteris kunthii (Desv.) C.V.Morton (Aspleniaceae) IPNI: 251400-2 Habit: Herb Use: ME Origin: Endemic Region: Andean, Caribbean Dept.: ATL, BOY, HUI, LAG Elev.: 100–1500 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 100 Thelypteris nicaraguensis (E.Fourn.) C.V.Morton (Aspleniaceae) IPNI: 252198-2 Habit: Herb Use: ME Origin: Native Region: Pacific Dept.: CHO Elev.: 100–100 m a.s.l. Conservation status: (IUCN) NE 92 Diplazium subserratum (Blume) T.Moore (Aspleniaceae) IPNI: 17088960-1 Use: HF Conservation status: (IUCN) NE 93 Hymenasplenium obtusifolium (L.) L.Regalado & Prada (Aspleniaceae) IPNI: 60458652-2 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: CUN, MAG, NSA Elev.: 900–1600 m a.s.l. IUCN Habitat code: 1,2,14 Conservation status: (IUCN) NE CATALOGUE OF USEFUL PLANTS OF COLOMBIA 101 Thelypteris opulenta (Kaulf.) Fosberg (Aspleniaceae) IPNI: 17460390-1 Habit: Herb Use: HF, ME Origin: Naturalised Region: Amazonia, Guiana Shield, Orinoquia, Pacific Dept.: AMA, ANT, CAQ, CAU, CHO, MET, NAR, PUT, TOL, VAC Elev.: 0–500 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) NE 102 Cystopteris fragilis (L.) Bernh. (Cystopteridaceae) IPNI: 30013429-2 Habit: Herb Use: EU, GS, HF, ME Origin: Native Region: Andean Dept.: ANT, ARA, BOG, BOY, CAL, CAU, CUN, HUI, MAG, MET, NAR, NSA, PUT, RIS, SAN, TOL Elev.: CHECKLIST OF USEFUL PLANTS OF COLOMBIA 1400-4500 m.a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) NE (CPSG) Potential LC 103 Dennstaedtia cicutaria (Sw.) T.Moore (Dennstaedtiaceae) IPNI: 17081410-1 Habit: Herb Use: EU Origin: Native Region: Andean, Pacific Dept.: ANT, CAL, CHO, CUN, HUI, MAG, MET, NAR, NSA, SAN, TOL Elev.: 0–2600 m a.s.l. IUCN Habitat code: 1,2,3,5,14 Conservation status: (IUCN) NE 104 Hypolepis hostilis (Kunze) C.Presl (Dennstaedtiaceae) IPNI: 17126480-1 Habit: Herb Use: HF Origin: Native Region: Amazonia, Andean, Guiana Shield, Pacific Dept.: AMA, ANT, CAQ, CAU, MET, NAR, NSA, QUI, SAN, VAU Elev.: 100–1900 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) NE 105 Hypolepis repens (L.) C.Presl (Dennstaedtiaceae) IPNI: 1031307-2 Habit: Herb Use: HF, ME Origin: Native Region: Andean, Pacific Dept.: ANT, CAQ, CHO, NAR, NSA, VAC Elev.: 0–1650 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 106 Pteridium aquilinum (L.) Kuhn (Dennstaedtiaceae) IPNI: 17210060-1 Common name: caramo, helecho de marrano, helecho liso, helecho marranero, milile, mililí, pata de cuervo Use: AF, GS, HF, MA, ME, PO, SU Dept.: BOG IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE 107 Pteridium arachnoideum (Kaulf.) Maxon (Dennstaedtiaceae) IPNI: 213798-2 Common name: helecho de marrano, helecho marranero Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, BOG, BOY, CAQ, CAU, CES, CHO, CUN, HUI, MAG, MET, NAR, NSA, SAN, VAU Elev.: 2403200 m.a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE (CPSG) Potential LC 108 Pteridium caudatum Maxon (Dennstaedtiaceae) IPNI: 213799-2 Habit: Herb Use: HF, ME, PO Origin: Native Region: Amazonia, Andean, Caribbean, Pacific Dept.: AMA, ANT, BOG, BOY, CAL, CAQ, CUN, HUI, SAP, SAN Elev.: 0– 2600 m a.s.l. IUCN Habitat code: 1,2,3,4,5,8,14 Conservation status: (IUCN) NE 109 Campyloneurum albopunctatissimum (Lellinger) Christenh. (Polypodiaceae) IPNI: 77183020-1 Habit: Herb Use: ME Origin: Native Region: Andean, Guiana Shield, Pacific Dept.: ANT, CAU, CHO, CUN, MET, NAR, SAN Elev.: 500–3050 m a.s.l. Conservation status: (IUCN) NE 110 Campyloneurum angustifolium Fée (Polypodiaceae) IPNI: 17063000-1 Common name: calaguala, pecosa Habit: Herb Use: EU, ME Origin: Native Region: Amazonia, Andean, Guiana Shield Dept.: AMA, ANT, BOG, BOY, CAL, CAS, CAU, CES, CUN, HUI, MAG, MET, NAR, QUI, RIS, SAN, VAC Elev.: 450-3500 m.a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 111 Campyloneurum crassifolium (L.) Christenh. (Polypodiaceae) IPNI: 77183236-1 Common name: maiku jat Habit: Herb Use: EU, ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, BOL, BOY, CAQ, CAU, CHO, CUN, HUI, MET, PUT, QUI, RIS, SAN, TOL, VAC, VAU Elev.: 0-3300 m.a.s.l. Conservation status: (IUCN) NE (CPSG) Potential LC 119 Dryopteris wallichiana (Spreng.) Hyl. (Polypodiaceae) IPNI: 17392950-1 Common name: helecho de la suerte, helecho macho Habit: Herb Use: EU, ME, SU Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CAL, CAU, CES, CUN, MAG, MET, NAR, NSA, QUI, RIS, SAN, TOL Elev.: 350-3600 m.a.s.l. IUCN Habitat code: 1,3,4,8,14 Conservation status: (IUCN) NE (CPSG) Potential LC 112 Campyloneurum densifolium (Hieron.) Lellinger (Polypodiaceae) IPNI: 274004-2 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CAU, CES, CUN, MAG, NAR, QUI, RIS, SAN, TOL Elev.: 1200–3400 m a.s.l. IUCN Habitat code: 1,3,4,8,14 Conservation status: (IUCN) NE 120 Elaphoglossum gayanum (Fée) T.Moore (Polypodiaceae) IPNI: 17103530-1 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, ARA, BOG, BOY, CAL, CAU, CES, CUN, MAG, MET, NAR, NSA, SAN Elev.: 1700-4300 m.a.s.l. IUCN Habitat code: 1,3,4,8,14 Conservation status: (IUCN) NE (CPSG) Potential LC 113 Campyloneurum phyllitidis (L.) C.Presl (Polypodiaceae) IPNI: 17063380-1 Common name: helecha Habit: Herb Use: EU, ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, BOY, CAL, CAQ, CAU, CES, CHO, COR, CUN, HUI, LAG, MAG, MET, NAR, NSA, TOL, VAC, VAU, VID Elev.: 0-2800 m.a.s.l. IUCN Habitat code: 1,2,3,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 121 Elaphoglossum metallicum Mickel (Polypodiaceae) IPNI: 276374-2 Habit: Herb Use: EU Origin: Native Region: Andean Dept.: ANT Elev.: 2800–2800 m a.s.l. IUCN Habitat code: 14 Conservation status: (IUCN) NE 114 Cyclopeltis semicordata (Sw.) J.Sm. (Polypodiaceae) IPNI: 1148359-2 Habit: Herb Use: ME Origin: Native Region: Andean, Guiana Shield, Caribbean, Pacific Dept.: ANT, BOL, BOY, CAL, CES, CHO, CUN, LAG, MAG, MET, NSA, SAN, SUC, TOL Elev.: 0-1600 m.a.s.l. IUCN Habitat code: 1,2,3,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 115 Davallia canariensis (L.) Sm. (Polypodiaceae) IPNI: 17078300-1 Common name: helecho encaje, rabo de mico Habit: Herb Use: EU Origin: Cultivated Dept.: ANT, BOG IUCN Habitat code: 1,14 Conservation status: (IUCN) NE 116 Davallia solida (G.Forst.) Sw. (Polypodiaceae) IPNI: 17080730-1 Habit: Herb Use: ME Origin: Cultivated Region: Andean Dept.: CUN Elev.: 2500–2600 m a.s.l. Conservation status: (IUCN) NE 117 Didymochlaena truncatula (Sw.) J.Sm. (Polypodiaceae) IPNI: 17084790-1 Habit: Herb Use: EU, ME Origin: Native Region: Amazonia, Andean, Caribbean, Pacific Dept.: AMA, ANT, CAU, CES, CHO, CUN, LAG, MAG, NAR, NSA, PUT, QUI, RIS, SAN, VAC Elev.: 50-2350 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) LC (CPSG) Potential LC 118 Dryopteris fengiana (Ching) ined. (Polypodiaceae) IPNI: 77205539-1 Habit: Herb Use: EU Origin: Cultivated Region: Andean Dept.: TOL Elev.: 1300–1300 m a.s.l. Conservation status: (IUCN) NE CATALOGUE OF USEFUL PLANTS OF COLOMBIA 122 Elaphoglossum minutum (Pohl ex Fée) T.Moore (Polypodiaceae) IPNI: 17104440-1 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, BOG, CAU, CUN, MAG, NAR Elev.: 2800– 3650 m a.s.l. Conservation status: (IUCN) NE 123 Elaphoglossum petiolatum Urb. (Polypodiaceae) IPNI: 89270-2 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: CHO, TOL Elev.: 1900–3000 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) NE 124 Lomariopsis japurensis (C.Mart.) J.Sm. (Polypodiaceae) IPNI: 17442740-1 Common name: bagre pajo, charapa huasca Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean Dept.: AMA, ANT, CAQ, MET, PUT Elev.: 200–1000 m a.s.l. IUCN Habitat code: 1,3,5,14 Conservation status: (IUCN) NE 125 Microgramma lycopodioides (L.) Copel. (Polypodiaceae) IPNI: 160142-2 Habit: Herb Use: HF, ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean, Pacific Dept.: AMA, ANT, BOY, CAL, CAU, CES, CHO, CUN, HUI, LAG, MAG, MET, NAR, PUT, SAN, TOL, VAC, VAU Elev.: 0-1600 m.a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 126 Microgramma nitida (J.Sm.) A.R.Sm. (Polypodiaceae) IPNI: 17444940-1 Habit: Herb Use: ME Origin: Native Region: Caribbean Dept.: SAP Conservation status: (IUCN) NE 127 Microgramma percussa (Cav.) de la Sota (Polypodiaceae) IPNI: 17546710-1 Common name: calaguala, helecho de bejuco, herb de lagartija, planta de sanguijuela Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean, Pacific Dept.: AMA, ANT, BOY, CAL, CAQ, CES, CHO, CUN, HUI, LAG, MAG, MET, NAR, NSA, PUT, RIS, SAN, TOL, VAC, VID Elev.: 0-2600 m.a.s.l. 175 CHECKLIST OF USEFUL PLANTS OF COLOMBIA IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 128 Microgramma vacciniifolia (Langsd. & Fisch.) Copel. (Polypodiaceae) IPNI: 17406250-1 Habit: Herb Use: ME Origin: Native Region: Caribbean Dept.: LAG, MAG Elev.: 350–450 m a.s.l. Conservation status: (IUCN) NE 129 Microsorum punctatum Copel. (Polypodiaceae) IPNI: 17341250-1 Common name: manoetigre Habit: Herb Use: EU, HF, ME Origin: Cultivated, Naturalised Dept.: ANT IUCN Habitat code: 14 Conservation status: (IUCN) NE 130 Nephrolepis biserrata (Sw.) Schott (Polypodiaceae) IPNI: 17160710-1 Common name: helecho cola de pez Habit: Herb Use: EU, GS, HF, MA, ME, PO, SU Origin: Native Region: Amazonia, Andean, Caribbean, Pacific Dept.: AMA, ANT, CAQ, CHO, COR, CUN, MET, SAP, SAN, VAC, VID Elev.: 100–1150 m a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE 131 Nephrolepis brownii (Desv.) Hovenkamp & Miyam. (Polypodiaceae) IPNI: 77069554-1 Habit: Herb Use: ME Origin: Naturalised Region: Amazonia, Andean, Pacific Dept.: ANT, CAU, CHO, RIS, VAC, VAU Elev.: 0–2000 m a.s.l. Conservation status: (IUCN) NE 132 Nephrolepis cordifolia (L.) C.Presl (Polypodiaceae) IPNI: 17160730-1 Habit: Herb Use: EU, HF, ME, SU Origin: Cultivated, Naturalised Region: Amazonia, Andean, Pacific Dept.: ANT, BOG, BOY, CAL, CAU, CHO, CUN, HUI, MAG, NSA, PUT, SAN, TOL, VAC, VAU Elev.: 0–2800 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 133 Nephrolepis exaltata (L.) Schott (Polypodiaceae) IPNI: 17160830-1 Common name: aliento de niño, encaje, espárrago fino, helecho churco, helecho crespo, llanto de niño, sueño, sueño de niño Habit: Herb Use: EU, ME Origin: Native Region: Caribbean, Pacific Dept.: ANT, BOG, CUN, LAG, VAC Elev.: 250–2500 m a.s.l. IUCN Habitat code: 1,2,3,14 Conservation status: (IUCN) NE 134 Nephrolepis pectinata (Willd.) Schott (Polypodiaceae) IPNI: 17161100-1 Habit: Herb Use: EU Origin: Native Region: Andean, Pacific Dept.: ANT, BOG, BOY, CAQ, CHO, CUN, NAR, NSA, PUT, SAN Elev.: 50–2470 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 135 Nephrolepis undulata J.Sm. (Polypodiaceae) IPNI: 17161310-1 Habit: Herb Use: HF, ME, SU Origin: Native Region: Andean Dept.: ANT, BOY, CHO, HUI, MAG Elev.: 650– 2450 m a.s.l. IUCN Habitat code: 1,3 Conservation status: (IUCN) LC 176 136 Oleandra articulata (Sw.) C.Presl (Polypodiaceae) IPNI: 1138771-2 Common name: yerba de ojo Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Pacific Dept.: AMA, ANT, CAU, CHO, MET, NAR, NSA, PUT, VAC, VAU Elev.: 0–1800 m a.s.l. IUCN Habitat code: 1,3,5,14 Conservation status: (IUCN) NE 137 Oleandra pilosa Hook. (Polypodiaceae) IPNI: 173662-2 Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Pacific Dept.: AMA, ANT, BOY, CAQ, CAU, CHO, CUN, MAG, MET, NSA, SAN, VAC, VAU Elev.: 500-1900 m.a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 138 Phlebodium areolatum (Willd.) J.Sm. (Polypodiaceae) IPNI: 17175380-1 Common name: calaguala Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, BOY, CAU, CUN, MAG, NSA, SAN Elev.: 800–2300 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 139 Phlebodium aureum (L.) J.Sm. (Polypodiaceae) IPNI: 17175390-1 Common name: calaguala, calahuala, cartagenero, helecho costeño, helecho cuero, polipodio Habit: Herb Use: EU, HF, MA, ME Origin: Native Region: Andean Dept.: ANT, BOG Elev.: 450–2000 m a.s.l. IUCN Habitat code: 1,2,3,5,14 Conservation status: (IUCN) NE MET, NAR, NSA, PUT, RIS, SAN, TOL, VAC Elev.: 14003600 m.a.s.l. IUCN Habitat code: 1,2,3,4,8,14 Conservation status: (IUCN) NE (CPSG) Potential LC 145 Pleopeltis polypodioides (L.) E.G.Andrews & Windham (Polypodiaceae) IPNI: 1039912-2 Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean Dept.: ANT, BOG, BOY, CAQ, CES, CUN, MAG, MET, NSA, TOL Elev.: 250–3000 m a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) NE 146 Polybotrya botryoides (Baker) C.Chr. (Polypodiaceae) IPNI: 17180780-1 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT Elev.: 1800–2000 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) NE 147 Polybotrya osmundacea Humb. & Bonpl. ex Willd. (Polypodiaceae) IPNI: 30017613-2 Habit: Herb Use: MA Origin: Native Region: Amazonia, Andean, Caribbean, Pacific Dept.: AMA, ANT, BOY, CAU, CHO, CUN, MAG, MET, PUT, RIS, SAN, VAC Elev.: 100-2500 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 148 Polybotrya serratifolia (Fée) Klotzsch (Polypodiaceae) IPNI: 17181340-1 Habit: Herb Use: ME Origin: Native Region: Andean, Orinoquia Dept.: ANT, MET Elev.: 500–1500 m a.s.l. IUCN Habitat code: 1 Conservation status: (IUCN) NE 140 Phlebodium decumanum (Willd.) J.Sm. (Polypodiaceae) IPNI: 17175400-1 Common name: copudo, rabo de ardita Habit: Herb Use: EU, ME Origin: Native Region: Amazonia, Andean, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, BOL, BOY, CAQ, CES, CHO, COR, CUN, MAG, MET, NAR, NSA, SAN, TOL, VAU, VID Elev.: 100-1200 m.a.s.l. IUCN Habitat code: 1,2,3,4,14 Conservation status: (IUCN) NE (CPSG) Potential LC 141 Platycerium bifurcatum (Cav.) C.Chr. (Polypodiaceae) IPNI: 17177460-1 Common name: cacho de venado, cachoevenado, cuerno, helecho cuerno Habit: Herb Use: EU, ME Origin: Cultivated, Naturalised Dept.: ANT, BOG IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 142 Platycerium elephantotis Schweinf. (Polypodiaceae) IPNI: 17177470-1 Habit: Herb Use: EU, ME Origin: Cultivated, Naturalised Region: Andean Dept.: QUI, VAC Conservation status: (IUCN) NE 143 Pleopeltis balaonensis (Hieron.) A.R.Sm. (Polypodiaceae) IPNI: 77141144-1 Habit: Herb Use: ME Origin: Native Conservation status: (IUCN) NE 144 Pleopeltis macrocarpa (Willd.) Kaulf. (Polypodiaceae) IPNI: 17448650-1 Common name: calaguala Habit: Herb Use: ME, PO Origin: Native Region: Andean, Guiana Shield Dept.: ANT, BOG, BOY, CAL, CAU, CES, CUN, HUI, MAG, CATALOGUE OF USEFUL PLANTS OF COLOMBIA 149 Polypodium fraternum Schltdl. & Cham. (Polypodiaceae) IPNI: 17189260-1 Use: ME Origin: Native IUCN Habitat code: 14 Conservation status: (IUCN) NE 150 Polypodium vulgare L. (Polypodiaceae) IPNI: 300720-2 Common name: helecho peine, polipodio Use: EU, GS, HF, ME IUCN Habitat code: 3,14 Conservation status: (IUCN) NE 151 Polystichum montevidense Rosenst. (Polypodiaceae) IPNI: 208026-2 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CAL, CAU, CUN, HUI, NAR, SAN, VAC Elev.: 2100–4050 m a.s.l. IUCN Habitat code: 3,4 Conservation status: (IUCN) NE 152 Rumohra adiantiformis (G.Forst.) Ching (Polypodiaceae) IPNI: 224543-2 Common name: helecho cuero, helecho de cuero Habit: Herb Use: EU, ME Origin: Native, Cultivated Dept.: BOG IUCN Habitat code: 3,14 Conservation status: (IUCN) NE 153 Serpocaulon attenuatum (Humb. & Bonpl. ex Willd.) A.R.Sm. (Polypodiaceae) IPNI: 77075087-1 Habit: Herb Use: ME Origin: Native Region: Caribbean Dept.: CES Elev.: 100–100 m a.s.l. Conservation status: (IUCN) NE CHECKLIST OF USEFUL PLANTS OF COLOMBIA 154 Serpocaulon levigatum (Cav.) A.R.Sm. (Polypodiaceae) IPNI: 77075105-1 Common name: calaguala Habit: Herb Use: ME Origin: Native Region: Andean, Guiana Shield, Caribbean, Pacific Dept.: ANT, BOG, BOY, CAL, CAU, CES, CHO, CUN, HUI, LAG, MAG, NAR, NSA, PUT, QUI, RIS, SAN, VAC Elev.: 200-2850 m.a.s.l. IUCN Habitat code: 1,2,3,4,14 Conservation status: (IUCN) NE (CPSG) Potential LC 155 Serpocaulon maritimum (Hieron.) A.R.Sm. (Polypodiaceae) IPNI: 77075108-1 Habit: Herb Use: ME Origin: Native Region: Andean, Caribbean, Pacific Dept.: ANT, CAU, CHO, LAG, NAR, VAC Elev.: 0–1650 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 156 Serpocaulon triseriale (Sw.) A.R.Sm. (Polypodiaceae) IPNI: 77075127-1 Common name: helecho, nujubujupiali Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Orinoquia, Pacific Dept.: AMA, ANT, BOG, BOY, CAQ, CAU, CHO, CUN, GUA, HUI, MAG, MET, NAR, SAN, VAC, VAU Elev.: 40-2800 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 157 Tectaria heracleifolia Underw. (Polypodiaceae) IPNI: 249899-2 Habit: Herb Use: EU, ME Origin: Native Region: Amazonia, Andean, Caribbean Dept.: AMA, ANT, BOL, BOY, CAL, CUN, LAG, MAG Elev.: 50–1300 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 158 Tectaria incisa Cav. (Polypodiaceae) IPNI: 17226640-1 Habit: Herb Use: EU, ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, ATL, BOL, BOY, CAL, CAQ, CES, CHO, COR, CUN, GUV, LAG, MAG, MET, NAR, NSA, PUT, QUI, RIS, SAN, TOL, VAC, VAU Elev.: 01900 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 159 Tectaria mexicana (Fée) C.V.Morton (Polypodiaceae) IPNI: 249914-2 Habit: Herb Use: ME Origin: Native Region: Andean, Pacific Dept.: ANT, CHO, MAG, NAR Elev.: 0–700 m a.s.l. IUCN Habitat code: 1,2,3,5,14 Conservation status: (IUCN) NE 160 Triplophyllum funestum (Kunze) Holttum (Polypodiaceae) IPNI: 286436-2 Habit: Herb Use: ME Origin: Native Region: Amazonia, Guiana Shield, Pacific Dept.: AMA, ANT, CAQ, CHO, GUA, GUV, MET, NAR, SAN, VAC, VAU, VID Elev.: 01000 m.a.s.l. IUCN Habitat code: 1,3,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 161 Acrostichum aureum L. (Pteridaceae) IPNI: 319061-2 Common name: carranconcha, chicharrón, chiguamacho, corocilla, corozo de ciénaga, helecho chiguamacho, helecho mano de tigre, madrevieja, matatigre, racancha, ranconcha Habit: Herb Use: AF, EU, GS, HF, MA, ME Origin: Native Region: Andean, Caribbean, Pacific Dept.: ANT, ATL, BOL, CAU, CHO, COR, MAG, NAR, SUC, VAC Elev.: 0–950 m a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) LC 162 Acrostichum danaeifolium Langsd. & Fisch. (Pteridaceae) IPNI: 17002360-1 Habit: Herb Use: HF, ME Origin: Native Region: Andean, Caribbean, Pacific Dept.: ANT, CHO, LAG, MAG, VAC Elev.: 0–500 m a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) LC 163 Adiantum anceps Maxon & C.V.Morton (Pteridaceae) IPNI: 4862-2 Habit: Herb Use: EU Origin: Native Conservation status: (IUCN) NE 164 Adiantum capillus-veneris L. (Pteridaceae) IPNI: 325030-2 Common name: capilera, cilantrillo, culantrillo, helecho cilantro, helecho granizo Use: EU, GS, HF, ME, SU Dept.: BOG IUCN Habitat code: 1,3,14 Conservation status: (IUCN) LC 165 Adiantum concinnum Humb. & Bonpl. ex Willd. (Pteridaceae) IPNI: 315077-2 Common name: cilantrillo, culantrillo Habit: Herb Use: EU, ME Origin: Native Region: Andean, Guiana Shield, Caribbean, Pacific Dept.: BOG, BOY, CAL, CAU, CUN, HUI, LAG, MAG, MET, NAR, NSA, VAC Elev.: 250– 2000 m a.s.l. IUCN Habitat code: 1,2,3,14 Conservation status: (IUCN) NE 166 Adiantum deflectens Mart. (Pteridaceae) IPNI: 17009750-1 Habit: Herb Use: HF Origin: Native Region: Caribbean, Pacific Dept.: LAG, VAC Elev.: 700–700 m a.s.l. IUCN Habitat code: 1 Conservation status: (IUCN) NE 167 Adiantum latifolium Lam. (Pteridaceae) IPNI: 271633-2 Common name: culantrillo negro, helechillo, helecho de diablo, helecho de perfume Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Orinoquia, Pacific Dept.: AMA, ANT, BOY, CAQ, CAU, CHO, CUN, GUV, MET, NAR, NSA, PUT, VAC, VAU, VID Elev.: 100-1500 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 168 Adiantum lucidum (Cav.) Sw. (Pteridaceae) IPNI: 17011270-1 Common name: nabugui Habit: Herb Use: ME Origin: Native Region: Andean, Caribbean, Pacific Dept.: ANT, ATL, CHO, COR, MAG Elev.: 0–300 m a.s.l. IUCN Habitat code: 1,2,3,4,14 Conservation status: (IUCN) NE 169 Adiantum macrophyllum Sw. (Pteridaceae) IPNI: 17011380-1 Habit: Herb Use: EU Origin: Native Region: Andean, Guiana Shield, Caribbean, Pacific Dept.: ANT, CAL, CHO, CUN, LAG, MAG, MET, NAR, NSA, RIS, SAN, VAC Elev.: 0-1600 m.a.s.l. IUCN Habitat code: 1,2,3,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC CATALOGUE OF USEFUL PLANTS OF COLOMBIA 170 Adiantum orbignyanum Mett. (Pteridaceae) IPNI: 319723-2 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: BOY, CAU Elev.: 1900–2200 m a.s.l. IUCN Habitat code: 1 Conservation status: (IUCN) NE 171 Adiantum philippense L. (Pteridaceae) IPNI: 325052-2 Habit: Herb Use: ME Origin: Native Region: Andean, Caribbean Dept.: BOY, MAG Elev.: 450–650 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) NE 172 Adiantum poiretii Wikstr. (Pteridaceae) IPNI: 17012320-1 Habit: Herb Use: EU, ME Origin: Native Region: Andean Dept.: BOG, BOY, CUN, NAR Elev.: 1500–3000 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) NE 173 Adiantum raddianum C.Presl (Pteridaceae) IPNI: 17012580-1 Common name: culantrillo Habit: Herb Use: EU, ME Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CAL, CAU, CUN, HUI, MAG, NAR, QUI, SAN, TOL, VAC Elev.: 1000-3450 m.a.s.l. IUCN Habitat code: 1,2,3,4,14 Conservation status: (IUCN) NE (CPSG) Potential LC 174 Adiantum tenerum Sw. (Pteridaceae) IPNI: 17013290-1 Habit: Herb Use: EU, ME Origin: Native Region: Andean, Caribbean Dept.: BOY, CES, CUN, MAG Elev.: 800–1650 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 175 Adiantum tetraphyllum Humb. & Bonpl. ex Willd. (Pteridaceae) IPNI: 315261-2 Common name: culantrillo Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Caribbean, Pacific Dept.: ANT, BOY, CAL, CAU, CHO, CUN, MAG, MET, NSA, PUT, RIS, TOL, VAC, VAU Elev.: 50-1800 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 176 Adiantum tinctum T.Moore (Pteridaceae) IPNI: 17013420-1 Use: ME Origin: Native Conservation status: (IUCN) NE 177 Adiantum villosum L. (Pteridaceae) IPNI: 315262-2 Habit: Herb Use: ME Origin: Native Region: Andean, Caribbean, Pacific Dept.: LAG, MAG, NAR Elev.: 850– 850 m a.s.l. IUCN Habitat code: 1,2,3,14 Conservation status: (IUCN) NE 178 Adiantum wilsonii Hook. (Pteridaceae) IPNI: 17013890-1 Habit: Herb Use: ME Origin: Native Region: Pacific Dept.: ANT, CHO, SAN Elev.: 0–500 m a.s.l. IUCN Habitat code: 1,3 Conservation status: (IUCN) NE 177 CHECKLIST OF USEFUL PLANTS OF COLOMBIA 179 Ceratopteris pteridoides (Hook.) Hieron. (Pteridaceae) IPNI: 17242590-1 Common name: helecho acuático Habit: Herb, Aquatic Use: EU, HF, ME Origin: Native Region: Amazonia, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, ARA, ATL, BOL, CHO, COR, SAN, SUC, VID Elev.: 0–150 m a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE 180 Ceratopteris thalictroides (L.) Brongn. (Pteridaceae) IPNI: 17064190-1 Habit: Herb, Aquatic Use: EU, GS, HF, ME Origin: Native Region: Caribbean, Orinoquia, Pacific Dept.: ANT, BOL, CHO, MET Elev.: 50–450 m a.s.l. IUCN Habitat code: 1,5,14 Conservation status: (IUCN) LC 181 Hemionitis chlorophylla (Sw.) Christenh. (Pteridaceae) IPNI: 77182519-1 Habit: Herb Use: ME Origin: Native Conservation status: (IUCN) NE 182 Hemionitis concolor (Langsd. & Fisch.) Christenh. (Pteridaceae) IPNI: 77182373-1 Habit: Herb Use: ME Origin: Native Region: Caribbean Dept.: CES, MAG Elev.: 50–50 m a.s.l. Conservation status: (IUCN) NE 183 Hemionitis farinosa (Forssk.) Christenh. (Pteridaceae) IPNI: 77182808-1 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: CUN Elev.: 1900–3000 m a.s.l. Conservation status: (IUCN) NE 184 Hemionitis marginata (Kunth) Christenh. (Pteridaceae) IPNI: 77182463-1 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, BOY, CAU, CUN, NAR Elev.: 1800–3800 m a.s.l. Conservation status: (IUCN) NE 185 Hemionitis microphylla (Sw.) Christenh. (Pteridaceae) IPNI: 77183177-1 Habit: Herb Use: EU Origin: Native Region: Andean, Caribbean Dept.: ANT, BOY, CAL, CES, CUN, HUI, LAG, MAG, NAR, SAN, VAC Elev.: 250–1600 m a.s.l. Conservation status: (IUCN) NE 186 Hemionitis myriophylla (Desv.) Christenh. (Pteridaceae) IPNI: 77183191-1 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: BOY, CUN, MAG, NAR, SAN Elev.: 1000–3700 m a.s.l. Conservation status: (IUCN) NE 187 Hemionitis nivea (Poir.) Christenh. (Pteridaceae) IPNI: 77182941-1 Habit: Herb Use: ME Origin: Native Conservation status: (IUCN) NE 188 Hemionitis obducta (Mett.) Christenh. (Pteridaceae) IPNI: 77182303-1 Habit: Herb Use: ME Origin: Native Region: Caribbean Dept.: ANT, CES, CUN, HUI, MAG Elev.: 150–600 m a.s.l. Conservation status: (IUCN) NE 178 189 Hemionitis ovata (Desv.) Christenh. (Pteridaceae) IPNI: 77183106-1 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, BOY, CES, CUN, NAR, PUT, SAN Elev.: 1000–2600 m a.s.l. Conservation status: (IUCN) NE 190 Hemionitis ternifolia (Cav.) Christenh. (Pteridaceae) IPNI: 77182326-1 Habit: Herb Use: ME Origin: Native Region: Andean Dept.: BOY, CUN, MAG, NAR, SAN, VAC Elev.: 2000– 3350 m a.s.l. Conservation status: (IUCN) NE 191 Jamesonia bogotensis H.Karst. (Pteridaceae) IPNI: 17127250-1 Habit: Herb Use: EU, MA Origin: Native Region: Andean Dept.: ANT, BOG, BOL, BOY, CUN, NAR, RIS, SAN Elev.: 2950–4400 m a.s.l. IUCN Habitat code: 1,3,4,8,14 Conservation status: (IUCN) NE 192 Jamesonia flexuosa (Kunth) Christenh. (Pteridaceae) IPNI: 77110633-1 Common name: helechillo negro Habit: Herb Use: ME Origin: Native Region: Andean Dept.: ANT, BOG, BOL, BOY, CAL, CAU, CHO, CUN, HUI, MAG, MET, NAR, NSA, PUT, QUI, RIS, SAN, TOL Elev.: 1100–3700 m a.s.l. IUCN Habitat code: 1,3,4,8,14 Conservation status: (IUCN) NE 193 Pityrogramma calomelanos (L.) Link (Pteridaceae) IPNI: 17345640-1 Common name: helecho negro, yerba de la Virgen, yerba de la virgen Habit: Herb Use: EU, MA, ME, PO, SU Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, BOL, BOY, CAL, CAQ, CAU, CHO, CUN, GUA, GUV, HUI, LAG, MAG, MET, NAR, NSA, PUT, QUI, RIS, SAP, SAN, TOL, VAC, VAU, VID Elev.: 0-2700 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 194 Pityrogramma ebenea (L.) Proctor (Pteridaceae) IPNI: 200935-2 Common name: helecho blanco Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Pacific Dept.: ANT, BOY, CAL, CAQ, CAS, CAU, CHO, CUN, HUI, NAR, NSA, PUT, SAN, TOL, VAC Elev.: 200-3200 m.a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE (CPSG) Potential LC 195 Polytaenium guayanense (Hieron.) Alston (Pteridaceae) IPNI: 17355560-1 Common name: calaguala, yamichuí Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Pacific Dept.: AMA, ANT, CAQ, CHO, GUV, HUI, MET, NSA Elev.: 100–1700 m a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE 196 Polytaenium lineatum J.Sm. (Pteridaceae) IPNI: 17209080-1 Habit: Herb Use: ME Origin: Native Region: Andean, Caribbean Dept.: ANT, CAL, CES, CUN, HUI, LAG, MAG, NAR, NSA, RIS, SAN, TOL Elev.: 1500-2800 m.a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE (CPSG) Potential LC CATALOGUE OF USEFUL PLANTS OF COLOMBIA 197 Pteris biaurita L. (Pteridaceae) IPNI: 213840-2 Habit: Herb Use: HF, ME Origin: Native Region: Andean, Guiana Shield, Caribbean, Orinoquia Dept.: BOY, CAQ, COR, CUN, MAG, MET Elev.: 50–1300 m a.s.l. IUCN Habitat code: 1,2,3,14 Conservation status: (IUCN) NE 198 Pteris cretica L. (Pteridaceae) IPNI: 17211870-1 Habit: Herb Use: EU, HF, ME Origin: Naturalised Region: Amazonia, Andean Dept.: AMA, BOG, CUN, SAN Elev.: 200–2700 m a.s.l. IUCN Habitat code: 1,3,5,14 Conservation status: (IUCN) NE 199 Pteris grandifolia L. (Pteridaceae) IPNI: 313710-2 Habit: Herb Use: EU, ME Origin: Native Region: Andean, Caribbean Dept.: ANT, ATL, BOY, CAL, CUN, HUI, TOL Elev.: 350–1800 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 200 Pteris quadriaurita Retz. (Pteridaceae) IPNI: 17215900-1 Habit: Herb Use: EU, ME Origin: Native Region: Andean, Pacific Dept.: ANT, BOY, CAL, CAU, CES, CUN, HUI, MAG, VAC Elev.: 900–2700 m a.s.l. IUCN Habitat code: 1,2,3,14 Conservation status: (IUCN) NE 201 Pteris tripartita Sw. (Pteridaceae) IPNI: 17217170-1 Habit: Herb Use: EU, ME Origin: Naturalised Region: Amazonia, Pacific Dept.: AMA, CAQ, CHO, PUT Elev.: 50–700 m a.s.l. IUCN Habitat code: 1,3,5 Conservation status: (IUCN) NE 202 Pteris vittata L. (Pteridaceae) IPNI: 17217500-1 Habit: Herb Use: EU, HF, ME, PO Origin: Naturalised Region: Andean Dept.: ANT, VAC Elev.: 1500–1750 m a.s.l. IUCN Habitat code: 1,2,3,14 Conservation status: (IUCN) LC 203 Pterozonium reniforme (Mart.) Fée (Pteridaceae) IPNI: 17217830-1 Habit: Herb Use: ME Origin: Native Region: Amazonia, Guiana Shield Dept.: AMA, CAQ, VAU Elev.: 100– 650 m a.s.l. IUCN Habitat code: 1,4,14 Conservation status: (IUCN) NE 204 Vittaria graminifolia Kaulf. (Pteridaceae) IPNI: 266216-2 Habit: Herb Use: ME Origin: Native Region: Andean, Guiana Shield, Orinoquia, Pacific Dept.: ANT, BOG, BOY, CAU, CES, CHO, CUN, LAG, MAG, MET, NAR, NSA, PUT, SAN, VAC Elev.: 100-3200 m.a.s.l. IUCN Habitat code: 1,2,3,4,14 Conservation status: (IUCN) NE (CPSG) Potential LC 205 Vittaria lineata (L.) Sm. (Pteridaceae) IPNI: 17234040-1 Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Caribbean, Pacific Dept.: AMA, ANT, BOG, BOY, CAQ, CAU, CHO, LAG, MET, NAR, NSA, SAN, VAC Elev.: 0-3450 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC CHECKLIST OF USEFUL PLANTS OF COLOMBIA Psilotales 206 Psilotum complanatum Sw. (Psilotaceae) IPNI: 114220-3 Habit: Herb Use: ME Origin: Native Dept.: BOG Conservation status: (IUCN) NE 207 Psilotum nudum (L.) P.Beauv. (Psilotaceae) IPNI: 212103-2 Habit: Herb Use: EU, HF, IF, ME Origin: Native Region: Andean, Pacific Dept.: ANT, CHO, VAC Elev.: 0–1500 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) NE Salviniales 208 Marsilea polycarpa Hook. & Grev. (Marsileaceae) IPNI: 280439-2 Common name: trébol de agua Habit: Herb Use: HF, ME Origin: Native Region: Caribbean Dept.: ATL, BOL, CAS, COR, HUI Elev.: 0–650 m a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE 209 Azolla caroliniana Willd. (Salviniaceae) IPNI: 17056910-1 Use: EU Origin: Native IUCN Habitat code: 5,14 Conservation status: (IUCN) NE 210 Azolla filiculoides Lam. (Salviniaceae) IPNI: 17056950-1 Common name: alfombra de agua, azola Habit: Herb, Aquatic Use: AF, EU, ME Origin: Native Region: Amazonia, Andean, Caribbean Dept.: AMA, ANT, BOG, BOY, COR, CUN, HUI, MET, RIS, VAU Elev.: 50–4200 m a.s.l. IUCN Habitat code: 1,3,4,5,8,14 Conservation status: (IUCN) NE 211 Azolla microphylla Kaulf. (Salviniaceae) IPNI: 17057000-1 Habit: Herb, Aquatic Use: EU Origin: Native Region: Amazonia, Pacific Dept.: AMA, CHO Elev.: 100–100 m a.s.l. IUCN Habitat code: 1,5 Conservation status: (IUCN) LC 212 Salvinia auriculata Aubl. (Salviniaceae) IPNI: 17219800-1 Habit: Herb, Aquatic Use: EU Origin: Native Region: Amazonia, Caribbean, Orinoquia, Pacific Dept.: AMA, ANT, ATL, CHO, COR, MET, TOL, VAC, VID Elev.: 50–400 m a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE Schizaeales 213 Anemia adiantifolia (L.) Sw. (Schizaeaceae) IPNI: 1031990-2 Habit: Herb Use: EU Origin: Native Conservation status: (IUCN) NE 214 Anemia phyllitidis (L.) Sw. (Schizaeaceae) IPNI: 1202412-2 Habit: Herb Use: EU, ME Origin: Native Region: Andean, Guiana Shield, Caribbean Dept.: ANT, BOY, CAL, CAU, CES, CUN, LAG, MAG, QUI, RIS, SAN, VAC Elev.: 0-2500 m.a.s.l. IUCN Habitat code: 1,2,3,14 Conservation status: (IUCN) NE (CPSG) Potential LC 100-250 m.a.s.l. IUCN Habitat code: 1 Conservation status: (IUCN) CR (CPSG) EN 215 Lygodium venustum Sw. (Schizaeaceae) IPNI: 17143510-1 Common name: bejuco alambre Habit: Climbing Use: ME Origin: Native Region: Andean, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: ANT, ATL, BOL, BOY, CAS, CAU, CES, CHO, COR, CUN, HUI, LAG, MAG, MET, NSA, SAP, SUC, TOL, VAC, VID Elev.: 02750 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC 223 Zamia melanorrhachis D.W.Stev. (Zamiaceae) IPNI: 20009736-1 Common name: corocita, corocito, helecho, ibaracú Habit: Herb Use: HF, ME Origin: Endemic Dept.: ANT, COR, SAN Elev.: 30-500 m.a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) EN (CPSG) EN 216 Lygodium volubile Sw. (Schizaeaceae) IPNI: 17143520-1 Habit: Climbing Use: MA, ME Origin: Native Region: Amazonia, Andean Dept.: AMA, ANT, CAQ, MAG, NSA Elev.: 200–700 m a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) NE 217 Schizaea elegans (Vahl) Sw. (Schizaeaceae) IPNI: 17220460-1 Common name: helecho de madremonte, la diabla, nolocrea macho, sombrillita Habit: Herb Use: ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Caribbean, Pacific Dept.: AMA, ANT, BOL, CAQ, CAS, CAU, CES, CHO, COR, GUA, GUV, MAG, MET, NAR, NSA, PUT, SAN, TOL, VAC Elev.: 50-1400 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) NE (CPSG) Potential LC Cycadopsida Cycadales 218 Cycas revoluta Thunb. (Cycadaceae) IPNI: 328823-2 Common name: cícara, palma de sagú, palma funeral Habit: Shrub Use: EU, GS, HF, MA, ME Origin: Cultivated Region: Andean, Caribbean Dept.: ANT, BOG, CUN, QUI, SAP, TOL Elev.: 500–1600 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) LC 219 Cycas rumphii Miq. (Cycadaceae) IPNI: 326820-2 Common name: cica, palma cica, palma sagú Habit: Shrub Use: EU, GS, HF, IF, MA, ME, PO, SU Origin: Cultivated Region: Andean, Caribbean Dept.: ANT, MET, QUI, TOL Elev.: 500–2000 m a.s.l. IUCN Habitat code: 14 Conservation status: (IUCN) NT 220 Zamia amazonum D.W.Stev. (Zamiaceae) IPNI: 20009732-1 Common name: quiripa, yuca de monte Habit: Herb Use: HF Origin: Native Region: Amazonia Dept.: AMA, VAU Elev.: 100-500 m.a.s.l. IUCN Habitat code: 1 Conservation status: (IUCN) NT (CPSG) VU 221 Zamia chigua Seem. (Zamiaceae) IPNI: 297256-1 Common name: chigua, chigua macho, helecho Habit: Shrub Use: AF, EU, HF Origin: Endemic Region: Pacific Dept.: CHO, VAC Elev.: 30-250 m.a.s.l. IUCN Habitat code: 1,5,14 Conservation status: (IUCN) NT (CPSG) NT 222 Zamia hymenophyllidia D.W.Stev. (Zamiaceae) IPNI: 20009735-1 Habit: Herb Use: HF Origin: Native Region: Amazonia Dept.: AMA Elev.: CATALOGUE OF USEFUL PLANTS OF COLOMBIA 224 Zamia muricata Willd. (Zamiaceae) IPNI: 270552-2 Habit: Herb Use: EU Origin: Native Region: Caribbean Dept.: BOY, LAG, SAN Elev.: 0-700 m.a.s.l. IUCN Habitat code: 1,4 Conservation status: (IUCN) NT (CPSG) EN 225 Zamia obliqua A.Braun (Zamiaceae) IPNI: 297347-1 Common name: chigua, maicito, zamia Habit: Shrub Use: HF Origin: Native Region: Pacific Dept.: ANT, CHO, VAC Elev.: 0-600 m.a.s.l. IUCN Habitat code: 1,3 Conservation status: (IUCN) NT (CPSG) VU 226 Zamia poeppigiana Mart. & Eichler (Zamiaceae) IPNI: 270556-2 Use: HF IUCN Habitat code: 1 Conservation status: (IUCN) NT 227 Zamia roezlii Regel ex Linden (Zamiaceae) IPNI: 297372-1 Common name: chigua Habit: Shrub Use: EU, HF Origin: Native Region: Pacific Dept.: CHO, NAR, VAC Elev.: 0-200 m.a.s.l. IUCN Habitat code: 1 Conservation status: (IUCN) NT (CPSG) VU Ginkgoopsida Ginkgoales 228 Ginkgo biloba L. (Ginkgoaceae) IPNI: 262125-1 Common name: ginkgo biloba Use: EU, HF, MA, ME Conservation status: (IUCN) EN Gnetopsida Gnetales 229 Gnetum leyboldii Tul. (Gnetaceae) IPNI: 111368-2 Common name: plátano, purgante de picón Habit: Liana Use: HF Origin: Native Region: Amazonia, Orinoquia, Pacific Dept.: AMA, ANT, CAQ, CHO, GUV, MET, PUT, VAU, VID Elev.: 40–770 m a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) LC 230 Gnetum nodiflorum Brongn. (Gnetaceae) IPNI: 111373-2 Habit: Climbing Use: HF Origin: Native Region: Amazonia, Guiana Shield Dept.: AMA, CAQ, GUA, MET, VAU, VID Elev.: 100–250 m a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) LC 231 Gnetum paniculatum Spruce ex Benth. (Gnetaceae) IPNI: 111375-2 Common name: reventilla Habit: Climbing Use: HF Origin: Native Region: Amazonia Dept.: AMA, GUA Elev.: 150–240 m a.s.l. IUCN Habitat code: 1 Conservation status: (IUCN) LC 179 CHECKLIST OF USEFUL PLANTS OF COLOMBIA 232 Gnetum schwackeanum Taub. ex Schenck (Gnetaceae) IPNI: 111377-2 Habit: Climbing Use: HF Origin: Native Region: Amazonia, Guiana Shield, Orinoquia Dept.: AMA, CAQ, GUA, VID Elev.: 100–250 m a.s.l. IUCN Habitat code: 1,3 Conservation status: (IUCN) LC 233 Gnetum urens (Aubl.) Blume (Gnetaceae) IPNI: 383578-1 Habit: Climbing Use: HF Origin: Native Region: Amazonia, Guiana Shield Dept.: AMA, CAQ, PUT, VID Elev.: 120–300 m a.s.l. IUCN Habitat code: 1,3,5 Conservation status: (IUCN) LC Pinopsida Pinales 234 Araucaria angustifolia (Bertol.) Kuntze (Araucariaceae) IPNI: 17518-2 Common name: araucaria, araucaria crespa Habit: Tree Use: EU, HF, MA, ME Origin: Cultivated Region: Andean Dept.: ANT, BOG, CAL, CUN, QUI, RIS Elev.: 1200–2750 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) CR 235 Araucaria araucana (Molina) K.Koch (Araucariaceae) IPNI: 261681-1 Habit: Tree Use: EU, HF, MA, ME Origin: Cultivated Region: Andean Dept.: ANT, BOG Elev.: 2000–2500 m a.s.l. IUCN Habitat code: 1,3 Conservation status: (IUCN) EN 236 Araucaria columnaris (G.Forst.) Hook. (Araucariaceae) IPNI: 261688-1 Use: EU, HF, MA, ME Region: Andean Dept.: BOG IUCN Habitat code: 1,3,14 Conservation status: (IUCN) LC 237 Araucaria heterophylla (Salisb.) Franco (Araucariaceae) IPNI: 676669-1 Habit: Tree Use: EU, HF, MA, ME Origin: Cultivated Region: Andean, Caribbean Dept.: ANT, ATL, BOG, CUN, HUI, SAP, TOL Elev.: 10–2750 m a.s.l. IUCN Habitat code: 3,14 Conservation status: (IUCN) VU 238 Chamaecyparis lawsoniana (A.Murray bis) Parl. (Cupressaceae) IPNI: 261839-1 Use: EU, MA, ME Region: Andean Dept.: BOG IUCN Habitat code: 14 Conservation status: (IUCN) NT 239 Cryptomeria japonica (Thunb. ex L.f.) D.Don (Cupressaceae) IPNI: 261870-1 Habit: Tree Use: EU, HF, MA, ME Origin: Cultivated Region: Andean Dept.: ANT, BOG, CUN Elev.: 1500– 2600 m a.s.l. IUCN Habitat code: 14 Conservation status: (IUCN) NT 240 Cupressus sempervirens L. (Cupressaceae) IPNI: 261974-1 Common name: ciprés, pino Habit: Tree Use: EU, FU, MA, ME, PO Origin: Cultivated Region: Andean Dept.: ANT, BOG, CUN Elev.: 1300–2700 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) LC 180 241 Hesperocyparis arizonica (Greene) Bartel (Cupressaceae) IPNI: 60451546-2 Habit: Tree Use: EU, FU, MA, ME, SU Origin: Cultivated Region: Andean Dept.: ANT Elev.: 1500–2000 m a.s.l. Conservation status: (IUCN) NE 242 Hesperocyparis lusitanica (Mill.) Bartel (Cupressaceae) IPNI: 60451554-2 Common name: pino común Habit: Tree Use: EU, HF, MA, ME Origin: Cultivated Region: Andean Dept.: ANT, BOG, CHO, CUN, QUI, RIS, TOL, VAC Elev.: 900– 3000 m a.s.l. IUCN Habitat code: 3,14 Conservation status: (IUCN) NE 243 Hesperocyparis macrocarpa (Hartw.) Bartel (Cupressaceae) IPNI: 60451558-2 Habit: Tree Use: EU, FU, MA, ME, PO Origin: Cultivated Region: Andean Dept.: BOY Elev.: 3600–3600 m a.s.l. Conservation status: (IUCN) NE 244 Platycladus orientalis (L.) Franco (Cupressaceae) IPNI: 677124-1 Common name: pino, pino libro Habit: Tree Use: EU, HF, MA, ME Origin: Cultivated Region: Andean, Caribbean Dept.: BOG, CUN, SAP Elev.: 2620–2620 m a.s.l. IUCN Habitat code: 3,14 Conservation status: (IUCN) NT 245 Sequoia sempervirens (D.Don) Endl. (Cupressaceae) IPNI: 263741-1 Habit: Tree Use: EU, MA, ME Origin: Cultivated Region: Andean Dept.: ANT, BOG Elev.: 2475–2475 m a.s.l. IUCN Habitat code: 14 Conservation status: (IUCN) EN 246 Thuja occidentalis L. (Cupressaceae) IPNI: 263956-1 Common name: pino, tuya, tuya del Canadá Habit: Tree Use: EU, HF, MA, ME Origin: Cultivated Region: Andean Dept.: ANT, VAC Elev.: 900–2100 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) LC 247 Pinus ayacahuite Ehrenb. ex Schltdl. (Pinaceae) IPNI: 314973-2 Use: MA Region: Andean Dept.: BOG Conservation status: (IUCN) LC 248 Pinus caribaea Morelet (Pinaceae) IPNI: 314859-2 Common name: pino Habit: Tree Use: EU, GS, HF, MA, ME Origin: Cultivated Region: Andean, Caribbean Dept.: ANT, CAS, MET, VID Elev.: 40–2300 m a.s.l. IUCN Habitat code: 3,14 Conservation status: (IUCN) LC 249 Pinus montezumae Lamb. (Pinaceae) IPNI: 263125-1 Use: MA, ME Region: Andean Dept.: BOG IUCN Habitat code: 3,14 Conservation status: (IUCN) LC 250 Pinus mugo Turra (Pinaceae) IPNI: 677083-1 Use: EU, HF, MA, ME Region: Andean Dept.: BOG Conservation status: (IUCN) LC CATALOGUE OF USEFUL PLANTS OF COLOMBIA 251 Pinus patula Schiede ex Schltdl. & Cham. (Pinaceae) IPNI: 314961-2 Common name: barbas de chivo, pino, pino espátula, pino pátula Habit: Tree Use: EU, FU, HF, MA, ME Origin: Cultivated Region: Andean Dept.: ANT, BOG, CAL, CAU, CUN, HUI, NAR, NSA, PUT, QUI, RIS, SAN, TOL, VAC Elev.: 2000–3000 m a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) LC 252 Pinus pinaster Aiton (Pinaceae) IPNI: 263216-1 Common name: pino marítimo, pino negral Use: EU, HF, MA, ME Dept.: BOG IUCN Habitat code: 14 Conservation status: (IUCN) LC 253 Pinus radiata D.Don (Pinaceae) IPNI: 30020085-2 Common name: pino, pino candelabro, pino ocarpo Habit: Tree Use: EU, HF, MA, ME Origin: Cultivated Region: Andean Dept.: BOG, CUN Elev.: 2600–3000 m a.s.l. IUCN Habitat code: 1,3,5,14 Conservation status: (IUCN) EN 254 Pinus sylvestris L. (Pinaceae) IPNI: 263353-1 Common name: pino Use: EU, FU, HF, MA, ME, SU Conservation status: (IUCN) LC 255 Pectinopitys harmsiana (Pilg.) C.N.Page (Podocarpaceae) IPNI: 77203787-1 Common name: chaquiro, pino, pino colombiano, pino negro Habit: Tree Use: EU Origin: Native, Cultivated Region: Andean Dept.: CAU, MAG, QUI, RIS Elev.: 1800–2200 m a.s.l. Conservation status: (IUCN) NE 256 Podocarpus guatemalensis Standl. (Podocarpaceae) IPNI: 204582-2 Common name: ají, chaquiro, chaquiro dulce, pino, pino amarillo, pino chaquiro, pino colombiano Habit: Tree Use: MA Origin: Native Region: Andean, Pacific Dept.: ANT, CAQ, CAU, CHO, NAR, PUT, SAN Elev.: 0-1200 m.a.s.l. IUCN Habitat code: 1,3,5,14 Conservation status: (IUCN) LC (CPSG) VU 257 Podocarpus oleifolius D.Don (Podocarpaceae) IPNI: 263553-1 Common name: ají, chaquiro, hayuelo, pino, pino amarillo, pino chaquiro, pino colombiano, pino criollo, pino de peña, pino hayuelo, pino real, pino romerillo caleño, pino romerón Habit: Tree Use: HF, MA, ME Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CAL, CAU, CES, CHO, CUN, HUI, LAG, MAG, MET, NAR, NSA, PUT, QUI, RIS, SAN, TOL, VAC Elev.: 1900-3800 m.a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) LC (CPSG) VU 258 Podocarpus parlatorei Pilg. (Podocarpaceae) IPNI: 204598-2 Use: MA, ME Region: Andean Dept.: BOG Conservation status: (IUCN) NT 259 Retrophyllum rospigliosii (Pilg.) C.N.Page (Podocarpaceae) IPNI: 946471-1 Common name: chaquiro, pino, pino chaquiro, pino colombiano, pino de Pacho, pino de montaña, pino hayuelo, pino montañero, pino negro, pino romerillo, pino romerón Habit: Tree Use: EU Origin: Native, Cultivated Region: Andean Dept.: ANT, BOG, CUN, HUI, CHECKLIST OF USEFUL PLANTS OF COLOMBIA MAG, NSA, QUI, SAN Elev.: 1200–3750 m a.s.l. Conservation status: (IUCN) VU Magnoliopsida Nymphaeales 260 Cabomba caroliniana A.Gray (Cabombaceae) IPNI: 273581-2 Common name: majate Habit: Herb, Aquatic Use: EU, ME Origin: Cultivated Region: Andean Dept.: ANT Elev.: 1500–1500 m a.s.l. IUCN Habitat code: 2,4 Conservation status: (IUCN) NE 261 Nymphaea amazonum Mart. & Zucc. (Nymphaeaceae) IPNI: 605491-1 Habit: Herb, Aquatic Use: HF, ME Origin: Native Region: Andean, Caribbean Dept.: CAU, COR, CUN, MAG, TOL, VAC Elev.: 6–1600 m a.s.l. IUCN Habitat code: 1,2,14 Conservation status: (IUCN) NE 262 Nymphaea ampla (Salisb.) DC. (Nymphaeaceae) IPNI: 605494-1 Habit: Herb, Aquatic Use: HF, ME Origin: Native Region: Andean, Caribbean, Pacific Dept.: CHO, COR, CUN, MAG Elev.: 100–1200 m a.s.l. IUCN Habitat code: 1,2,3,5,14 Conservation status: (IUCN) NE 263 Nymphaea elegans Hook. (Nymphaeaceae) IPNI: 171178-2 Common name: loto Habit: Herb, Aquatic Use: HF, ME Origin: Naturalised Region: Andean Dept.: ANT, CUN Elev.: 1600–2125 m a.s.l. IUCN Habitat code: 3,14 Conservation status: (IUCN) NE 264 Nymphaea glandulifera Rodschied (Nymphaeaceae) IPNI: 281434-2 Habit: Herb, Aquatic Use: EU, MA Origin: Native Region: Pacific Dept.: CHO Elev.: 150–150 m a.s.l. IUCN Habitat code: 1 Conservation status: (IUCN) NE 265 Nymphaea jamesoniana Planch. (Nymphaeaceae) IPNI: 605588-1 Habit: Herb, Aquatic Use: HF Origin: Native Region: Caribbean, Pacific Dept.: ANT, ATL, CHO Elev.: 20–200 m a.s.l. IUCN Habitat code: 14 Conservation status: (IUCN) NE 266 Nymphaea lotus L. (Nymphaeaceae) IPNI: 605604-1 Habit: Herb, Aquatic Use: AF, EU, GS, HF, MA, ME, SU Origin: Cultivated, Naturalised Region: Andean, Pacific Dept.: ANT, CUN, HUI Elev.: 110–1600 m a.s.l. IUCN Habitat code: 1,2,3,14 Conservation status: (IUCN) LC 267 Nymphaea pulchella DC. (Nymphaeaceae) IPNI: 605675-1 Common name: torta Habit: Herb, Aquatic Use: ME Origin: Native Region: Andean, Caribbean, Pacific Dept.: ANT, CHO, SUC Elev.: 0–180 m a.s.l. IUCN Habitat code: 2,14 Conservation status: (IUCN) NE 268 Nymphaea rudgeana G.Mey. (Nymphaeaceae) IPNI: 281442-2 Habit: Herb, Aquatic Use: ME Origin: Native Region: Caribbean Dept.: MAG Elev.: 50–50 m a.s.l. Conservation status: (IUCN) NE 269 Victoria amazonica (Poepp.) Klotzsch (Nymphaeaceae) IPNI: 605776-1 Common name: victoria regia Habit: Herb, Aquatic Use: EU, HF, MA Origin: Native Region: Amazonia Dept.: AMA Elev.: 180–360 m a.s.l. IUCN Habitat code: 1,5,14 Conservation status: (IUCN) NE Austrobaileyales 270 Illicium verum Hook.f. (Schisandraceae) IPNI: 554553-1 Common name: anís estrella, anís estrellado Use: HF, MA, ME Conservation status: (IUCN) NE Canellales 271 Drimys granadensis L.f. (Winteraceae) IPNI: 84100-2 Common name: ají, ají de páramo, bocadillo, canela de páramo, canelo, canelo de páramo, canelón, cupis, palo de ají, quijón, quinón, quirón, quiñón Habit: Small tree, Tree Use: EU, HF, ME Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CAL, CAQ, CAU, CUN, HUI, NAR, NSA, PUT, QUI, RIS, SAN, TOL, VAC Elev.: 1800–3900 m a.s.l. IUCN Habitat code: 1,3,4,5,8,14 Conservation status: (IUCN) NE 272 Drimys winteri J.R.Forst. & G.Forst. (Winteraceae) IPNI: 330748-2 Common name: ajicillo, ajizuelo, ají, ají de páramo, boighe, boique, canela, canelo, canelo de monte, canelo de páramo, cascarilla, cupis, foiye, fuñe, fuñe boighe, palo de ají, quinón Use: EU, HF, MA, ME IUCN Habitat code: 1,4 Conservation status: (IUCN) LC Chloranthales 273 Hedyosmum angustifolium (Ruiz & Pav.) Solms (Chloranthaceae) IPNI: 167832-1 Habit: Tree Use: HF Origin: Native Region: Andean Dept.: NAR Elev.: 900–2990 m a.s.l. IUCN Habitat code: 1,4 Conservation status: (IUCN) LC 274 Hedyosmum anisodorum Todzia (Chloranthaceae) IPNI: 277965-2 Habit: Shrub, Small tree, Tree Use: HF, ME Origin: Native Region: Andean Dept.: HUI, NAR, PUT Elev.: 1000–2800 m a.s.l. Conservation status: (IUCN) LC IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) NE 277 Hedyosmum cuatrecazanum Occhioni (Chloranthaceae) IPNI: 297228-2 Common name: canelo, canelón, chuchuguasa, granizo, guayusa Habit: Shrub, Small tree, Tree Use: HF Origin: Native Region: Andean Dept.: ANT, BOY, CAL, CAU, CUN, HUI, MAG, NAR, PUT, QUI, RIS, SAN, VAC Elev.: 1400-3700 m.a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) LC (CPSG) Potential LC 278 Hedyosmum cumbalense H.Karst. (Chloranthaceae) IPNI: 167850-1 Common name: granizo Habit: Shrub, Small tree, Tree Use: HF, ME Origin: Native Region: Andean Dept.: ANT, BOG, CAU, HUI, NAR, PUT, QUI, RIS, VAC Elev.: 1950–3800 m a.s.l. IUCN Habitat code: 1,3,4,8,14 Conservation status: (IUCN) LC 279 Hedyosmum goudotianum Solms (Chloranthaceae) IPNI: 277980-2 Common name: chiflador, granizo, silva silva, silvador, árnica de árbol Habit: Shrub, Small tree, Tree Use: HF, ME Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CAL, CAU, CHO, CUN, HUI, MET, NAR, NSA, PUT, QUI, RIS, SAN, TOL, VAC Elev.: 1050-3300 m.a.s.l. IUCN Habitat code: 1,3,4,8,14 Conservation status: (IUCN) LC (CPSG) Potential LC 280 Hedyosmum luteynii Todzia (Chloranthaceae) IPNI: 277988-2 Common name: borracho, cashco, granecillo, granizo, guayusa andina, majagua, pururug colorado, tarqui Habit: Shrub, Small tree, Tree Use: HF, ME Origin: Native Region: Andean Dept.: CAU, HUI, NAR, QUI, RIS, TOL, VAC Elev.: 2600–3600 m a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) NE 281 Hedyosmum maximum (Kuntze) K.Schum. (Chloranthaceae) IPNI: 118340-2 Use: HF IUCN Habitat code: 14 Conservation status: (IUCN) VU 282 Hedyosmum mexicanum C.Cordem. (Chloranthaceae) IPNI: 167876-1 Use: HF Origin: Native Conservation status: (IUCN) VU 275 Hedyosmum bonplandianum Kunth (Chloranthaceae) IPNI: 277969-2 Common name: aguanoso, almizcle, anisillo, canelo, canelón, chiflachifla, chocó, colchón de pobre, gallinazo, granicillo, granizo, guayusa, hurón, llorón, motilón, planta de soldado, planta de soledad, planta del soldado, silbador, silbasilba, silbato, silbo silbo, si Habit: Shrub, Small tree, Tree Use: HF, ME Origin: Native Region: Andean Dept.: ANT, BOG, CAU, CHO, QUI, RIS, VAC Elev.: 900–2400 m a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) LC 283 Hedyosmum racemosum (Ruiz & Pav.) G.Don (Chloranthaceae) IPNI: 167884-1 Common name: canelo, chiflichifli, chuchuguasa, granizo, guayusa, hojas de granizo, palo de agua, paloepiedra, pito, silbosilbo Habit: Shrub, Small tree, Tree Use: HF Origin: Native Region: Amazonia, Andean, Guiana Shield, Orinoquia Dept.: ANT, BOG, BOY, CAL, CAQ, CAU, CES, CUN, GUV, HUI, MAG, MET, NAR, NSA, PUT, QUI, RIS, SAN, TOL, VAC Elev.: 2503350 m.a.s.l. IUCN Habitat code: 1,2,3,4,8,14 Conservation status: (IUCN) LC (CPSG) Potential LC 276 Hedyosmum crenatum Occhioni (Chloranthaceae) IPNI: 118320-2 Common name: aguanoso, granizo, silbasilba, silva silva Habit: Shrub, Subshrub, Tree Use: ME Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CUN, LAG, MAG, MET, NSA, SAN Elev.: 1990–3700 m a.s.l. 284 Hedyosmum scaberrimum Standl. (Chloranthaceae) IPNI: 167886-1 Common name: granizo, rodilla de pollo, zancazanca Habit: Shrub, Tree Use: HF Origin: Native Region: Andean, Pacific Dept.: ANT, CHO, NAR, VAC Elev.: 0– CATALOGUE OF USEFUL PLANTS OF COLOMBIA 181 CHECKLIST OF USEFUL PLANTS OF COLOMBIA 2150 m a.s.l. IUCN Habitat code: 1,3,5,14 Conservation status: (IUCN) LC 285 Hedyosmum scabrum (Ruiz & Pav.) Solms (Chloranthaceae) IPNI: 167887-1 Habit: Shrub, Tree Use: MA Origin: Native Region: Andean Dept.: NAR, PUT Elev.: 2090–3130 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) LC 286 Hedyosmum sprucei Solms (Chloranthaceae) IPNI: 277998-2 Common name: granizo Habit: Shrub, Small tree Use: HF Origin: Native Region: Andean Dept.: CAU, PUT Elev.: 630–1050 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) LC 287 Hedyosmum translucidum Cuatrec. (Chloranthaceae) IPNI: 118351-2 Common name: aguaquín, granizo, silbasilba Habit: Shrub, Small tree, Tree Use: HF, ME Origin: Native Region: Andean Dept.: ANT, BOG, BOY, CAQ, CAU, CUN, HUI, MET, NAR, NSA, PUT, VAC Elev.: 2000– 3100 m a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) LC 294 Aniba affinis (Meisn.) Mez (Lauraceae) IPNI: 13808-2 Habit: Tree Use: MA Origin: Native Region: Amazonia, Guiana Shield Dept.: CAQ Elev.: 100–500 m a.s.l. Conservation status: (IUCN) LC 295 Aniba burchellii Kosterm. (Lauraceae) IPNI: 13815-2 Habit: Tree Use: MA Origin: Native Region: Amazonia, Guiana Shield Dept.: VAU Elev.: 100–200 m a.s.l. IUCN Habitat code: 1 Conservation status: (IUCN) LC Laurales 296 Aniba canelilla (Kunth) Mez (Lauraceae) IPNI: 13816-2 Common name: azafrás, canela de Quijos, canela muena, canelo, canelo de Andaquíes, cáscara, cáscara preciosa, sasafrás, tuabe Habit: Tree Use: HF, MA, ME, PO Origin: Native Region: Amazonia Dept.: CAQ Elev.: 400–400 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) LC 288 Gyrocarpus americanus Jacq. (Hernandiaceae) IPNI: 170637-1 Common name: banco, volador Habit: Tree Use: FU, MA, ME, SU Origin: Native Region: Caribbean Dept.: ANT, ATL, BOL, COR, LAG, MAG, SAN, SUC Elev.: 5–1000 m a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) LC 297 Aniba coto (Rusby) Kosterm. (Lauraceae) IPNI: 13822-2 Common name: cordillero, laurel, laurel comino Habit: Tree Use: MA, ME Origin: Native Region: Andean Dept.: QUI, TOL, VAC Elev.: 1200–2600 m a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) NE 289 Hernandia didymantha Donn.Sm. (Hernandiaceae) IPNI: 120657-2 Common name: aguacatón, banco, musana, palo blanco, tambor Habit: Tree Use: MA Origin: Native Region: Pacific Dept.: ANT, CHO Elev.: 0–100 m a.s.l. IUCN Habitat code: 1,3,5,14 Conservation status: (IUCN) NT 298 Aniba hostmanniana (Nees) Mez (Lauraceae) IPNI: 13841-2 Common name: amarillo, canelo Habit: Tree Use: MA, PO Origin: Native Region: Amazonia, Andean Dept.: ANT, MET Elev.: 100–1200 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) LC 290 Hernandia sonora L. (Hernandiaceae) IPNI: 316357-2 Common name: hernandia Use: MA, ME Conservation status: (IUCN) NE 291 Sparattanthelium amazonum Mart. (Hernandiaceae) IPNI: 170915-1 Habit: Shrub, Climbing Use: ME Origin: Native Dept.: ANT Elev.: 120– 600 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 292 Sparattanthelium glabrum Rusby (Hernandiaceae) IPNI: 241044-2 Common name: canelón Habit: Liana Use: ME Origin: Native Region: Amazonia Dept.: AMA, COR, CUN, NAR Elev.: 200–750 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) NE 293 Aiouea montana (Sw.) R.Rohde (Lauraceae) IPNI: 77173628-1 Common name: aguacatillo, amarillo laurel, chambolán de lo frío, chaviaco, comino, guacharaco, gualpante, jigua, jigua laurel, laurel, laurel colorado, muena, opachiro, queso Habit: Tree Use: ME Origin: 182 Native Region: Amazonia, Andean, Caribbean, Pacific Dept.: AMA, ANT, BOL, CAL, CAU, CUN, HUI, LAG, MAG, NAR, QUI, RIS, VAC Elev.: 0-2600 m.a.s.l. IUCN Habitat code: 1,2,3,4,5,14 Conservation status: (IUCN) LC (CPSG) Potential LC 299 Aniba panurensis (Meisn.) Mez (Lauraceae) IPNI: 13863-2 Common name: amarillo, amarillo oloroso, comino, laurel, laurel oloroso, loiro, mediocomino, miratava, palo rosa, yema de huevo Habit: Tree Use: MA, ME Origin: Native Region: Amazonia, Andean, Guiana Shield, Orinoquia Dept.: AMA, CUN, MET, TOL, VAU, VID Elev.: 200–1000 m a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) LC 300 Aniba perutilis Hemsl. (Lauraceae) IPNI: 13869-2 Common name: caucho, chachajo, comino, comino crespo, comino real, laurel, laurel comino, medio comino, mediocomino, tuno Habit: Tree Use: MA, ME, SU Origin: Native Region: Andean Dept.: ANT, HUI, MET, SAN, VAC Elev.: 100-2600 m.a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) VU (CPSG) CR 301 Aniba puchury-minor (Mart.) Mez (Lauraceae) IPNI: 13872-2 Common name: amarillo corazón negro, canelo, guacharaco, jigua negro, laurel, laurel comino, tuabe, tuabe canelo, tuave Habit: Tree Use: MA, ME Origin: Native Region: Amazonia, Andean, Pacific Dept.: AMA, ANT, BOY, CAQ, MET, NAR, QUI, SAN, TOL, VAC Elev.: CATALOGUE OF USEFUL PLANTS OF COLOMBIA 200–2000 m a.s.l. IUCN Habitat code: 1,2,3,5,14 Conservation status: (IUCN) LC 302 Aniba riparia (Nees) Mez (Lauraceae) IPNI: 13878-2 Common name: canelo, comino Habit: Tree Use: MA, ME Origin: Native Region: Amazonia, Andean Dept.: AMA, ANT Elev.: 100–800 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) LC 303 Aniba rosodora Ducke (Lauraceae) IPNI: 13880-2 Common name: palo de rosa Habit: Tree Use: HF, MA, ME, PO Origin: Native Region: Amazonia Dept.: AMA Elev.: 100–500 m a.s.l. Conservation status: (IUCN) EN 304 Beilschmiedia alloiophylla (Rusby) Kosterm. (Lauraceae) IPNI: 3010840-1 Habit: Tree Use: MA, ME, PO Origin: Native Region: Andean, Pacific Dept.: MAG, VAC Elev.: 100–1900 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) LC 305 Beilschmiedia brasiliensis (Kosterm.) Kosterm. (Lauraceae) IPNI: 31042-2 Common name: abacachirana, aguacatillo, emege, emehe, uflé, uflé-aguacatillo, yurajicù Habit: Tree Use: HF, MA, ME Origin: Native Region: Amazonia Dept.: AMA Elev.: 100–200 m a.s.l. Conservation status: (IUCN) NE 306 Beilschmiedia pendula (Sw.) Hemsl. (Lauraceae) IPNI: 1057986-2 Common name: aguacate cimarrón, laurel Habit: Tree Use: MA, PO Origin: Native Region: Andean Dept.: ANT, NAR Elev.: 0–1700 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 307 Beilschmiedia sulcata (Ruiz & Pav.) Kosterm. (Lauraceae) IPNI: 31062-2 Use: HF, ME Origin: Native IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 308 Beilschmiedia tovarensis (Klotzsch & H.Karst. ex Meisn.) Sachiko Nishida (Lauraceae) IPNI: 1116029-2 Common name: aguacatillo, amarillo tara, laurel Habit: Tree Use: HF, ME Origin: Native Region: Andean Dept.: ANT, BOY, CAL, CAU, MAG, MET, VAC Elev.: 500–3000 m a.s.l. IUCN Habitat code: 1,3,4,14 Conservation status: (IUCN) LC 309 Caryodaphnopsis cogolloi van der Werff (Lauraceae) IPNI: 274159-2 Common name: yumbé, yumbé aguacatillo y zumbí Habit: Tree Use: MA Origin: Endemic Dept.: ANT Elev.: 300-500 m.a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) EN (CPSG) EN 310 Cassytha filiformis L. (Lauraceae) IPNI: 463202-1 Common name: bejuco de rema Habit: Herb, Parasitic Use: HF, MA, ME, PO, SU Origin: Native Region: Amazonia, Guiana Shield, Caribbean, Orinoquia, Pacific Dept.: ANT, CAQ, GUA, GUV, VAU, VID Elev.: 0– CHECKLIST OF USEFUL PLANTS OF COLOMBIA 1500 m a.s.l. IUCN Habitat code: 1,2,3,4,14 Conservation status: (IUCN) NE 311 Chlorocardium venenosum (Kosterm. & Pinkley) Rohwer, H.G.Richt. & van der Werff (Lauraceae) IPNI: 274529-2 Habit: Tree Use: ME, PO Origin: Native Region: Amazonia, Pacific Dept.: AMA, CAQ, PUT, VAC Elev.: 0–300 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) LC 312 Cinnamomum camphora (L.) J.Presl (Lauraceae) IPNI: 463336-1 Common name: alcanfor Use: EU, GS, HF, MA, ME, PO Conservation status: (IUCN) NE 313 Cinnamomum verum J.Presl (Lauraceae) IPNI: 463752-1 Common name: canela, canelo Habit: Tree Use: EU, FU, HF, MA, ME Origin: Cultivated Region: Amazonia, Orinoquia Dept.: CAQ, CAS, TOL Elev.: 200–800 m a.s.l. IUCN Habitat code: 1,3,14 Conservation status: (IUCN) NE 314 Damburneya martinicensis (Mez) Trofimov (Lauraceae) IPNI: 77158859-1 Habit: Tree Use: MA Origin: Native Region: Pacific Dept.: CHO Elev.: 0– 100 m a.s.l. IUCN Habitat code: 1,2,14 Conservation status: (IUCN) LC 315 Damburneya purpurea (Ruiz & Pav.) Trofimov (Lauraceae) IPNI: 77158867-1 Common name: amarillo, laurel Habit: Tree Use: MA, ME Origin: Native Region: Andean, Pacific Dept.: ANT, CAL, CAU, CHO, CUN, HUI, LAG, MAG, NSA, QUI, VAC Elev.: 100–2400 m a.s.l. IUCN Habitat code: 1,2,3,4,8,14 Conservation status: (IUCN) LC 316 Endlicheria anomala (Nees) Mez (Lauraceae) IPNI: 464481-1 Habit: Tree Use: MA, ME Origin: Native Region: Amazonia, Andean, Orinoquia Dept.: AMA, CAQ, MET, VAC, VAU, VID Elev.: 100–1400 m a.s.l. IUCN Habitat code: 1,3,4,5,14 Conservation status: (IUCN) LC 317 Endlicheria arenosa Chanderb. (Lauraceae) IPNI: 60440993-2 Habit: Tree Use: MA, ME, PO Origin: Native Region: Amazonia Dept.: AMA Elev.: 100–400 m a.s.l. IUCN Habitat code: 1,4 Conservation status: (IUCN) LC 318 Endlicheria canescens Chanderbali (Lauraceae) IPNI: 307011-2 Habit: Tree Use: MA Origin: Native Region: Amazonia, Andean Dept.: AMA, PUT Elev.: 100–1700 m a.s.l. IUCN Habitat code: 1,3 Conservation status: (IUCN) LC 319 Endlicheria citriodora van der Werff (Lauraceae) IPNI: 276551-2 Habit: Tree Use: MA, ME Origin: Native Region: Amazonia Dept.: AMA, VAU Elev.: 100–200 m a.s.l. IUCN Habitat code: 1 Conservation status: (IUCN) VU 320 Endlicheria dictifarinosa C.K.Allen (Lauraceae) IPNI: 90945-2 Habit: Tree Use: HF Origin: Native Region: Guiana Shield, Orinoquia Dept.: GUA, VID Elev.: 80–120 m a.s.l. IUCN Habitat code: 1 Conservation status: (IUCN) LC 321 Endlicheria multiflora (Miq.) Mez (Lauraceae) IPNI: 90972-2 Use: MA Origin: Native IUCN Habitat code: 3 Conservation status: (IUCN) LC 322 Endlicheria paniculata (Spreng.) J.F.Macbr. (Lauraceae) IPNI: 90974-2 Habit: Tree Use: MA Origin: Native Region: Andean Dept.: CUN, HUI, MAG, MET Elev.: 100–2000 m a.s.l. IUCN Habitat code: 1,14 Conservation status: (IUCN) LC 323 Endlicheria sericea Nees (Lauraceae) IPNI: 464535-1 Common name: amarillo co