Professional Documents
Culture Documents
Fungi of Colombia
Edited by
Rafael F. de Almeida
Robert Lücking
Aída Vasco-Palacios
Ester Gaya
Mauricio Diazgranados
Catalogue of
Fungi of Colombia
Catalogue of
Fungi of Colombia
Edited by
Rafael F. de Almeida
Robert Lücking
Aída Vasco-Palacios
Ester Gaya
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
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Designs and Patents Act 1988.
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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 978-1-84246-790-9
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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: Fungi mosaic made from photographs included in this book, based on the photograph of the species Cyptotrama asprata
(Basidiomycota) by Nataly Gómez-Montoya. Design cover by R.F. Almeida; back cover background photo credit: Ganoderma (Mauricio Diazgranados
© RBG Kew); back cover fungi photo credits: Lepidostroma calocerum (Robert Lücking ©), Tetrapyrgos alba (Nataly Gómez-Montoya © RBG Kew),
Pleurotus djamor (Nataly Gómez-Montoya © RBG Kew), Cordyceps nidus (Robert Lücking ©), Dacryopinax spathularia (Kent Brothers © RBG Kew),
Cyathus striatus (Aída Marcela Vasco-Palacios ©), Aseroe rubra (Nataly Gómez-Montoya © RBG Kew), Xylaria multiplex (Robert Lücking ©), Phyllobaeis
imbricata (Robert Lücking ©), Cladonia secundana (Robert Lücking ©)
Preface 7
Executive Summary 11
Foreword 13
Acknowledgements 15
List of Contributors 17
Chapter 1 The Useful Plants and Fungi of Colombia (UPFC) project: delivering mycological 19
knowledge to support conservation and sustainable development
Chapter 11 Diversity, Functional Groups, and Community Structure of Fungi of Colombia 139
Checklist 216
Laminae 427
Fungi have played an essential role in shaping the Earth’s of botany in academia until 1969, when the North American
biosphere for the past one billion years. They might have ecologist Robert Whittaker proposed that they should be
been one of the few life forms inhabiting landmasses over placed in their own kingdom, separate from plants. Twenty
500 million years ago, perhaps associated with terrestrial years later, the first DNA-based studies of fungi revealed that
algae, even before early land plant lineages arose. Lacking fungi are indeed more closely related to animals than plants.
roots, early plants may have relied on fungal partners for The knowledge on fungi accumulated during the past
endophytic mycorrhiza-like associations, allowing them to 500 years led to ground-breaking discoveries for modern
thrive on primordial mineral soil. In a process known as science, such as antibiotics or the use of fungi in food
biological weathering, fungal hyphae would secrete organic production and biotechnology. Nonetheless, most of this
acids to dissolve rocks and extract nutrients held within, knowledge was based on temperate fungas from Europe
and for water acquisition. In return, plants would transfer or North America. The diversity of Neotropical funga and
carbohydrates produced through photosynthesis to their their potential uses started to be uncovered only in the
fungal partners. Widespread today in associations known 1800s when European nations began to study the natural
as mycorrhizae or lichens, this exchange of resources resources of their colonies in the New World. Since then,
between fungi and early land plants or other photosynthetic several floras have been published for different countries
organisms helped to ignite the Earth’s biosphere growth, in Central and South America or for larger regions, such
evolution, and diversification into ever more complex as Mesoamerica, the Caribbean, or the entire Neotropics,
organisms, communities, and ecosystems. Nowadays, over which in some cases covered groups of fungi such as
ninety per cent of known land plants associate with at least lichens. Yet, compared to vascular plant and even bryophyte
one fungal partner, and some are entirely dependent on floras, megadiverse countries in South America, including
fungal assistance to survive. The symbiotic rise of land Colombia, Venezuela, Ecuador, Peru, and Bolivia, still lack
plants and their fungal partners also had dramatic effects any comprehensive taxonomic study of their funga.
on our atmosphere, with the increased absorption of Through the Catalogue of Fungi of Colombia, we want to
carbon dioxide by plants over millions of years producing showcase the significant progress that has resulted from
a massive rise in oxygen concentrations, supporting the a partnership between Colombia and the UK (see Chapter
emergence of much larger, more complex animal life forms, 1). These joint efforts enabled us to compile the first broad
from early tetrapods to humans. assessment on the history of mycology in the country
Fungi have also directly impacted the daily life of human (Chapter 2), expand on the general diversity of the Colombian
society along with their cultural evolution during the past
funga (Chapter 3) and on the diversity of the main groups
300,000 years, being used as food, for ritualistic purposes,
of true fungi (Chapters 4–9), assess the biogeography
as medicinal sources, or by causing diseases affecting
of selected taxa (Chapter 10), and elaborate on fungal
our crops, livestock, and own bodies. Nonetheless, the
ecology (Chapter 11), uses (Chapter 12), biotechnological
documented history of the uses and impact of fungi through
potential (Chapter 13), conservation (Chapter 14), and
time within human societies is greatly fragmented, not the
biological collections (Chapter 15). The first checklist of
least because there have always been major doubts regarding
fungi and fungi-like organisms for Colombia (Chapter 16) is
their nature. In medieval European societies, naturalists
complemented by indexes of families, genera, synonyms,
diverged in assessing the relationships of fungi with other life
misapplied and dubious names, and a laminae section with
forms, some considering them closer to animals and others
photographs of selected taxa for proper illustration.
as a “mixture” of animal and algal traits, best positioned
within plants. It was only in 1728 that Antoine Jussieu read
a paper to the Paris Academy stating the urgent need to USING THIS CATALOGUE AND THE COLFUNGI ONLINE PORTAL
recognise fungi as a section of plants. Since then, even if Free access to a large amount of information on the diversity
their separate nature had been well established at the turn of fungi in Colombia is available to everyone for the first
of the 19th century, fungi remained confined as a section time for over 7,000 accepted species. Accessing the web
portal from this Catalogue is made easy by clicking on the while promoting the exchange and transfer of knowledge
hyperlink on the species name or by reading the QR code of between researchers from the UK and Colombia.
the species, or by typing a scientific or common name of the
species, genus, family, or any keyword that describes the SHARING KNOWLEDGE FAIRLY
fungus or its uses, in the search bar on the homepage of By gathering data from different sources, ColFungi enables
colfungi.org access to valuable information, enriches the data and
The search results show all the relevant fungi species its potential uses, and maximises the visibility of local
profiles that the user can explore. The pages in each profile resources. Only publicly available content is used and, as
show the classification and status of the fungal name part of the content collection process, we aim to ensure that
(whether it is an accepted name, synonym, misapplied or publishers have respected the Nagoya Protocol (a part of the
dubious) and the habit, description, ecology, distribution, Convention on Biological Diversity that aims to ensure the
uses, common names, and conservation status when fair and equitable sharing of benefits obtained from the use
known and available. Images of fungi in the field and from of genetic resources), as well as Colombian laws regarding
fungarium specimens are also featured when available, the dissemination of traditional knowledge.
along with any other information about the species, as well ColFungi is part of Kew’s Plants of the World Online portal
as links to other sources. The ColFungi portal has advanced (plantsoftheworldonline.org) and therefore offers structured
search and filtering options tailored to the fungi of Colombia. content adjusted to standards that facilitate the distribution
Advanced search allows users to build customised queries and exchange of information with global data resources. It
on the data based on their interests. is also part of Kew’s initiative on Useful Plants and Fungi
The taxonomic backbone of ColFungi and the Catalogue (UPF), benefiting from UPF databases and its standards and
of Fungi of Colombia is based on the Index Fungorum applying them to fungi in Colombia. These actions make
(IF: indexfungorum.org) and Species Fungorum (SF: information about fungi of Colombia more widely available,
speciesfungorum.org), adjusted by consultation of recent both in Colombia and among the international community.
reference works, and will be updated over time. Other
metadata, such as distribution, have been extracted COLLABORATION AND CAPACITY BUILDING
from public occurrence repositories, such as the Global
ColFungi is building a strong collaborative network between
Biodiversity Information Facility (GBIF, gbif.org) and
the UK and Colombian researchers and institutions,
institutional and personal databases from digitised
including universities, fungal collections, non-governmental
Colombian herbaria/fungaria.
organisations, associations, and national institutes. The
project involves 37 researchers and collaborators from
INTRODUCING COLFUNGI TO YOU
Kew’s scientific departments, 35 researchers from the
Access to information on fungi is essential to expand our Asociación Colombiana de Micología (ASCOLMIC) and 25
knowledge of fungal diversity, allowing us to conserve Colombian partner institutions that cover most regions and
them, along with their habitats and ecosystems, and to cities, and international experts studying the Colombian
make sustainable use of them, achieving the benefits they funga from countries such as Brazil, Mexico, the USA,
provide to our societies. ColFungi is a free access online Canada, and Germany.
portal built by experts that offer accredited information on The Catalogue of Fungi of Colombia and ColFungi
Colombian fungi. It is a detailed resource that provides for gather and link information from other portals (such as
the first time high-quality taxonomic information along with the Information System on Biodiversity of Colombia - SIB
a wide range of data and information for all groups of fungi Colombia), the Catalogue of Plants and Lichens of Colombia,
in a single resource. the Global Biodiversity Information Facility (GBIF), Index
In order to have information on all known Colombian fungi Fungorum, databases from national and international
in a single web page, ColFungi also shares a large amount fungarium and strain collections, and data generated by
of information, such as herbarium/fungarium specimens, mycological researchers during their investigations, providing
species descriptions, maps, images, information on an additional window to the available data and expanding
traditional and potential uses and sustainable practices, the access to information on Colombian fungi. As a result,
and relevant literature and links when available. As a the visibility of all linked resources is expanded, providing
dynamic tool with a rapidly growing amount of information, it reciprocal benefits to RBG Kew, the Humboldt Institute,
provides a valuable resource for a wide variety of audiences the Asociación Colombiana de Micología (ASCOLMIC), and
and users, ranging from researchers and practitioners to other academic and environmental institutions in Colombia
legislators, teachers, and schoolchildren. and abroad.
ColFungi contributes directly to the objectives of Both the Catalogue and the portal provide opportunities
Colombian governmental organisations to develop national for Colombian mycologists and their international
and regional biodiversity catalogues while supporting the collaborators and volunteers and interns to contribute to the
transformation of the Colombian economy towards green project. The exchange of skills and experience has provided
growth. It also maximises the impact of scientific research great benefits to all the institutions involved and has helped
and highlights the value of fungal diversity in Colombia to strengthen capacities in Colombia.
ENRICHING INFORMATION
The value of species profiles in ColFungi is increased with
expanded data on the biology and ecology of the species when
available. To this end, different resources are used to access
information on the distribution, threats, uses, and availability
of the underlying resources, such as herbarium/fungarium
specimens and strain collections. These specimens are dried
fungi or parts thereof labelled with a wealth of information
about the species and its environment. Digitising specimens
includes assigning a unique barcode, capturing an image
of the sample and labels, and transcribing data from the
labels. A compendium of 7,241 species of Colombian fungi
has been currently recorded, most from Colombian herbaria,
fungaria, strain cultures, and international collections. How
many additional uncatalogued collections exist is unknown.
CONSERVATION STATUS
Extinction risk assessments depend on knowledge of
species distribution and the threats they face. The IUCN
Red List of Threatened Species Categories and Criteria data
available for fungi to date are assessed through the ColFungi
profiles and in this Catalogue. Also, some species are
placed in extinction risk categories according to the regional
or national lists of threatened species in Colombia.
USES
The Catalogue of Fungi of Colombia and ColFungi teams have
been collecting information on useful fungi from different
sources to enrich the content of the species profiles. The
data and images from these collections were compiled from
published sources and added to the species profiles to
provide a valuable information resource on the various uses
of Colombian fungi – from culinary to cultural.
Rafael F. de Almeida
Robert Lücking
Aída M. Vasco-Palacios
Ester Gaya
Mauricio Diazgranados
Editors
Colombia has a great comparative advantage over other In addition to the checklist, we have included a series
countries in being the second most biodiverse country of chapters written by specialists to provide some context
globally. Its biological resources are a key asset for for the actual knowledge about funga in Colombia. These
changes in its productive matrix. However, to plan a solid perspectives on the state of knowledge on the Fungi of
bioeconomic strategy, the first step is to consolidate the Colombia, covering the diversity of the main groups of fungi
current knowledge of the rich diversity of plants and fungi found in the country and the history of mycological studies
in the country. Arising from the collective efforts of the in Colombia, are presented alongside chapters on their
Alexander von Humboldt Biological Resources Research biogeography, biotechnology, conservation, ecology, uses,
Institute (Colombia), the Royal Botanic Gardens, Kew— and presence in national and international collections. The
under the Useful Plants and Fungi of Colombia Project supplementary materials that follow the checklist will allow
and Colombia Bio—and the mycological community in users, to explore open questions and opportunities in order to
Colombia, this Catalogue is presented as a source of develop new ideas on consumption/production alternatives
baseline information about the great diversity of fungal and pattern changes, species conservation, and new
species in Colombia, now linked and accessible through a technologies, to influence positive social and environmental
single online portal—ColFungi (Colombian Fungi Resources impacts, and to inspire new generations. Importantly, this
made Accessible). The Useful Plants and Fungi of Colombia compilation also points out the information gaps we have
Project Team and national and international mycologists on the diversity, conservation, and sustainable uses of the
have consolidated expert generated and accredited country’s fungal diversity, which will serve as the basis for
information on Colombian fungi. For the first time, high- planning actions to fill these gaps, thereby supporting the
quality and up-date taxonomic information is presented conservation and sustainable use of the Colombian funga
alongside data on uses, morphology, known geographic for the benefit of society.
distributions, habitats, life forms, vernacular names, and We hope that this Catalogue and ColFungi, as its dynamic
field images taken from the field by several mycologists who resource, will be used by the audience as a useful reference
have donated to the project. This goal required generating for decision-making, production, and demand building.
a list of the species that occur in Colombia and associating We expect a broad audience (from scientists, students,
all reported taxonomic information for those species from interested public, politicians, stakeholders, regulators,
all available databases and literature sources. For further producers, and consumers), who together can work to
details, check Chapter 16. allow Colombia as a Nation to achieve its Mission on Green
Tracking species as their names change through time and Growth, and to use and conserve its natural resources in an
maintaining links to their uses, distributions, conservation integrated, equitable and sustainable way.
status, and common names is an endless task. It is the
reason for linking the printed version of the Catalogue to a
dynamic online portal. We still need to gain scale on those
tasks. Still, we are confident that technological advances
in artificial intelligence and in curation fields will accelerate
both the taxonomy and the integration of traditional and
scientific knowledge needed to support alternative and
sustainable uses for Colombia’s plant and fungal resources
and to facilitate their conservation. It is also important to
remember that we currently know less than 5% of Colombia’s
expected diversity of fungi. For this reason, this is a base list
of species representing the funga of the country, but our
knowledge will increase over time, and this information will
be updated on the ColFungi portal.
Colombia houses some of the most diverse ecosystems of fungi and humanity. Medicinal, edible, tinctorial, and more
on earth. This diversity is not only of animals and plants uses are thoroughly described in this book, which is a delight
but also of fungi, the mysterious Kingdom of life that feeds, to read and a delight to look through.
heals, clothes, harms, and certainly delights us. In this But this project is not only about describing and
extraordinary, publication which represents the joint effort illustrating species. This project is about collaboration, and
of many scientists and researchers worldwide, you will be interconnection is the essence of fungi. This pioneering effort
submerged in the wonder and magic that the fungal Kingdom to document the species of fungi in Colombia and their uses
provides through concise data and superb photography. for human benefit forms the basis of an ongoing investment
However, what makes this edition so extraordinary is that in building platforms for digital access to information about
it not only presents the charismatic mushrooms and conks fungal species and their uses, as well as a digital platform
but, more uniquely and delightfully, it takes us on a deep dive to create and promote value chains that facilitate the
into the invisible world of yeasts, rusts, and smuts, as well sustainable use of fungi in Colombia. This information is not
as the intriguing and fundamental diversity of endophytes. only useful in the context of the difficulties that humanity
The latter is seldom shown in any books on fungi. is facing on earth today, but also as proof of how fungal
With over 7,200 species of macro- and microscopic fungi technologies discovered in the past hold the keys to the
found in Colombia listed in this book, you will have the most health and sustainability of people and the planet in the
in-depth induction into the funga of this unique part of the present and future. The virtuous symbiosis of international
world. This book will provide you with a unique view of how researchers from Kew, which houses the largest fungarium
species are intertwined and sometimes are used by their globally, and Colombian and Latin American researchers
plant and animal allies, including us humans. Through well- who know their habitats like no other can only be a two-
curated and analysed data that give you a sophisticated times virtuous collaboration. Although it seems obvious,
understanding of the role of fungi in Colombian ecosystems, these symbioses are not frequent and are seldom enabled
and most importantly, their role in ecosystems at large, your by many institutions at the same time. Therefore, this
appreciation for the fungi can only grow. With the data on endeavour is truly pioneering and reveals a multi-faceted
the conservation status of fungi—and detailed infographic view of the diversity of Colombian fungi.
illustrations that convey these powerful data--you will gain an Fungi are not exempt from or immune to the same threats
integral understanding of these extraordinary and fundamental animals and plants face. Habitat loss, habitat fragmentation,
organisms. But most importantly, this book highlights the overuse of pesticides and fungicides, and climate change
information gaps in our knowledge of the funga of Colombia affect the funga, fauna and flora equally. Understanding
and proposes a way forward for different groups of fungi. the world beneath our footsteps, a world in which fungi
The photography you will see in this book is astonishing. play symbiotic and decomposing roles, enables us to look
Rarely do we get to see so many species of fungi in such at forests, prairies, city parks and even your garden in a
perfect conditions and at such high magnification. Through different light. With this understanding comes care, and care
this book, you will observe structures rarely glimpsed triggers action. This book invites us to come together to
while walking through a forest. With every photograph, the protect the habitats that house this unique fungal diversity,
importance of these organisms for Colombian ecosystems and it is a call we should all answer.
are portrayed with delight, making the book a versatile
publication that is not only a guide for mycologists or people Giuliana Furci – Founder & CEO, Fungi Foundation
with a developed understanding of these organisms but also Harvard University Herbaria Associate
a guide for beginners and nature enthusiasts of all ages and Dame, Order of the Star of Italy
from anywhere and everywhere in the world. Co-Chair, IUCN SSC Fungal Conservation Committee
The way in which the ethnic groups and indigenous Advisory Board member – Society for the Protection of
peoples of Colombia use different species of fungi provide Underground Networks SPUN
a unique opportunity to delve into the cultural co-evolution Santiago, Chile, February 2022
The Catalogue of Fungi of Colombia is the result of the efforts numerous specialists from several countries who contributed
of countless people and institutions, representing one of to the elaboration or revision of the updated checklist or
the main products of the project Useful Plants and Fungi with valuable information on local or regional uses of the
of Colombia (UPFC), led by the Royal Botanical Gardens species. We also thank those colleagues who contributed to
Kew and the Alexander von Humboldt Biological Resources the book or its chapters with metadata or images, besides
Research Institute (IAvH) in Colombia. the various chapter authors also Álvaro Rúa, Ana Cristina
This book is substantially based on the ColFungi portal, Bolaños, and Gineth Adriana Calderón, and those who acted
developed and maintained by the Biodiversity Informatics as academic reviewers of the various chapters and made
and Spatial Analysis team (BISA) at RBG Kew. This portal valuable suggestions, especially Bianca R. da Hora, Celeste
uses several databases as primary sources, such as the Heisecke, Manuela Dal Forno, Patricia Fiuza, Raquel Negrão,
Catalogue of Plants and Lichens of Colombia (Bernal et al., Tiara Cabral, and Tom Prescott. Additionally, we thank
2019), the Global Biodiversity Information Facility (GBIF), Giuliana Furci for the wonderful foreword that conveys the
The Mycology Collections data Portal (MyCoPortal), Index importance of this book.
Fungorum, databases from Colombian fungal collections We would also like to thank our Project Manager David
(HUA - Ana Esperanza Franco, COL - Lauren Raz, ANDES-F, Hammond, Project Officer Kaitalin White, and Communication
and UBDC - Bibiana Moncada) and personal data generated Specialists Camila Gutierrez and Maria del Pilar Mira Ponton
by Colombian mycologists during years of research. Special for their enormous support of the Useful Plants and Fungi of
thanks to the BISA team, especially Joaquim d’Souza and Colombia project through to the entire process leading up
Malcolm Stone, for making all the information available from to the publication of the present Catalogue. Additionally, we
the ColFungi portal and providing the PURLs, which allowed would like to thank the successive directors and researchers
the production of the QR codes. The data central repository of the RBG Kew, IAvH, and the Ministry of Environment and
team, especially Bob Allkin, is also acknowledged for working Development Sustainable of Colombia. We hope that they
on consolidating the data and the taxonomic backbone. continue to support future mycological studies in Colombia.
Also, many thanks to those that provided invaluable Special thanks to all institutions and research groups that
support towards the compilation of the preliminary lists, dedicated their time and data to the project. Countless
references, specimens, common names, and all information people and institutions supported this Catalogue in different
related to species and their respective uses and conservation ways, and to all of them, our deepest gratitude.
status, integrating these metadata into the central data Finally, we are very grateful for the support of the funders
repository. This refers especially to Kew’s digitisation team regarding the production costs of this book, RBG Kew for
(Laura Green, Ellie Graves, and Priscila Reis), Julia Carretero, providing the research support and internal services, the
and the colleagues from IAvH (Alejandra Aguilar-Giraldo, IAvH for contributing field data and literature resources,
Daniel Jiménez-Pastrana, Lina Isabel Guevara-Ruiz, Henry all mycologists and their institutions/fungaria for their
Agudelo, and Fabio Ávila). A special thanks to the Asociación contribution of databases and images of fungal diversity and
Colombiana de Micología (ASCOLMIC) for the scientific for the curation of all catalogue records, and Kew Publishing
support and constant advice during the project execution. for the final copy editing and layout. This book was especially
We further thank the “Spatial Analysis and data” team supported by a Professional Development & Engagement
from RBG Kew for producing the species distribution maps grant under the Newton-Caldas Fund partnership. The
available in the ColFungi species profiles, especially Carolina project grant is funded by the UK Department for Business,
Tovar, Ian Ondo, and Justin Moat. We thank all staff, past and Energy and Industrial Strategy (BEIS) alongside Colombia’s
present, of their respective institutions for logistic support Minciencias and delivered by the British Council. For further
in developing the contents of this book, as well as the information, please visit www.newtonfund.ac.uk.
CONTRIBUTORS INSTITUTION
Adriana Corrales Universidad del Rosario, Colombia
Aída M. Vasco-Palacios Universidad de Antioquia, Colombia
Ana Esperanza Franco-Molano Universidad de Antioquia, Colombia
Anabel Barona-Colorado Universidad Santiago de Cali, Colombia
Angelis Marbello Universidad de los Andes, Colombia
Bibiana Moncada Universidad Distrital Francisco Jose Caldas, Colombia
Carolina Castellanos-Castro Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Colombia
Cristina Benjumea Universidad de Antioquia, Colombia
Dahyana Bolanos-Burbano Universidad Santiago de Cali, Colombia
David Diaz-Escandon University of Alberta, Canada
David Hammond Royal Botanic Gardens, Kew, UK
Diego Simijaca-Salcedo Universidad Autonoma de Aguascalientes, Mexico
Edier Soto-Medina Universidad del Valle, Colombia
Ester Gaya Royal Botanic Gardens, Kew, UK
Fabiola Eugenia Gonzalez-Cuellar Universidad de Cauca, Colombia
German Torres-Morales Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Colombia
Gregory Mueller Chicago Botanic Garden, USA
Joaquim de Souza Royal Botanic Gardens, Kew, UK
Julia Carretero Royal Botanic Gardens, Kew, UK
Kaitalin White Royal Botanic Gardens, Kew, UK
Kent Brothers Grupo Micólogos de Colombia, Colombia
Luis Fernando Coca Herbario Universidad de Caldas, Colombia
Marcela Guevara-Suárez Universidad de los Andes, Colombia
Margarita Jaramillo-Ciro Universidad de San Buenaventura, Colombia
Maria Camila Bados-Lopez Universidad Santiago de Cali, Colombia
Maria del Pilar Mira Royal Botanic Gardens, Kew, UK
Martha Cardenas Universidad de los Andes, Colombia
Martha L. Ortiz-Moreno Universidad de los Llanos, Colombia
Mauricio Diazgranados Royal Botanic Gardens, Kew, UK
Mauricio Ramírez-Castrillón Universidad Icesi, Colombia
Mauricio Salazar-Yepes Universidad Nacional de Colombia, Colombia
Meike Piepenbring Goethe University Frankfurt am Main, Germany
Melissa Palacio Universidad Federal do Rio Grande do Sul, Brazil
Natalia Bolaños Universidad de los Andes, Colombia
Natalia Vargas-Estupiñan Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Colombia
Nataly Gomez-Montoya Universidad de Antioquia, Colombia
Paul Kirk Royal Botanic Gardens, Kew, UK
Pedro Jimenez Universidad Militar Nueva Grana, Colombia
Rafael Felipe de Almeida Royal Botanic Gardens, Kew, UK
Robert Lucking Botanischer Garten, Germany
Rocío Peña-Cañon Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Colombia
Silvia Restrepo Universidad de los Andes, Colombia
Tatiana Rojas Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Colombia
Tatiana Sanjuan Grupo Micólogos de Colombia, Colombia
Tiziana Antonella Cossu Royal Botanic Gardens, Kew, UK
Tiziana Ulian Royal Botanic Gardens, Kew, UK
Viviana Motato-Vasquez Universidade del Valle, Colombia
Yeina Milena Nino Fernandez Universidad Pedagógica y Tecnológica de Colombia - Tunja, Colombia
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 socio-economic growth, which could
represent either a threat to Colombian biodiversity or an opportunity for sustainable development based on the country's
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 through 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 while 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,
developed pathways for tackling socio-environmental challenges and contributed 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 store of global carbon among other functions (Millennium
Our life and well-being rely on the environmental goods Ecosystem Assessment, 2005). However, the conversion
and services provided by plants and fungi. Not only do of natural habitats—primarily for farming—is fuelling land
they provide us with products, such as food, medicines, degradation, undermining the well-being of two-fifths of the
natural fibres, fuel, building materials, and cosmetics, human population, and raising the risk of migration and
but they also act to purify the air, enrich the soil, protect conflict (IPBES, 2018). Therefore, the preservation and
against erosion, regulate water flow and quality, and sustainable use of this biodiversity are essential to human
provide habitats for animals while acting as a significant well-being and future economic prosperity.
TABLE 1. Origin and conservation status of plant and fungal diversity in Colombia, based on data collected in this project.
Plants Useful plants Fungi Useful fungi Plants & fungi Useful Plants & fungi
Cultivated 1,077 1,077 21 N/A N/A N/A
Naturalised 677 638 20 N/A N/A N/A
Native 27,134 5,830 7,200 25 28,398 5,855
Endemic 9,473 478 203 8 9,679 486
Threatened 1,061 249 63 3 1,071 252
TABLE 2. Plant and fungal diversity in Colombia, based on data collected by this project (useful vs non-useful).
Plants Useful plants Fungi Useful fungi Plants & fungi Useful Plants & fungi
Species 28,947 7,472 7,241 374 36,087 7,790
Genera 3,583 2,140 1,790 192 5,346 2,318
Families 395 258 455 99 843 356
Colombia is one of the most biodiverse countries on As deforestation proceeds, the disappearance of traditional
Earth, hosting at least 28,947 plant species (24% endemic; knowledge on the conservation, use and management of
>1,000 threatened), 3,583 genera, and 395 families plants and fungi accelerates with biodiversity loss.
(Table 1). The country ranks third in the diversity of plants With its unique and extraordinary biocultural diversity
worldwide and is a centre of origin of tomatoes, peppers, and richness, Colombia can become an exemplar for
potatoes, chillies and many other crops and their wild achieving Sustainable Development Goals (SDGs) (United
relatives (Diazgranados et al., 2020, Negrão et al., 2022; Nations, 2015), by reducing inequality and poverty based
Khoury et al., 2014). The fungal diversity in Colombia is also on the sustainable use of its useful plants and fungi whilst
impressive, with at least 7,241 species belonging to 1,790 reducing the degradation of its natural resources. Although
genera and 455 families (Table 2). Given that the observed native non-crop plants and fungi have great potential to
ratio between fungi and plant species in well-studied areas improve livelihoods and to support economic development
is 9.8:1, Colombia could harbour up to 300,000 species, in the country, knowledge on the most useful native
of plants and fungi representing 9% of the global diversity species remains highly dispersed, largely inaccessible, and
(Gaya et al., 2021). susceptible to disappearing over time.
Colombia is also described as one of the most ethnically
diverse countries in the world, with 85 different ethnic CAN THE SUSTAINABLE USE OF BIODIVERSITY BECOME
groups and 68 recognised native languages. The knowledge PART OF THE SOLUTION?
associated with useful plants and fungi can be as rich as the Despite the unique biological and cultural richness of
cultural variety. The country has been considered a “cradle Colombia, vast social inequality and marked poverty in
for modern ethnobotany”, with a plethora of recent studies rural areas have triggered more than six decades of internal
on this topic (Bernal et al., 2011), although ethnobotanical conflict. Plant and fungal diversity and its associated
knowledge is still vastly under-documented in this region cultural knowledge could tackle socio-environmental
(Cámara-Leret et al., 2014). challenges by boosting rural employment and incomes,
Regrettably, the country lost just under six million hectares improving people’s livelihoods, reducing inequalities, and
of forest between 1990 and 2015, at an average rate of supporting peace. Improving access to critical biological
237,000 hectares per year (FAO, 2015). This deforestation information and the development of markets for natural
is mainly due to agricultural expansion, urban development, ingredients, while meeting sustainability benchmarks,
and illegal mining, driven by inadequate political decision- can boost the stake of marginalised communities in the
making, often not supported by science (Eufemia et al., 2019; national economy by adding value to their traditional
Sabater et al., 2017; Salazar et al., 2021). Land-use change knowledge. Simultaneously, this process would empower
is threatening Colombia’s impressive cultural and biological meaningful participation, leading ultimately to more stable
diversity, and researchers and conservationists are now in a and equitable socio-economic outcomes. This approach
race to protect it (Andrade-C., 2011; WWF-Colombia, 2017). tackles a number of SDGs, most notably those related to
poverty alleviation, health improvement, empowerment of 4. Apply a VCN framework to improve local communities’
the disenfranchised, sustainability of economic growth and livelihoods in three pilot areas (WP4).
promotion of peace (United Nations, 2015).
In recent years, Colombia has established several DOCUMENT KNOWLEDGE ON USEFUL PLANTS AND FUNGI
policies promoting the sustainable use of biodiversity OF COLOMBIA (WP1)
to reduce the social gap and to consolidate peace
whilst following a green growth approach to economic Native non-crop plants and fungi have great potential to
development (Baptiste et al., 2017). The country has improve livelihoods and economic development in the
stated its intention to develop as a bioeconomy, which country. However, high-quality information on their uses and
is “an economy that efficiently and sustainably manages properties remains scarce or inaccessible. This WP aimed
biodiversity and biomass to generate new value-added to gather and compile this information to boost Colombia’s
products, processes and services based on knowledge bioeconomy and to benefit local communities.
and innovation” (CONPES, 2018). The commitment of Significant effort to document Colombian floral and fungal
the Colombian government to green growth has been diversity has been carried out in recent years. Still, hundreds
demonstrated through the Colombia Bio programme, of sources of published information had not been compiled
and its principal aim to foster the development of the and made readily available. The project brought together a
bioeconomy through science, technology, and innovation. binational team of researchers who collected and compiled
As an offspring of this overarching national effort, the data from available datasets and publications; digitised
Useful Plants and Fungi of Colombia project (UPFC) was data that were not readily available; scanned specimens,
supported by a Professional Development & Engagement illustrations, and records from scientific collections (from
grant under the Newton-Caldas Fund partnership. The herbaria, fungaria, and libraries); generated an image bank of
£2.5 million grant was funded by the UK Department for the useful plants and fungi of Colombia; developed species
Business, Energy, and Industrial Strategy (BEIS) and the distribution maps and models; and created species profiles
Colombian Ministry of Science, Technology and Education that are available online through the project’s portals.
(MinCiencias) and was delivered by the British Council over The gathered information was analysed to identify
28 months from November 2019 to February 2021. taxonomic and geographic gaps and assessed for quality
and potential contribution to conservation practices, among
THE USEFUL PLANTS AND FUNGI OF COLOMBIA other aspects. In all, six research questions on useful
The overall goal of the UPFC was to develop pathways plants and fungi were addressed, each with a scientific
to enhance nature’s contribution to people in Colombia output (https://in-colombia.org/):
by increasing, consolidating, and making accessible the
knowledge on its useful plants and fungi for the benefit • How can we assess the comprehensiveness (in quantity
of local communities; and to promote a market for useful and quality) of the information available for each
indigenous species and their high-value products to motivate species?
the sustainable use of biodiversity whilst protecting the • Are there taxonomic gaps in the available information?
surrounding natural resources. • Are there geographic gaps?
This project documented and broadly disseminated
knowledge on useful plants and fungi of Colombia across • What are the most likely factors influencing the observed
a wide range of audiences. Outputs targeted policy and taxonomic and geographic patterns?
decision-makers in government, local NGOs, scientists, • Which geographic areas are important for conservation?
educators, and private and public companies, aiming to
• Can traditional knowledge erosion be mitigated through
boost bioeconomic development. The project’s long-term
Citizen Science?
vision was to support Colombia’s economic future by
promoting the sustainable expansion of its bioeconomy,
Results from initial activities helped us to identify
based on its unique plant and fungal diversity, whilst
poorly surveyed areas of the country for useful plants and
conserving its natural resources and associated
plants in general (Bystriakova et al., 2021), which informed
traditional knowledge.
subsequent fieldwork and ethnobotanical priorities for WP1
The UPFC project was structured as four work packages
(Figure 1).
(WP), each with a specific objective:
In addition to the research questions, three tools were
designed to manage and collect data on useful plants
1. Document knowledge on useful plants and fungi of
and fungi, specifically: 1) the shinyCCleaner tool, to clean
Colombia (WP1).
georeferenced records in the R environment (Ondo et al.,
2. Synthesise and disseminate knowledge on useful unpub.); 2) an online survey form built in ESRI Survey123,
plants and fungi of Colombia using various platforms for to collect ethnobotanical information from mobile devices;
different audiences (WP2). and 3) a Plant Use Data Management tool, with modules for
3. Develop a framework for creating a sustainable value data entry and exportation, taxonomic reconciliation, and
chain network (VCN) from plant and fungal diversity (WP3). classification of uses, among other functions.
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
well-surveyed grid cells, defined as those with ≥ 25 observations, calculated by bioregion.
SYNTHESISING AND DISSEMINATING KNOWLEDGE ON The project built two websites (https://www.kew.org/
USEFUL PLANTS AND FUNGI OF COLOMBIA (WP2) upfc, in English; and https://in-colombia.org/, in Spanish)
Access to information about plants and fungi and their uses to inform the public and disseminate project outputs. In
underpins our collective effort to further understand their addition, it built on the previously created ColPlantA portal
diversity in Colombia, allowing us to better conserve species, (http://colplanta.org/), further synthesising knowledge
habitats, and ecosystems and to use them sustainably to the of Colombian plants and improving the user experience,
benefit of society. Knowledge must be available for different content and infrastructure (Diazgranados et al., 2020). It
audiences to empower Colombia to protect and sustainably also created a companion portal for Colombian fungi, i.e.,
utilise its valuable plants and fungi. This WP aimed to build ColFungi (http://colfungi.org/) (Figure 2). Each portal was
a comprehensive synthesis and to disseminate this through accompanied by an explanatory booklet (Diazgranados et al.,
various media, including online portals, scientific journal 2020; Gaya et al., 2021) designed for a general audience.
publications, books, booklets, reports, magazine and news Comprehensive annotated checklists for both plants
articles, social media streams and educational tools. and fungi, along with a range of thematic chapters, are
This wide range of dissemination outputs and formats presented here and in the Catalogue of Useful Plants of
was aimed strategically to reach a broad audience of Colombia as a reference for scientists and for the long-term
policy-makers, researchers, and practitioners, including preservation of knowledge.
those in governmental, monitoring and developmental aid A survey analysis was conducted to understand how
agencies, regulatory bodies, conservation organisations, much Colombian society knows about their native diversity
biobusinesses, schools and universities. Potential of plants and fungi and their uses as part of this WP. An
users included: innovators, developers, bioeconomists, online national survey was developed to collect information
herbal and food entrepreneurs (including plant and fungi on how Colombians generally use plants and fungi. Semi-
importers, suppliers and retailers), biotechnologists, structured interviews of 20 experts from different sectors
bioenergy experts, agronomists, forestry engineers, crop of society, including academia, civil society, NGOs, and
specialists, horticulturalists, arborists, weed scientists, the private sector, were also undertaken. These experts
forest managers, wildlife managers, health professionals, provided information regarding the state of knowledge of
biopharmaceutical scientists, medicine regulators, useful plants and fungi, advising on the limitations and
conservationists, naturalists, photographers, environmental opportunities of various studies and the inclusion of plants
consultants, library managers, students, professors, and fungi in value chains.
researchers (biologists, botanists, ecologists, chemists, In consideration of socio-cultural components, a
agronomists, biomedical researchers, ethnobiologists, Spanish-language storytelling series, “Somos Historias”,
among others), scientific editors, people interested in was developed, which was designed to identify and promote
culture and gardening, and the general public. practices and relationships that underlie the value chains of
key plants. The co-creation of short videos that portrayed the their habitats and ecosystems, use them sustainably, and
practices, knowledge, and relationships of local producers enjoy the benefits they provide to society. In 2020, the UPFC
with these plants and their environment promoted value project delivered ColFungi, an authoritative, expert-driven,
chains and increased project visibility to a broader audience open-access online portal for information on Colombian
while also gathering qualitative information on the local use fungi and their uses. ColFungi is an entirely new tool backed
of the species. by the expertise of Colombian mycologists and presenting
Last, the project published over 50 blogs and notes in the integrated content on the fungi of Colombia. For the first time
press to raise awareness on the importance of useful plants ColFungi is a content aggregator site—it collects
and fungi for the well-being and prosperity of Colombians information from Kew’s databases and various external
and humanity. sources, displaying it in one place while preserving the
appropriate links and citations. By bringing together data
COLFUNGI, A NEW PORTAL DEDICATED TO THE FUNGI OF from a wide range of sources, ColFungi aims to synthesise
COLOMBIA and unlock valuable information, including potential uses
Access to information about fungi and their uses, it is of fungi, while maximising the visibility of local resources.
essential to understand their diversity, conserve them and Only publicly available content is used and, as part of the
content gathering process, steps are taken to ensure that research and the distribution of species, collections,
publishers have respected the Nagoya Protocol (part of the taxonomy, ecology, geography, and economic uses.
international Convention on Biological Diversity that aims to ColFungi contributes directly to the Colombian
ensure the fair and equitable sharing of benefits arising from government’s goals of developing national and regional
the utilisation of genetic resources) as well as Colombian catalogues of biodiversity while supporting the transformation
laws relating to traditional knowledge. of the economy to drive green growth. It also maximises the
ColFungi is also part of Kew’s Useful Plants and Fungi impact of research and highlights the value of fungal diversity
project (UPF), and takes advantage of UPF’s databases and in the country, promoting knowledge exchange and transfer
standards by applying them to Colombian fungi data. This between the UK and Colombian researchers and end-users.
infrastructure provides a platform for reliable information on
Colombia’s useful fungi to be made more widely available. DIGITAL OUTREACH OPPORTUNITIES
The UPFC bioinformatics work team developed The project reached a range of audiences and created new
ColFungi based on the initial infrastructure of ColPlantA alliances and opportunities for collaboration through an
and incorporated recommendations from a detailed UX array of varying virtual events and social media activities.
analysis. The current version contains 9,400 scientific Despite the recent challenges presented by the coronavirus
names, 7,241 species profiles including 308 with images, pandemic, the project found new methods to engage with its
and 251 full species and uses descriptions. Alternative target audiences through virtual events, such as Hay Festival
names (including common names) and local checklists Cartagena Virtual, Hay Festival Digital Queretaro 2020,
are also available. The taxonomic backbone of ColFungi is Sabor Barranquilla 2021, Hay Festival Cartagena 2022, the
taken from the Index Fungorum and Mycobank databases, Colombian Chef Masterclass series, and virtual workshops,
and common names from published material, especially reaching over one million viewers. In addition, it delivered
field guides. more than 23 virtual conferences, five seminar presentations,
The UPFC project team, in collaboration with more than three teaching modules, 30 socialisation workshops, and
a dozen Colombian and European mycologists, produced a 15 radio interviews. It had more than 90,000 engagements
companion booklet to the ColFungi portal (Gaya et al., 2021). in its social media accounts (Facebook and Twitter). It also
The booklet contains a significant overview of the current published ten blogs and more than 90 popular articles and
state of knowledge of Colombian fungi and mycological media notes, which helped broaden the project’s recognition.
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.
BUILDING A VALUE CHAIN NETWORK TO SUPPORT agencies related to the production, distribution, and
COLOMBIA’S BIOECONOMY (WP3) exportation of bioproducts from Colombia. Access to
Innovative projects to deliver natural ingredients or other the live quiz platform in Kahoot can be found through
bioproducts to the market often fail because a viable the link https://kahoot.it/challenge/00115304 or in the
market has not been established. The project developed practical guide.
a value chain network (VCN) centred on a digital platform
to provide pathways to develop such markets, facilitating To establish a baseline about the current status of the
interaction between local, national and international actors Colombian bioeconomy, researchers produced a 124-page
in the production, refinement, distribution and exportation report (Rojas et al., 2020), also freely accessible at https://
of plant- and fungi-based products. The VCN aims to enable in-colombia.org/. Highlights included:
the symbiosis between plant and fungal diversity, knowledge
of its uses, and local livelihoods that depend on it. It is the 1. A prioritisation exercise for potential species in Colombia’s
first VCN focused on useful plants and fungi for Colombia bioeconomy.
and South America 2. A broad directory of companies engaged in the use of
The VCN connects scientists, technologists, innovators, prioritised plant species of native plants and fungi (25
producers, and consumers, enabling the flow of materials in total).
downstream (e.g., the supply of natural ingredients produced
3. A Science, Technology, and Innovation capability analysis
by local communities to consumers) and information
around the Bioeconomy and Natural Ingredients sector.
upstream (e.g., consumer demand to producers) whilst linking
products to species names based on a taxonomic backbone. 4. A multi-criteria value chain infrastructure for biodiversity-
This functionality allows two-way linking between the VCN based products.
platform and the scientific portals (ColPlantA and ColFungi), 5. A case study of the value chain analysis for the first
enabling users to browse specific uses, read exciting plants prioritised species, naidí (Euterpe oleracea), in the Pacific
and fungi species profiles, and contact product providers coast region of Colombia.
directly. A website for the VCN (https://www.redin-colombia.
org/) was created in Spanish to maximise outreach among 6. A search for and analysis of patents and publications
the Colombian audience (primary users) (Figure 3). related to naidí in Colombia and the rest of the world.
In addition, the project delivered the first comprehensive Finally, to inform the VCN, a standard for controlled
practical guide (174 pages) to understanding the Colombian vocabulary on natural ingredients and bioproducts was
legal processes for developing and commercialising natural- developed, which included a harmonised table and
ingredient products from non-crop plants and fungi (Rojas et accompanying explanatory report. The standard will allow the
al., 2021, available at https://redin-colombia.org/guide/). This classification, indexing, cataloguing, description, and analysis
freely available guide is in Spanish and is organised around of these ingredients and products. It was based on the
four central roles within the value chain: producer, refiner, harmonisation of three classification structures linking natural
national wholesaler/retailer, and exporter. Content focuses ingredient products: Kew’s Economic Botany Data Collection
on permissions, regulatory compliance, and certification Standard (EBDCS), the UN Central Product Classification
concerning three types of natural ingredients: harvested scheme (CPC) and the Departamento Administrativo Nacional
non-timber forest products, nursery-grown products, and de Estadistica’s Clasificacion Central de Productos (DANE
genetic material products. It articulates processes related to CPC). These three structures were reconciled based on
intellectual property rights, benefit-sharing agreements, and the concept, definition, and rules to identify a common,
threatened species, among others related to national and core standard linking botanical (e.g., species), national and
international regulations, compliance, and market access. international bioproduct classificatory structures.
Eleven semi-structured interviews were conducted with staff
from the Ministry of Environment and Sustainable Development, IMPLEMENTING A VALUE CHAIN NETWORK IN THREE PILOT
ProColombia, natural product companies, and NGOs. AREAS OF COLOMBIA (WP4)
Two online tools were created to accompany the guide: Three biodiverse pilot areas of Colombia recently affected by
the armed conflict were selected to test the potential of the
1. A serving BOT, which is an automated tool designed to VCN framework to improve the livelyhoods of impoverished
guide users to obtain information from within the guide communities (food security, health, and prosperity): Bahía
by answering a series of targeted questions regarding Solano (Chocó), Becerril (Cesar), and Otanche (Boyacá)
their needs and the type of commercial role they play (Figure 4).
within the value chain (e.g., producer, refiner, distributor, The selection of the areas considered several criteria
retailer, exporter). The serving BOT is accessible via the within five categories: environmental, social, governance,
VCN website (https://redin-colombia.org/). business, and economic factors (Diazgranados et al., 2021).
2. An online quiz in Spanish, drawing upon the information The implementation included:
contained within the guide, it tests users’ knowledge
regarding documentation, certification and responsible 1. Assessment of the useful local plants and fungi resources.
FIGURE 4. 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).
2. Evaluation of the market status based on local native among others) and analysis of statistics from the potential
diversity. implementation. The project published a booklet to connect
3. Assessment and modelling of current and potential experiences in local communities within the three pilot areas
outcomes of the VCN implementation. with a broader, more general audience. It included highlights,
notes on the prioritisation of species, stories from local
4. Identification of the most promising species and types of initiatives and experiences (Quiñones-Hoyos et al., 2021).
natural ingredients.
5. The gleaning of lessons from the implementation and THE UPFC TEAM
community feedback. The UPFC project was led by RBG Kew in collaboration
with the Instituto de Investigación de Recursos Biológicos
The project expected that implementing the VCN would Alexander von Humboldt in Colombia. It directly involved
support an improvement in family livelihoods over the 85 people (50 from Kew and 35 from Humboldt Institute)
middle- to long-term through its role in facilitating diversified, and received contributions from 34 other researchers from
market-based approaches to income generation. 26 institutions across Colombia and Europe, including the
The social impact assessment carried out during the Asociación Colombiana de Micología, Ascolmic. The Diverse
implementation of the VCN will allow the measurement and gender-balanced participation was achieved in all
of middle- and long-term changes in cultural, economic, activities to ensure a more inclusive environment and in line
and social outcomes. The outcome monitoring activities with SDG5 (Empower women). We have been collaborating
implemented at the time of writing include participant surveys to include minorities such as indigenous people, Afro-
(gender, age, locality, ethnic group, family composition, Colombians and the LGBT+ community, always following
RBG Kew’s and Humboldt Institute’s policies on diversity the University of Cali (GIM/CICBA), the Research Group in
and inclusion. From the total of 115 project participants, Environmental Sustainability (SUSA) of the Universidad de
64 (56%) were female, and 51 (44%) were male. The project los Llanos, the Research Group of Biology and Conservation
deemed this gender balance appropriate given the traditional of the Universidad Pedagógica y Tecnológica de Colombia in
tendency for science to be a male-dominated discipline. Tunja, the Research Group of Biology and Microorganisms
(BPM) of the Universidad del Valle, Universidad Santiago de
REACHING COMMUNITIES ACROSS SOCIETY Cali, Colombian Mycology group and the Colombian Group
The UPFC project built and maintained connections with of Lichenology (GCOL).
other government and non-government initiatives. Through Fieldwork was carried out through a participatory
various fora and media, objectives and outcomes were approach involving local organizations, social leaders, and
communicated to the Presidency of the Republic, the communities from the veredas of Caño Rodrigo and Río
National Committee of Sustainability, the Green Business Maracas in Becerril (Cesar), and Altazor, Betania, Camilo,
Bureau of the Ministry of Environment, the Programa de Cunchalita, Curubita, El Carmen and Nazareth in Otanche
Transformación Productiva (PTP) of the Ministry of Commerce, (Boyacá) (Figure 5).
ProColombia, the Instituto de Investigaciones Ambientales In conclusion, the UPFC project played a pivotal role
del Pacífico John von Neumann (IIAP), and the Green Growth by reaching communities across society, promoting the
Mission (GGM) created by the Colombian National Planning conservation and sustainable use of the country’s plants
Department in 2017. and fungi, combining traditional and scientific knowledge
The GGM, together with Biointropic Corporation and to boost the national bioeconomy, and raising awareness
Suricata Consulting, identified seven factors to boost about the importance of these resources for the country. On
the country’s bioeconomy: 1) research and technological a broader scale, it represents a sustainable development
development; 2) regulatory framework; 3) market dynamics; 4) model of green growth in megadiverse nations worldwide.
human capital; 5) financing and investment; 6) environmental
or ecosystem services; and 7) infrastructure. By contributing UNEXPECTED IMPACTS AND CHALLENGES DUE TO COVID-19
directly to factors 1–4, the project supported GGW’s mission The pandemic of 2020–21 had significant impacts on
and the Green Growth Policy, i.e., “Promote by 2030 the where and how the UPFC project team carried out their
increase in productivity and economic competitiveness of work due to the restriction of travel and facility access,
the country, while ensuring the sustainable use of natural recruitment delays, and the shortage of human resources.
capital and social inclusion, in a compatible way with the Mitigation plans proved effective in buffering some of the
climate”. effects, especially regarding WP1, WP2 and WP3 workflows
It engaged the Colombian National Board of Natural and timelines, through increased teleconferencing and
Ingredients (Programa de Interés estratégico-PINE), an online stakeholder consultations with several cross-cutting
initiative led by the Program Colombia+ Competitiva of contingency actions. However, the capacity building (e.g., the
SwissContact as part of a 5-year plan to boost natural internship and research visitor programmes) and fieldwork
ingredients for cosmetics. The Humboldt Institute, leader of activities had to be considerably scaled down, and future
the board’s six sub-activities regarding the governance and funds could help to strengthen the implementation of the
competitiveness of value chains from native fungal and plant VCN at the national scale.
species, was vital in maintaining this interaction.
The project also connected with private-sector PATHWAY TO IMPACT ON ECONOMIC DEVELOPMENT AND
organisations such as Crepes & Waffles, the Chamber SOCIAL WELFARE
of Food Industry, the National Association of Industrials This project has developed novel pathways to enhance
of Colombia (ANDI), Corpocampo, Naidí Pacífico SAS, nature’s contribution to people in Colombia. It has made
Fondo Acción, Planeta CHB SA, Partnerships for Forest, available ‘state-of-the-art’ knowledge on the nation’s useful
MUCHOCOL, UNIANDES (Faculty of Economics), Apsacesar, plants and fungi while unveiling how this information can be
Envol-vert, Selvacéutica S.A.S., E3 Asesorías, Corporación used to promote and develop markets for these species and
Boyapaz, Alianza Quinchas, among many others. their high-value products.
The mycological work was made possible through Beyond the delivery of specific outputs, a broader set of
extensive collaboration with the mycological community project outcomes were achieved, such as:
in Colombia and Europe, including the Colombian
Association of Mycology (ASCOLMIC), Botanischer Garten • A significant increase in online, open-access knowledge
und Botanisches Museum in Germany, BioMicro Group at on the useful plants and fungi of Colombia.
School of Microbiology of the Universidad de Antioquia,
the Group of Taxonomy and Ecology of Fungi of the • Increased capacity of Colombian scientists to research
Universidad de Antioquia (TEHO), Universidad de los plant and fungal diversity and uses.
Andes, Universidad del Cauca, Universidad del Rosario, • Increased public perception and awareness about the
Universidad Distrital Francisco José de Caldas, Universidad importance of plant and fungal diversity and their uses to
de San Buenaventura, the Research Group in Mycology of support green growth in Colombia.
FIGURE 5. Map of urban centres of main collaborators, 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).
• Improved opportunities for alternative local income of Colombia's diversity of useful plants and fungi and
streams based on value-added to bioproducts through will inform policy and decision-makers in government
VCN development, at least in the three pilot areas. and non-government organisations to boost sustainable
• Supported expansion of a natural ingredients market at biobusinesses and support national and international-
the national level. scale connections. The VCN may improve the livelihoods
of communities at the national scale, providing
• Opportunities opened for further improvement to new opportunities to reduce poverty, reduce gender
people’s livelihoods in the three pilot areas. inequality, and improve health and well-being.
• Informed development of a bioeconomy, bringing • Long-term (10–15 years): Colombia will rise as a
together science, policy, and industry. successful exemplar of a developing country that is
• The predicted spillover effects included: expanding its bioeconomy based on its unique plant and
fungal diversity whilst preserving its natural environment.
• Increased interest by Colombian institutions in
expanding plant and especially fungal research.
FUTURE PERSPECTIVES AND REMARKS
• Enhanced recognition of market possibilities for
natural ingredients based on native species of plants The project has contributed significantly to the Colombian
and fungi at the Colombian national level. Green Growth Policy’s scope to “Promote by 2030 an
increase in productivity and economic competitiveness of
• The incentive to develop specific areas of commercial
the country, while ensuring the sustainable use of natural
businesses based on natural ingredients from native
capital and social inclusion, in a compatible way with
Colombian plants and fungi, e.g., by the British and
the climate”. While the transformation of a country must
Colombian Chamber of Commerce, ProColombia,
be seen as a very complex, long-term process, usually
among others.
involving all levels of society, the UPFC project is confident
• Expansion of Colombian bioeconomy by strengthening that its outputs and outcomes have supported Colombia’s
industrial endeavours to add value to useful plant transformation towards a greener, more sustainable
and fungal products. economy by enhancing critical baseline knowledge on
plants and fungi, and by re-imagining how access to and
In addition to the contributions to the United Nations practical use of this information could be improved through
SDGs, the project outputs and activities also contributed open-access digital platforms and tools. The impact of the
to several objectives of the CBDs Global Strategy for Plant UPFC project will continue, with the information generated
Conservation (GSPC), particularly those linked to ecosystem here being made broadly available to Colombian companies
goods and services (GSPC1), recognition, respect and in the agricultural, pharmaceutical, food, human health,
maintenance of indigenous and local community knowledge and personal care sectors that have an interest in the
(GSPC 5), and improved awareness of the urgency of plant sustainable development of biodiversity-based (bio-based)
conservation and public participation (GSPC 6). products. Colombia’s green transformation has begun!
As most communities in rural Colombia are male-
dominated, with men focused on farming, mining and
construction, the project emphasised pathways for women Acknowledgements
to play leading roles in obtaining and commercialising The UPFC project thanks all the researchers and contributors who
natural ingredients, potentially bringing additional income made this project possible: Adriana Corrales, Aida Marcela Vasco-
to their household. Thus, the implementation of the VCN Palacios, Alba Nohemi Téllez A., Alejandra Aguilar, Alejandra Osejo,
could improve gender equality and opportunities, supporting Alex Monro, Aline Dufat, Amalia Díaz, Ana Cristina Bolaños-Rojas,
SDG5 (Empower women). Ana Esperanza Franco-Molano, Ana Garrido, Andrea Baquero,
Angélica Herrera, Angie Karina Rengifo Fernandez, Anna Haigh,
The project pathway to impact on economic development
Benedetta Gori, Bibiana Moncada, Bob Allkin, Camilo Garzón,
and social welfare was based on a stepwise approach:
Carlos Cortés, Carolina Quiñones, Carolina Soto, Carolina Tovar,
Catalina Mesa Sánchez, Clara Morales, Cristian Lasso-Benavides,
• Short-term (1–3 years): the project documented and Dairon López, Daniel Jiménez, Daniel López, David Bishop, David
broadly disseminated knowledge on the useful plants and Díaz-Escandón, Diana Marcella Moreno, Diego Simijaca-Salcedo,
fungi of Colombia, reaching both general and academic Edier Soto-Medina, Eduardo Toledo Romero, Edwin Rios, Ehidy
audiences, as well as policy and decision-makers. It also Rocio Peña-Cañón, Elizabeth Hodson, Ellie Graves, Emmerson
contributed to capacity building. Last, by implementing Pastás, Ester Gaya, Fabiola Eugenia González-Cueller, Felipe
the VCN in three pilot areas, the project aimed to García, Germán Torres, Giuliana Furci, Harrie J.M. Sipman, Henry
support improvements to communities’ livelihoods by Agudelo, Henry Yesid Bernal M., Hugh Pritchard, Ian Ondo, James
establishing a baseline that can be monitored and by Morley, Jenny Williams, Jennyfer Andrea Aldana Mejía, Joaquim
d’Souza, Joseph Ruff, Julia Carretero, Justin Moat, Laura
providing some initial training to communities.
Green, Laura Kor, Lee Davies, Lina Guevara, Louise Colville,
• Mid-term (3–10 years): the project outputs will increase Luis Fernando Coca, Lynn Parker, Malcolm Stone, Marcela
awareness in Colombian society about the importance Cely, Margarita María Jaramilo-Ciro, Mariana Pizano Noguera,
Mario Murcia, Mark Nesbitt, Martha Lucía Ortiz-Moreno, Mateo Gaya E, Vasco-Palacios AM, Vargas-Estupiñán N, Lücking R, Carretero
Fernández, Mauricio Ramírez-Castillón, Mauricio Salazar-Yepes, J, Sanjuan T, Moncada BBA, Bolaños-Rojas AC, Castellanos-Castro
Meike Piepenbring, Melanie-Jayne Howes, Mimi Tanimoto, Nadia C, Coca LF, Corrales A, Cossu T, Davis L, Dsouza J, Dufat A, Franco-
Molano AE, García F, Gómez-Montoya N, González-Cuellar FE,
Bystriakova, Natalia Vargas-Estupiñán, Nataly Gómez-Montoya,
Hammond D, Herrera A, Jaramillo-Ciro MM, Lasso-Benavides C. Mira
Nestor García, Nestor R. Salinas, Nicolas Black, Pablo Hendigo, MP, Morley J, Motato-Vásquez V, Niño-Fernández Y, Ortiz-Moreno ML,
Paola Acosta, Paola Andrea Torres-Andrade, Paul Kersey, Paul Peña-Cañón ER, Ramírez-Castrillón M, Rojas T, Ruff J, Simijaca D,
Wilkin, Priscilla Reis, Rafael Felipe de Almeida, Raquel Negrão Sipman HJM, Soto-Medina E, Torres G, Torres-Andrade PA, Ulian T,
Baldoni, Rene Camacho, Robert Lücking, Rose Taylor, Sabina White K, Diazgranados M (2021) ColFungi: Colombian resources for
Bernal, Santi Gómez, Sarah Phillips, Silvia Restrepo, Tatiana Fungi Made Accessible. Richmond: Royal Botanic Gardens, Kew. 36p.
Sanjuan, Tim Utteridge, Tiziana Cossu, Vicky Murphy, Viviana IPBES (2018). The IPBES regional assessment report on biodiversity
and ecosystem services for the Americas. Secretariat of the
Motato-Vásquez, Wiebke Hillebrecht, William Baker, William
Intergovernmental Science-Policy Platform on Biodiversity and
Milliken, Yeina Milena Niño-Fernández, and Yih Wen Fung. The Ecosystem Services, Bonn, Germany. 656 p. doi: 10.5281/
project also thanks the local communities and authorities in the zenodo.3236252.
pilot areas, as well as all the stakeholders involved. Khoury CK, Bjorkman AD, Dempewolf H, Ramirez-Villegas J, Guarino L,
Jarvis A, Rieseberg LH, Struik PC (2014) Increasing homogeneity
in global food supplies and the implications for food security.
References Proceedings of the National Academy of Sciences U S A , 111, 4001-
4006. doi: 10.1073/pnas.1313490111
Andrade-C MG (2011) Estado del conocimiento de la biodiversidad Millennium Ecosystem Assessment (2005) Ecosystems and human well-
en Colombia y sus amenazas: Consideraciones para fortalecer la being: synthesis. Washington, DC: Island Press. 155p.
interacción Ciencia-Política. Revista de la Academia Colombiana de Negrão R, Monro AK, Castellanos-Castro C, Diazgranados M (2022)
Ciencias Exactas, Físicas y Naturales, 35, 491-507. Catalogue of Useful Plants of Colombia. Richmond: Royal Botanic
Baptiste B, Pinedo-Vasquez M, Gutierrez-Velez VH, Andrade GI, Vieira Gardens, Kew. 1,056p.
P, Estupiñán-Suárez LM, Londoño MC, Laurance W, Lee TM (2017) Ondo I, Zizka A, Pironon S, Antonelli A (unpublished) ShinyCCleaner: an
Greening peace in Colombia. Nature Ecology and Evolution, 1, 0102. interactive app for cleaning species occurrence records.
doi: 10.1038/s41559-017-0102 Quiñones-Hoyos C, Rengifo-Fernández A, Bernal-Galeano S, Peña R,
Bernal HY, Quevedo Sánchez GF, García Martínez H (2011) Pautas para el Fernández M, Rojas T, Diazgranados M (2021) Una mirada a las
conocimiento, conservación y uso sostenible de las plantas medicinales plantas y los hongos útiles en tres áreas biodiversas de Colombia.
nativas en Colombia: Estrategia nacional para la conservación de Richmond: Royal Botanic Gardens, Kew; Bogotá DC: Instituto de
plantas. Bogotá DC: Instituto de Investigación de Recursos Biológicos Investigaciones en Recursos Biológicos Alexander von Humboldt.
Alexander von Humboldt. 236p. Rojas MT, Cortés C, Noguera M, Acosta P, Diazgranados M (2021) Guía
Bernal R, Gradstein SR, Celis M (eds.). (2019) Catálogo de plantas práctica para potencializar el uso sostenible de los ingredientes
y líquenes de Colombia. Instituto de Ciencias Naturales, naturales en Colombia. Richmond: Royal Botanic Gardens ; Bogotá
Universidad Nacional de Colombia, Bogotá. Available at: http:// DC: Instituto de Investigaciones en Recursos Biológicos Alexander
catalogoasdecolombia.unal.edu.co von Humboldt. 172p.
Bystriakova N, Tovar C, Monro A, Moat J, Hendrigo P, Carretero J, Torres- Rojas T, Cortés C, Noguera M, Ulian T, Diazgranados M (2020) Evaluación
Morales G, Diazgranados M (2021) Colombia’s bioregions as a del estado de los desarrollos bioeconomicos colombianos en plantas
source of useful plants. PLoS One, 16(8), e0256457. doi: 10.1371/ y hongos. Kew: Royal Botanic Garden ; Bogotá DC: & Instituto de
journal.pone.0256457 Investigación de Recursos Biológicos Alexander von Humboldt. 125p.
Cámara-Leret R, Paniagua-Zambrana N, Balslev H, Macía MJ (2014) Sabater S, González-Trujillo JD, Elosegi A, Rondón JCD (2017) Colombian
Ethnobotanical knowledge is vastly under-documented in ecosystems at the crossroad after the new peace deal. Biodiversity
northwestern South America. PLoS One, 9(1), e85794. doi: 10.1371/ and Conservation, 26, 3505-3507. doi: 10.1007/s10531-017-1415-8
journal.pone.0085794 Salazar A, Salazar JF, Sánchez-Pacheco SJ, Sanchez A, Lasso E, Villegas
CONPES (2018) Política de Crecimiento Verde. Consejo Nacional de JC, Arias PA, Poveda G, Rendón ÁM, Uribe MR, Pérez JC, Dukes JS
Política Económica y Social (Documento CONPES 3934). Available at: (2021) Undermining Colombia’s peace and environment. Science,
https://colaboracion.dnp.gov.co/CDT/Conpes/Económicos/3934. 373, 289-290. doi: 10.1126/science.abj8367
pdf United Nations (2015) Transforming our world: the 2030 Agenda for
Diazgranados M, Allkin B, Ávila F, Baker W, Bishop D, Black N, Bystriakova Sustainable Development. Document A/70/L.1. New York: UNGA.
N, Carretero J, Castellanos-Castro C, Cely M, Colville L, Cossu T, 35p.
Davies L, Díaz A, Dsouza J, García F, Gaya E, Graves E, Green L, WWF-Colombia (2017) Colombia Viva: un país megadiverso de cara al
Haigh A, Hammond D, Hendrigo P, Herrera A, Hillebrecht W, Howes futuro, informe 2017. Cali: WWF-Colombia. 166p.
MJ, Iggulden D, Kersey P, Kor L, Mattana E, Milliken W, Mira M, Moat
J, Monro A, Morley J, Murphy V, Nesbitt M, Ondo I, Parker L, Phillips
S, Piquer M, Pritchard HW, Reis P, Rojas T, Ruff J, Stone M, Tanimoto
M, Torres G, Tovar C, Turner R, Taylor R, Utteridge T, Vargas N, Weech
MH, White K, Wilkin P, Williams J, Ulian T (2020) ColPlantA: Colombian
resources for Plants made Accessible. 2nd edition. Richmond: Royal
Botanic Gardens, Kew. 33p.
Diazgranados M, Hammond D, Rojas T, Castellanos-Castro C, Díaz A,
Hernández M, Quinoñez C, Kor L, Ulian T (2021) Selection of pilot
areas in Colombia for the implementation of a value chain network on
useful plants and fungi. UPFC Report Series. Richmond: Royal Botanic
Gardens, Kew; Bogotá DC: Instituto de Investigaciones en Recursos
Biológicos Alexander von Humboldt.
Eufemia L, Bonatti M, Castro-Nunez A, Lana M, Morales H, Sieber S
(2019) Colombia’s inadequate environmental goals. Science, 364,
444-445. doi: 10.1126/science.aax3494
Food and Agriculture Organization of the United Nations - FAO (2015)
Global Forest Resources Assessment 2015. Desk reference. Available Marasmius cladophyllus
at: http://www.fao.org/3/i4808e/i4808e.pdf [Nataly Gómez-Montoya]
Anthropozoomorphic pendant
[Robert Lücking]
32 CATALOGUE OF FUNGI OF COLOMBIA
Chapter 2
Two Centuries of Mycological History in Colombia
Aída M. Vasco-Palacios1,2 & Bibiana Moncada2,3,4,5
1
Grupo de Microbiología Ambiental y Grupo BioMicro, Escuela de Microbiología, Universidad de Antioquia, Calle 70 No. 52–21, Medellín 050010, Colombia.
2
Asociación Colombiana de Micología - ASCOLMIC, Grupo BioMicro.
3
Licenciatura en Biología, Universidad Distrital Francisco José de Caldas, Cra. 4 No. 26D–54, Torre de Laboratorios, Herbario, Bogotá D.C., Colombia.
4
Botanischer Garten, Freie Universität Berlin, Königin-Luise-Straße 6–8, 14195 Berlin, Germany.
5
Grupo Colombiano de Liquenología - GCOL.
ABSTRACT
Current knowledge about the Colombian funga results from countless expeditions, researchers, and national and foreign
institutions that have contributed to the development of mycology in Colombia for more than two centuries. This historical
timeline will probably remain incomplete because several historical accounts are difficult to track down, but four different
periods are highlighted: 1. Fungal knowledge from pre-Hispanic times is still incomplete despite efforts to compile the
biocultural heritage of ancestral knowledge of indigenous, root, and traditional farming populations; 2. The 19th century’s
study of the funga started with The Royal Botanical Expedition to New Granada and was followed by other important
expeditions; 3. The 20th century was a period marked by expeditions of European naturalists, but Colombian mycologists
also started to develop areas such as phytopathology and clinical mycology; 4. From the 1970s to today, the knowledge on
Colombian funga has grown due to the contributions of foreign and national mycologists, with about 80 Colombian research
groups studying fungi nowadays. In each of these periods, some notable events and influential figures left their mark on the
historical memory of Colombian mycology.
RESUMEN
El conocimiento sobre la funga colombiana es el resultado de innumerables expediciones, investigadores e instituciones
nacionales y extranjeras que por más de dos siglos han aportado a su desarrollo. En este capítulo se presenta un breve
recuento de esta historia dividiéndola en cuatro períodos, mencionando hechos destacados y personajes influyentes
que han dejado huella en la memoria histórica de la micología colombiana. En el primer periodo se reconoce la relación
de las diferentes comunidades ancestrales colombianas con los hongos, vestigios que se pueden encontrar en piezas
prehispánicas en el Museo del Oro de Bogotá y en publicaciones iconográficas. A pesar de los esfuerzos en la recopilación
del patrimonio biocultural de conocimientos ancestrales de poblaciones indígenas, raizales y campesinas tradicionales,
aún está lejos de completarse. El segundo período denominado siglo XIX, coincide con la participación de Francisco José
de Caldas en la Real Expedición Botánica del Nuevo Reino de Granada, quien reporta el primer hongo para Colombia.
Posteriormente Sir William Jackson Hooker describe varias especies de hongos y líquenes de las colecciones realizadas
por Alexander von Humboldt y Aimé Jacques Alexandre Bonpland durante sus viajes por el país. Y finalmente, la edición del
Prodromus Florae Novo-Granatensis de José Jerónimo Triana y Jules Émile Planchon (1863–1867), con las colaboraciones
de Joseph-Henri Léveillé (Hongos) y William Nylander (Líquenes). El tercer período, del siglo XX hasta 1970, inicia con la
Expedición de Otto Fuhrmann y Eugène Mayor conocida como la expedición Helvética, inspirando a micólogos como Carlos
Chardón y Rafael Toro, quienes lideraron el primer inventario de hongos de Colombia. Posteriormente, el establecimiento
de la carrera en ciencias agrícolas en la Universidad Nacional sede Medellín en 1927, contribuyó sustancialmente a la
formación de micólogos y al conocimiento de los hongos fitopatógenos. Así mismo, la micología clínica tuvo su inicio con las
investigaciones de Alfredo Correa y Alfonso Jaramillo; hacia los años sesenta aparece la primera micóloga médica Angela
Restrepo-Moreno, creadora de una de las más importantes escuelas micológicas de América Latina, posteriormente, en
Bogotá, la reconocida micóloga Elizabeth Castañeda ha liderado el estudio de los hongos que causan enfermedades. El
cuarto período comprendido entre los 70s y la segunda década del siglo XXI, debería considerarse el periodo de iluminación
micológica, pues no solamente fue el tiempo con mayores visitas de especialistas internacionales, sino también el despertar
de la micología nacional en todas las áreas conocidas hoy. Especialistas en diversos grupos y líneas conformaron sus
propias escuelas y han formado a las nuevas generaciones de investigadores. Reciente dos hechos han transformado
la historia de la micología, ambos involucrando el trabajo en equipo como fuente de progreso. El primero de ellos, la
conformación de grupos de investigación que ha fortalecido los procesos de investigación y formación de capital humano,
además de generar un reconocimiento colectivo de las actividades de investigación a nivel nacional e internacional. Y
Finalmente, en el 2019, la consolidación de la Asociación Colombiana de Micología (ASCOLMIC) que tiene como misión,
además de conectar a todos aquellos que investigan en hongos en Colombia, generar resultados científicos que puedan
integrarse a las políticas públicas nacionales, y conlleven a la inclusión de los hongos en las agendas nacionales a fin de
orientar y gestionar la protección, manejo y uso de la funga del país.
The traditional use of fungi by Colombian indigenous (Bolivar), and part of the Río Magdalena (1820–1821)
communities continues until the present. Colombia comprises (Delprete et al., 2002; Baldini & Guglielmone, 2012). In 1882,
an enormous cultural diversity, including 85 indigenous Sir William Jackson Hooker described 73 species of fungi,
peoples, three distinct Afro-Colombian ethnicities, the 12 new to science, collected by Alexander von Humboldt
Romani people, and traditional farming communities (DANE, and Aimé Jacques Alexandre Bonpland in Venezuela and
2007). These communities primarily depend on environmental deposited at the Kew Gardens herbarium (K).
resources, hunting, fishing, and gathering wild products such Shortly after the middle of the 19th century, José Jerónimo
as fruits, buds, and mushrooms, and producing food using Triana and Jules Émile Planchon (1863–1867) published
fermentation. The use of mushrooms as food, in traditional the Prodromus Florae Novo-Granatensis, including a lengthy
medicine, and for societal and religious rituals has been chapter on lichens (Lichenes) elaborated by the Finish
recorded for the Uitoto, Andoke, Muinane, Bora-Miraña, and lichenologist and mycologist William Nylander (Nylander,
Yukuna-Matapí of the Amazon (e.g., Landaburu & Pineda, 1863). He described numerous new species based on
1984; Urbina, 1986; Vasco-Palacios et al., 2008), as well the collections of the German botanist Alexander Lindig,
as for the Ingas in Putumayo (Sanjuan, 1999), the Embera- who lived in Bogotá and undertook botanical expeditions
chamí in the Pacific tropical forests (Cayón & Aristizábal, to the surroundings and other areas, including Choachí,
1980), the U'wa in the eastern foothills of the mountain Fusagasugá, Tibacuy, and Villeta (Cundinamarca), Honda
range in the Nevado del Cocuy (Rocheraux, 1959), the (Tolima), and Muzo (Boyacá). Another chapter of non-
Kokonucos in Cauca (Gonzalez-Cuellaret al., 2021), and the lichenised fungi, by the French mycologist Joseph-Henri
Wayuu in the upper Guajira, who use fungal spores as facial Léveillé (1863), enumerating 63 species, was based on
sunscreen (Villalobos et al., 2017). Traditional farmers from collections by Hooker, Justin Goudot, and Lindig (Herbarium,
Boyacá and Santander use mainly ectomycorrhizal species Muséum National d' Histoire Naturelle).
associated with oaks (Quercus humboldtii) as a food source
(Peña-Cañón & Enao-Mejia, 2014). 20TH CENTURY TO THE 1970S
At the beginning of the 20th century, field expeditions to
19TH CENTURY Colombia were very popular among European naturalists.
Until July 20th of 1810, Modern Colombian territory was The Swiss parasitologist Otto Fuhrmann and the Swiss
known as part of the Viceroyalty of the New Kingdom of mycologist Eugène Mayor undertook further scientific
Granada. The first inventory of Colombia’s biodiversity dates exploration in Colombia within the Helvetic Expedition
to 1783–1816 when The Royal Botanical Expedition to New (Fuhrmann & Mayor, 1914). Their results contributed to the
Granada was led by the Spanish priest José Celestino Mutis knowledge of various groups of fungi, mainly pathogens
(Díaz-Piedrahita, 1991). Around 20,000 plants and 7,000 and lichens. This work constituted the first extensive
animals were collected and documented. The specimens systematic treatment of Uredinales (rusts) studied by Mayor,
collected in this expedition were published in a series of and Pucciniales, by the German mycologist Paul Sydow and
books, in which volume 2, published a posteriori (Aguirre & his son Hans Sydow, from Berlin (Gómez-Gutiérrez, 2011;
Calonge 1985), entitled Algae, Lichens, Fungi, Liverworts, Buriticá-Céspedes et al., 2014). The collected materials
and Mosses, included 15 fungi (seven Basidiomycota and represent 156 species currently deposited at the University
eight Ascomycota, three of them being lichenised). The of Neuchatel herbarium (NEU) in Switzerland (Pardo-Cardona,
original illustrations were edited with complementary texts 2001; Gómez-Gutiérrez, 2011). The German lichenologist
and scientific determinations were done by Jaime Aguirre and mycologist Gustav Lindau, also from Berlin, studied
(Colombian bryologist and lichenologist) and Francisco de the lichens collected in this expedition reporting 64 taxa,
Diego Calonge (Spanish mycologist). Unfortunately, the including the then-new species Lecidea mayorii. The holotype
original plates of this volume do not have information on of this species was unfortunately destroyed during the 1943
the place, date, collector, or artist, except plate VI, called a bombing in the Berlin herbarium (Botanischer Garten und
posteriori Cookeina sulcipes, made by the Colombian artist Botanisches Museum Berlin, B).
Francisco Javier Matis on April 14th, 1785 (Figure 2e). This The Helvetic Expedition inspired the Puerto Rican
plate was documented for the first time by Calonge (1986). mycologists and pathologists Carlos Chardón and Rafael
As an update to Volume II, we have compiled plates III, IV, Toro to carry out the Mycological Explorations of Colombia
VI, and VIII, added photographs of the species “in situ,” and supported by the Colombian government (Chardón & Toro,
updated their taxonomy (Figure 2). 1930). They registered 610 species of fungi collected
Francisco José de Caldas (1803) provided the first between 1926 and 1930 in the surroundings of Medellín, the
known report on a phytopathogenic fungus, a description Magdalena Valley, the Pacific coast, the Valle del Cauca, and
that coincides with the wheat leaf rust, Puccinia graminis the mountain ranges of the departments of Cundinamarca
(Figure 2). Later, the Italian botanist and naturalist Luigi and Tolima. This work included mostly phytopathogenic
Carlo Giuseppe Bertero, during his expedition to the Antilles fungi and some macrofungi, later recognised as the first
and the Caribbean coast of Colombia (1816–1821), made modern inventory of Colombian fungi. Frank D. Kern and
considerable collections of plants, algae, and fungi from Herbert Whetzel (Uredinales), Julian Miller (Xylariaceae),
Santa Marta (Magdalena), Barranquilla (Atlántico), Mompóx and Jay Seaver (Discomycetes), Dr. L.O. Overholts (Higher
FIGURE 2. Species drawn in Flora de la Real Expedición Botánica Tomo II and their possibly corresponding or
similar species. A–B Cladonia meridensis, previously identified as Cladonia didyma. C–D Stereocaulon ramulosum.
E–F Cookeina sulcipes. and G–H Trametes versicolor (Jar. Bot. Mad. 86).
Basidiomycetes), Dr. 1Y. FI. Weston Jr. (Phycomycetes), (Figure 3). The collection of phytopathogenic fungi gave
Dr. Charles Chupp (Cercospora) collaborated in that rise to the Tibaitatá collection, currently at Corporación
publication. Most of these collections were deposited in Colombiana de Investigación Agropecuaria - AGROSAVIA.
foreign herbaria, especially at Cornell University (CUP) and A few years later, in 1936, the Spanish botanist and
Pennsylvania State University (PACMA) herbaria, the latter pharmacist José Cuatrecasas, motivated by the celebration
now transferred to the US National Fungus Collections, of the second centenary of the birth of Mutis, reported
USDA-ARS (BPI). In addition to the contributions of Chardon, two species of fungi and 22 lichens, collected by himself
in 1927, higher education in agricultural science was and currently held in the National Herbarium, Smithsonian
established at the Faculty of Agricultural Science in Medellín Institution (US) (Cuatrecasas, 1936). Between 1937–
and at the Plant Health Service (PHF) of Colombia (Ministry 1939, George Willard Martin (1937, 1938, 1939a, 1939b)
of Industries). These developments were fundamental to published a serial publication on the “New or noteworthy
the development of phytopathology in Colombia. Enrique fungi from Panama and Colombia”. His collections from the
Pérez Arbeláez, botanist and catholic priest, head of the Sierra Nevada of Santa Marta in the Magdalena department
Botany Department at the PHF, implemented a methodology enriched our knowledge about the funga of this biodiversity
suggested by Chardón and Toro, both recognised for initiating “hotspot”. Martin described eight new species of
phytopathology in Colombia. This method includes drawings microfungi from this area. The exploration of parasitic fungi
of plant host, the symptoms, and the different stages of associated with plants of agronomic importance continued
pathogenic fungi. Part of those drawings (1940–1944) was during the 1950s with Carlos Garcés and Rafael Obregón,
rescued by Pablo Buriticá-Céspedes and published in 1996 both professors at the Facultad de Agronomía (Faculty
of Agronomy), which was integrated into the Universidad
Nacional de Colombia, as their Medellín headquarters, and
of some of their students (Pardo-Cardona, 2001).
Clinical mycology arises from the need to know the fungi
that cause diseases, including endemic diseases. In the
1930s, the study of human pathogenic fungi in Colombia
focused on skin diseases and their annexes. One of the
first important contributions is the monograph Contribution
to the study of mycosis in Antioquia (1929), by doctors
Alfredo Correa and Alfonso Jaramillo, from the Universidad
de Antioquia, which included aspects of the classification,
laboratory science, and clinical relevance of medically
important fungi. In this same period, dermatologists were
particularly interested in systemic mycoses, the study of how
fungi affect internal organs. Around the 1940s, publications
on systemic mycosis started with the publications on
Blastomycosis by Miguel Serrano (Serrano, 1943). In this
first stage, microscopic examination and isolation in culture
were used. Still, the initiation of pathophysiology changed
the orientation of the diagnosis of mycoses, with clinicians
trying to identify the fungus causing the disease and its
mechanisms of action or alteration (Galvis-Pérez et al., 2013).
In the early 1960s, research in paracoccidioidomycosis
was consolidated thanks to Angela Restrepo-Moreno. She
was the first woman in medical mycology in Colombia, a
field previously exclusively male-dominated. In 1962, she
compiled a series of cases of various mycoses, giving
the first overview of the state of the art, including cases
caused by Cryptococcus and Sporothrix. Thanks to her
dedication to teaching, Restrepo-Moreno formed one of the
most important schools in this area in Latin America. In
Bogotá DC, Elizabeth Castañeda del Gordo studied fungi
that cause human diseases from a very young age. Later,
at the National Institute of Health, she and her team made
Colombia one of the leaders in Latin America in identifying
this type of agent. Currently, Colombia is well established
FIGURE 3. Illustration of Puccinia graminis Pers. (Pucciniaceae) and recognised worldwide in this area, with many research
made by Alberto Franco 1940. Iconography of the National groups dedicated to the study of mycoses, thanks to the
Phytopathology Service, personal collection of Dr. Pablo Buriticá. efforts of these two women.
THE 70S TO THE SECOND DECADE OF THE 21ST CENTURY Liquenología- GCOL (Colombian Group of Lichenology), with
During the last third of the 20th century, knowledge on the the support of the German mycologist Robert Lücking in 2010.
Colombian funga increased largely due to contributions This prompted the study of lichenology with great contributions
by foreign mycologists visiting the country. Some of such as: the consolidation of reference collections in several
these scientists left their legacy by training Colombian Colombian herbaria such as Herbario Forestal Gilberto Emilio
mycologists. Particularly, during this period, the expeditions Mahecha Vega, Sección no Vasculares, Universidad Distrital,
of Kent P. Dumont (New York Botanical Garden – NYBG) (UDBC), Herbario Luis Sigifredo Espinal-Tascón, Universidad
in Colombia during 1968 provided the base for further del Valle (CUVC), and Herbario Universidad de Caldas (FAUC);
unveiling of the richness of Colombian fungi. The project the description of new species (Lücking et al., 2017); the
Mycological Flora of Colombia was a program established publication of several ecological (Soto et al., 2012; Chilito
in 1974 between the New York Botanical Garden and et al., 2016; Simijaca et al., 2018), phylogenetic/systematic
the Instituto de Ciencias Naturales — ICN (Institute of (Lücking et al., 2014; Moncada et al., 2014; Coca et al., 2018),
Natural Sciences) at Universidad Nacional de Colombia. bioindication (Díaz-Escandón et al., 2016; Correa-Ochoa et
From 1978 to 1983, this project amassed about 4,000 al., 2020), microbiome (Sierra et al., 2020), and conservation
collections from the departments of Antioquia, Boyacá, (IAVH, 2019) studies; besides training specialised Colombian
Cauca, Cundinamarca Boyacá, Caquetá, and Valle del personnel (see Chapters 6, 14). At the same time, Norma
Cauca. It was the first international project for which the Valencia-Islas and Leopoldo Rojas, with their research group,
main collections were deposited in a Colombian herbarium began their studies in bioprospecting of lichen substances
(Herbario Nacional Colombiano, COL). As a result of this (e.g., Rojas et al., 2015; Leal et al., 2018). Lichens went from
project, ten publications have been produced under the title being one of the least known groups of fungi in Colombia to
Los Hongos de Colombia I–X, with descriptions and records one of the most well-studied (Lücking et al., 2021).
of macrofungi in the Ascomycota and Basidiomycota, In addition to the Mycological Flora of Colombia project
as well as phytopathogenic fungi, moulds, lichens, and and ECOANDES, numerous international researchers have
myxomycetes (e.g., Dumont et al., 1978; Guzmán & Varela, visited the country. One of the relevant mycologists in the
1978; Pulido-L., 1983). 20th century, Rolf Singer, travelled around the country and
Another significant contribution to mycology, particularly described more than 200 new species from Colombia (Mueller
lichenology and macrofungi in Colombia, was the binational & Wu, 1997). Dennis (1970) presented a brief contribution to
framework ECOANDES project between the Netherlands Colombia’s funga, including macrofungi species and various
and Colombia (1980–1983). This has been the most taxonomic groups and non-true fungi (myxomycetes). Later,
comprehensive large-scale ecological project developed in Roy Halling (NYBG, USA), Greg Mueller (Field Museum, USA),
the Colombian Andes to date. Systematic expeditions were and other colleagues conducted pioneering studies of fungi
developed along transects in four mountain ranges, including associated with montane oak in Colombia and Costa Rica
the three Cordilleras (Parque Nacional Natural los Nevados – (1986 – 1988). Collections of those expeditions are deposited
PNNN, Tatamá and Sumapaz) and the Sierra Nevada de Santa in the Herbaria of the Universidad de Antioquia (HUA), NY, and F
Marta. Lichens (Sipman, 1986, 1989, 2005; Aguirre, 2008b) (see Chapter 15). Other important mycologists that contributed
and macrofungi were studied (Pulido-L. & Boekhout, 1989a, to the knowledge of Colombian funga were Dennis Desjardin
1989b; Pulido-L., 2005, 2008), contributing to the knowledge (University of Tennessee, USA), Tim Baroni (SUNY Cortland,
of these groups in altitudinal gradients, and considerably USA), Gaston Guzmán (Instituto de Ecología de Xalapa, México),
increasing the number of species known for the country. a specialist in agaricoid fungi, and Leif Ryvarden, working with
The first Colombian studies on foliicolous lichens were done polypore fungi (University of Oslo, Norway). More recently,
by the German ecologist Sieghard Winkler and his student Colombian mycologists have carried out studies on macrofungi
Rainer Nowak, both in the Sierra Nevada de Santa Marta and (Chapters 4, 5). Important contributions to the funga from the
the Chocó region (Nowak & Winkler, 1970, 1975). During this Amazon ecosystem have been made since 2001 by a project
period, the Dutch lichenologist Harrie Sipman, based at the focused on exploring the diversity and ecology of saprotrophic,
Botanical Garden and Botanical Museum (BGBM) in Berlin, ectomycorrhizal, and entomopathogenic fungi (e.g. Franco-
emerged as a leader of lichenological studies in Colombia, Molano et al., 2005; López-Quintero et al., 2012; Vasco-
supporting the Dutch ecologist Jan Wolf in his pioneering Palacios et al., 2018, 2019, 2020; Sanjuan et al., 2015). This
studies of canopy epiphyte communities (Wolf, 1993, 1994, initiative has been widely supported by Dr Teun Boekhout, a
1995). Sipman also mentored and consolidated Jaime Aguirre Dutch mycologist who was part of the ECOANDES and visited
as the first Colombian lichenologist. Sipman and Aguirre made Colombia in the 80s. The best-studied regions are the Andean
important contributions among which are the first key to the region of the central, eastern, and western cordillera, mainly in
lichens of Colombia (Sipman & Aguirre, 1982), and three places close to the large cities and both ectomycorrhizal fungi
lichen catalogues and checklists (Aguirre, 2008a; Sipman et associated with Quercus humboldtii and saprotrophic fungi
al., 2008; Sipman & Aguirre, 2016). Towards the end of the (Vasco-Palacios & Franco-Molano, 2013; Peña-Venegas &
1980s, Juan Luis Rubiano began to work in bioindication using Vasco-Palacios, 2019; Vargas & Restrepo, 2019; Chapter 10).
lichens as model organisms (Rubiano, 1987, 1988). Colombian mycologists and lichenologists are actively working
One of the most important recent events in the study of on the threat assessment of fungal species and are among
lichenised fungi was the creation of the Grupo Colombiano de the leaders in these efforts in South America (Chapter 14).
Studies in arbuscular mycorrhizae (AM) also started in environments separated by important geographical barriers
the 1970s. Colombia was one of the first countries in South such as the Andes or different latitudes. Two years later,
America to study these organisms. At the end of the 1970s, Llanos & Kjoller (1976) presented microorganisms that
studies of AM in crops began at The International Centre for attack crude oil and hydrocarbons. As part of the series
Tropical Agriculture (CIAT) with Reinhardt Howeler and James Hongos de Colombia, John Veerkamp & Walter Gams (1983)
Spain, who isolated for the first time the species Glomus described three new species of soil fungi. Several species
manihotis, and Entrophospora colombiana (Schenck et al., of Penicillium, Trichoderma, and Talaromyces have been
1984). During the 1980s, Ewald Sieverding, considered the described from isolates of Amazonian soils (Houbraken et
father of AM studies in Colombia, in a cooperation project al., 2010; López-Quintero et al., 2013; Yilmaz et al., 2016).
supported by the German Technical Cooperation Agency Since the 1980s, Maria Caridad Cepero de García has
(GTZ), collected almost 1,200 ecotypes (Sieverding, 1984, dedicated her professional life to the study of fungal biology,
1989a, 1989b; Sieverding & Howeler, 1985). This was training several microbiologists and biologists in this area.
a starting point for the “Mycorrhizal Collection”, one of Biological control using microorganisms has a long history
the most important AM collections in the world and Latin in Colombia, emphasising the biological control of insects in
America during the eighties. The collection preserves 44 crops. The development of this field began in the 1970s when
species of AM, of which 86% are from Colombia (Garcia et microorganisms were imported from the USA to control insect
al., 2000). Those AM strains had been used as reference diseases in cotton (Cotes, 2014). Now, the entomopathogenic
cultures to develop a new AM fungal taxonomic classification fungus Lecanicillium lecanii is routinely used to control whitefly
that includes molecular data, resulting in 12 new species in this crop. Trichoderma was incorporated into the production
from Colombia (Peña-Venegas & Vasco-Palacios, 2019). of cut flowers in Colombia to control Fusarium oxysporum
Important contributions in AM have also been made in cloves. Later, in the 1990s, the Centro Nacional de
by Colombian researchers such as Marina Sanchez de Investigaciones de Café – Cenicafé (National Coffee Research
Prager (Marina Sanchez, 2007, 2010), Marina Correa de Centre) – promoted the production of Beauveria bassiana for
Restrepo (Romero et al., 2004) and Eduardo Guerrero Forero the control of the Coffee Berry Borer (Hipothenemus hampei).
(Guerrero, 1996), among others. Peña-Venegas et al. (2006) This project promoted the development of biological control
made an essential contribution to the Illustrated Catalogue of with microorganisms and encouraged producers and research
Arbuscular Mycorrhizae of the Colombian Amazon. institutes to dedicate a large part of their efforts to developing
With the rise of studies on pathogenic fungi in the country biopesticides (Cadena, 2005). The Corporación Colombiana
during the 1960s and early 1970s, the Asociación Colombiana de Investigaciones Agropecuarias (Colombian Agricultural
de Fitopatología y Ciencias Afines (Colombian Association Research Corporation), led by Dra Alba Marina Cotes
of Phytopathology and Related Sciences) was organised in since 1994, defined a comprehensive research strategy
1974, bringing together experts on the subject and generating for biological control, including a multidisciplinary team of
the Colombian Phytopathology journal, still active and with researchers. They also established infrastructure, including a
wide recognition in the country. Probably one of its most pilot plant, for the semi-commercial scaling of biopesticides,
prominent members was Dr. Pablo Buritacá Céspedes, who making Colombia the leading country in South America in
published on rusts from the late 1960s until a few years ago the development, production, and commercialisation of
(e.g., Buriticá-Céspedes, 1978, 1991; Buriticá-Céspedes & biopesticides (Cotes, 2014)
Hennen, 1980; Buriticá-Céspedes & Pardo-Cardona, 1996; For more than three decades, biotechnology has been a
Buriticá-Céspedes & Salazar, 2007; Buriticá-Céspedes et al., guide for developing areas such as medicine, agriculture,
2014). Buriticá-Céspedes (2014) was the first to describe and the food industry, among others, in Colombia. The use
genera and native species of Uredinales directly collected by of fungi in this area has been led by Dr Lucia Atehortua from
him throughout the Colombian territory and covering hitherto the Universidad de Antioquia and Dr Sandra Montoya from
unexplored areas. His work includes “Buriticá’s collection”, the Universidad de Caldas. Production of edible mushrooms
which constitutes more than 3,000 specimens deposited at has also been important in Colombia, and we can mention
the Museo Micológico, Universidad Nacional de Colombia, Carmenza Jaramillo from Manizales, Caldas in this context.
Medellin (MMUNM). Rusts are currently one of the best- This topic is developed in more detail in Chapters 12 and 13.
studied groups in Colombia, thanks also to Luis Molina-Valero, Fungi have been part of the biocultural heritage of pre-
Víctor Manuel Pardo-Cardona, and Mauricio Salazar-Yepes Hispanic and current groups of the country. At the beginning
(Chapter 9). The smut fungi (Ustilaginales) were studied by of this chapter, we referenced most of the current studies
Meike Piepenbring, who published an annotated checklist in published on this topic. Part of this traditional knowledge
2002, after visiting Colombia in 1998 (Chapter 9). has been lost due to the acculturation of communities and
Inquiries into the historical development of microfungi the loss of ecosystems. Therefore, collecting ancestral
studies are very difficult. This group of fungi includes soil knowledge has become important in different regions of
fungi, endophytes, crithidia, and moulds, which belong to the country. For example, at the Universidad Pedagógica y
various phylogenetic groups within the Fungi kingdom (e.g., Tecnológica de Tunja (UPTC), Universidad del Cauca, and
Ascomycota, Basidiomycota, Zygomycota, Mucoromycota, Universidad de Antioquia, there are research groups and
Chrytidiomycota) (Chapter 8). In 1971, Alvin Lee Rogers hotbeds in ethnomycology, and several training courses have
studied keratinophilic species comparing the mycobiota of been conducted in this discipline (Vasco-Palacios, 2021).
Delprete PG, Forneris G, Pistarino A (2002) Carlo Bertero (1789-1831) in López-Quintero CA, Straatsma G, Franco AE, Boekhout T (2012)
the New World. Sida, 20(2), 621-644. Macrofungal diversity in Colombian Amazon forests varies with
Dennis RW (1970) Fungus flora of Venezuela and adjacent countries. regions and regimes of disturbance. Biodiversity and Conservation,
London: Stationery Office Books. 584p. 21(9), 2221–2243. doi: 10.1007/s10531-012-0280-8
Díaz Piedrahita S (1991) La Botánica en Colombia, hechos notables en López-Quintero CA, Atanasova L, Franco-Molano AE, Gams W, Komon-
su desarrollo. Academia Colombiana de Ciencias Exactas, Físicas y Zelazowska M, Theelen B, Druzhinina I (2013) DNA barcoding survey
Naturales, Colección Enrique Pérez Arbeláez 6. Santafé de Bogotá. of Trichoderma diversity in soil and litter of the Colombian lowland
Diaz-Escandón D, Soto-Medina E, Lücking R, Silverstone-Sopkin PA (2016) Amazonian rainforest reveals Trichoderma strigosellum sp. nov. and
Corticolous lichens as environmental indicators of natural sulphur other species. Antonie van Leeuwenhoek, 104(5), 657-674.
emissions near the sulphur mine El Vinagre (Cauca, Colombia). The Lücking R, Dal Forno M, Moncada B, Coca LF, Vargas-Mendoza LY, Aptroot
Lichenologist, 48(2), 147-159. doi: 10.1017/S0024282915000535 A, Arias LJ, Besal B, Bungartz F, Cabrera-Amaya DM, Cáceres ME (2017)
Dumont KP, Buriticá P, Forero E (1978) Los Hongos De Colombia-I. Turbo-taxonomy to assemble a megadiverse lichen genus: seventy
Caldasia, 12(57),159-164. new species of Cora (Basidiomycota: Agaricales: Hygrophoraceae),
Franco-Molano AE, Vasco-Palacios AM, López-Quintero C, Boekhout T honouring David Leslie Hawksworth›s, seventieth birthday. Fungal
(2005) Macrohongos de la región del Medio Caquetá. Guía de campo. Diversity, 84(1), 139-207. doi: 10.1007/s13225-016-0374-9
Medellin: Multimpresos. 219p. Lücking R, Dal-Forno M, Sikaroodi M, Gillevet PM, Bungartz F, Moncada
Fuhrmann O, Mayor E (1914) Voyage d’exploration scientifique en B, Yánez-Ayabaca A, Chaves JL, Coca LF, Lawrey JD (2014) A
Colombie par Dr O. Fuhrmann et Dr Eug. Mayor Voyage d’exploration single macrolichen constitutes hundreds of unrecognised species.
scientifique en Colombie. Mémoires de la Société des Sciences Proceedings of the National Academy of Sciences U S A, 111(30),
Naturelles de Neuchâtel, 5, 1087. 11091-11096. doi: 10.1073/pnas.1403517111
Galvis-Pérez Y, Cano Restrepo LE, Vásquez Valencia MF (2013) History Lücking R, Moncada B, Soto-Medina E, Simijaca D, Sipman HJM (2021)
of medical micology in Colombia, 1930-1970. Iatreia, 26(2), Actualización nomenclatural y taxonómica del Catálogo de Líquenes
221-231. de Colombia. Revista de la Academia Colombiana de Ciencias Exactas,
Gaya E, Vasco-Palacios AM, Vargas-Estupiñán N, Lücking R, Carretero Físicas y Naturales, 45(174), 147-189.
J, Sanjuan T, Moncada B, Allkin B, Bolaños-Rojas AC, Castellanos- Llanos C, Kjøller A (1976) Changes in the flora of soil fungi following oil
Castro C, Coca LF, Corrales A, Cossu T, Davis L, dSouza J, Dufat A, waste application. Oikos, 27, 377-382. doi: 10.2307/3543456
Franco-Molano AE, García F, Gómez-Montoya N, González-Cuellar FE, Martin GW (1937) New or Noteworthy Fungi from Panama and Colombia. I.
Hammond D, Herrera A, Jaramillo-Ciro MM, Lasso-Benavides C, Mira Mycologia, 29, 618-625.
MP, Morley J, Motato-Vásquez V, Niño-Fernández Y, Ortiz-Moreno ML, Martin GW (1938) New or Noteworthy Fungi from Panama and Colombia II.
Peña-Cañón ER, Ramírez-Castrillón M, Rojas T, Ruff J, Simijaca D, Mycologia, 30, 431-441.
Sipman HJM, Soto-Medina E, Torres G, Torres-Andrade PA, Ulian T, Martin GW (1939a) New or Noteworthy Fungi from Panama and Colombia
White K, Diazgranados M (2021) ColFungi: Colombian resources for III. Mycologia, 31, 239-249.
Fungi Made Accessible. Richmond: Royal Botanic Gardens, Kew. 36p. Martin GW (1939b) New or Noteworthy Fungi from Panama and Colombia
Garcia C, Franco MA, Quintero J, Rao IM (2000) Catalogo de cepas de IV. Mycologia, 31, 507-518.
micorrizas arbusculares, documento de trabajo 182. Cali: Centro Moncada B, Lücking R, Suárez A (2014) Molecular phylogeny of the genus
Internacional de Agricultura Tropical. 78p. Sticta (lichenised Ascomycota: Lobariaceae) in Colombia. Fungal
Gómez-Gutiérrez A (2011) La expedición helvética: viaje de exploración Diversity, 64(1), 205-231. doi: 10.1007/s13225-013-0230-0
científica por Colombia en 1910 de los profesores Otto Fuhrmann y Mueller GM, Wu Q (1997) Mycological contributions of Rolf Singer: field
Eugène Mayor. Editorial Pontificia Universidad Javeriana. itinerary, index to new taxa, and list of publications. Publication/Field
González-Cuellar FE, Lasso-Benavides CM, Adrada-Gómez BY, Sanabria- Museum of Natural History.
Diago OL, Vasco-Palacios AM (2021) Estudio etnomicológico con tres Nowak R, Winkler S (1970) Foliicole flechten der sierra nevada de
comunidades rurales ubicadas en la zona andina del departamento santa marta (kolumbien). und ihre gegenseitigen beziehungen.
del Cauca, Colombia. Boletín de Antropología, 36(62), 147-164. Österreichische Botanische Zeitschrift, 118(5), 456-485.
Guerrero Forero E (1996) Micorriza: fundamentos biológicos y estado Nowak R, Winkler S (1975) Foliicolous lichens of Choco, Colombia, and
del arte (No. Doc. 15660). Bogotá DC: CO-BAC. their substrate abundances. The Lichenologist, 7(1), 53-58.
Guzmán G, Varela L (1978) Los Hongos de Colombia III. Observaciones Nylander W (1863) Enumeration des plantes de la Nouvelle Grenade. In:
sobre los hongos, líquenes y mixomicetos de Colombia. Caldasia, Triana J, Planchon JE. Prodromus Florae Novo Granatensis. Annales
12(58), 309-338. des Sciences Naturelles, série 4 (Botany), 302-305.
Hooker WJ (1822) Lichenes. In: Kunth CS. Synopsis plantarum, quas in Pardo-Cardona VM (2001) Historia, estado actual y perspectivas de la
itinere ad plagam aequinoctialem orbis novi, vol 1. Paris: FG Levraut. investigación de los Uredinales en Colombia. Revista Facultad Nacional
491p. de Agronomía Medellín, 54(1-2), 1333-1350.
Houbraken J, Lopez-Quintero CA, Frisvad, JC, Boekhout T, Theelen B, Peña-Cañón ER, Enao-Mejía LG (2014) Conocimiento y uso tradicional
Franco AE, Samson RA (2011). Penicillium araracuarense sp. nov., de Hongos silvestres de las comunidades campesinas asociadas a
Penicillium elleniae sp. nov., Penicillium penarojense sp. nov., bosques de roble (Quercus humboldtii) en la zona de influencia de la
Penicillium vanderhammenii sp. nov. and Penicillium wotroi sp. nov., Laguna de Fúquene. Etnobiología 12 (3): 28-40.
isolated from leaf litter. International Journal of Systematic and Peña-Venegas CP, Cardona GI, Mazorra A, Arguellez JH, Arcos AL (2006)
Evolutionary Microbiology, 61(6), 1462-1475. Micorrizas Arbusculares de la Amazonia Colombiana, 1st ed. Bogotá
IAVH - Instituto de Investigación de Recursos Biológicos Alexander von DC: Instituto Amazónico de Investigaciones Científicas.
Humboldt (2019) Especies de líquenes priorizadas en Colombia. Peña-Venegas CP, Vasco-Palacios AM (2019) Endo-and Ectomycorrhizas
Available at: http://i2d.humboldt.org.co/ceiba/resource.do?r=rrbb_ in tropical ecosystems of Colombia. In: Pagano M, Lugo M (eds).
liquenes_2019 Mycorrhizal Fungi in South America. Fungal Biology. Springer, Cham.
Landaburu J, Pineda R (1984) Tradiciones de la gente del hacha. Pp. 111-146. doi: 10.1007/978-3-030-15228-4_6
Mitología de los indios Andoques del Amazonas. Bogotá: Instituto Pérez de Barradas J (1954) Orfebrería prehispánica de Colombia, Estilo
Caro y Cuervo. Unesco. 290p. Calim. Tallerers Gráficos Jura. 367p.
Leal A, Rojas JL, Valencia-Islas NA, Castellanos L (2018) New Pulido-L MM (1983) Estudios en agaricales colombianos: los hongos de
β-orcinol depsides from Hypotrachyna caraccensis, a lichen Colombia IX. Biblioteca “Jose Jeronimo Triana”. Instituto de Ciencias
from the páramo ecosystem and their free radical scavenging Naturales. 143p.
activity. Natural Product Research, 32(12), 1375-1382. doi: Pulido-L MM, Boekhout T (1989a) Distribución de los macromycetos
10.1080/14786419.2017.1346639 en el Transecto Parque los Nevados. In: Van der Hammen T, Diaz-
Léveillé JH (1863) Fungi. In: Triana J, Planchon JE. Prodromus Florae Piedrahita S, Alvarez VJ (eds). La Cordillera Central Colombiana
Novo Granatensis. Annales des Sciences Naturelles, série 4 (Botany), Transecto Parque los Nevados (Segunda Parte). Berlin: Borntraeger
282-300. Verlagsbuchhandlung. Pp. 485-505.
Pulido-L MM, Boekhout T (1989b) La ocurrencia de los macromycetos y sus Simijaca D, Moncada B, Lücking R (2018) Bosque de roble o plantación
hábitats en vegetaciones a lo latgo del Transecto Parque Los Nevados. de coníferas, ¿qué prefieren los líquenes epífitos? Colombia Forestal,
In: Van der Hammen T, Diaz-Piedrahita S, Alvarez VJ (eds). La Cordillera 21(2), 123-141. doi: 10.14483/2256201X.12575
Central Colombiana Transecto Parque los Nevados (Segunda Parte). Sipman HJM, Aguirre-C J (1982) Contribución al conocimiento de los
Berlin: Borntraeger Verlagsbuchhandlung. Pp.507-532. líquenes de Colombia—I. Clave genérica para los líquenes foliosos y
Pulido-L MM (2005) Distribución y ecologia de los Macromicetes fruticosos de los páramos colombianos. Caldasia, 13(64), 603-634.
coleccionadas a lo largo del transecto de Tatamá (datos iniciales). En Sipman HJM (1986) Lichens of the Buritaca — la Cumbre Transect Sierra
La Cordillera Occidental Colombiana: Transecto Tatamá. Estudios de Nevada de Santa Marta, Colombia. Estudios de Sistemas Tropandinos,
Sistemas Tropandinos, 6. Berlin: Borntraeger Verlagsbuchhandlung. 2, 185-188.
Pp. 249-265. Sipman HJM (1989) Lichen zonation in the Parque Los Nevados transect.
Pulido-L MM (2008) Distribución y ecología de los macromycetes a Studies on Tropical Andean Ecosystems, 3, 461-483.
lo largo del transecto Sumapaz (datos iniciales). Distribution and Sipman HJM (2005) Los líquenes del transecto Tatamá. La Cordillera
ecology of the macromycetes along the Sumapaz transect (initial Occidental Colombiana: Transecto Tatamá. Estudios de Sistemas
data). In: der Hammen V (ed). Studies on Tropical Andean Ecosystems, Tropandinos, 6, 1-956.
7. Berlin: Borntraeger Verlagsbuchhandlung. Pp. 239-274. Sipman HJM, Aguirre-C J (2016) Líquenes. In: Bernal R, Gradstein R, Celis
Rocheraux H (1959) Documentos redactados en el dialecto de las tribus M (eds). Catálogo de Plantas y Líquenes de Colombia, 1st ed. Bogotá
Tunebas, radicadas en el triángulo de Cubugon, Royota, Nevado de DC: Universidad Nacional de Colombia.
Chita. Revista Colombiana de Antropología, 8, 15-119. Sipman HJM, Hekking W, Aguirre-C J (2008) Checklist of lichenised and
Rogers AL (1971) Isolation of keratinophilic fungi from soil in the vicinity lichenicolous fungi from Colombia. Bogota DC: Universidad Nacional
of Bogota, Colombia. Mycopathologia et Mycologia Applicata, 44(3), de Colombia. 242p.
261-264. Soto-Medina E, Lücking R, Bolaños Rojas A (2012) Especificidad de forófito
Rojas JL, Díaz-Santos M, Valencia-Islas NA (2015) Metabolites with y preferencias microambientales de los líquenes cortícolas en cinco
antioxidant and photo-protective properties from Usnea roccellina forófitos del bosque premontano de finca Zíngara, Cali, Colombia.
Motyka, a lichen from Colombian Andes. Journal of Pharmaceutical Revista de Biología Tropical, 60(2), 843-856.
and Biosciences, 3(4), 18-26. doi: 10.20510/ukjpb/3/i4/89454 Triana JJ, Planchon J (1863-1867) Prodromus florae novo-granatensis.
Romero JFG, Fernandez DCG, de Restrepo MC (2004) Incidencia de las Cryptogamie. Commission chorographique de la Nouvelle-Grenade.
micorrizas arbusculares y vesículo arbusculares como estrategia Paris.
adaptativa de especies de páramo y selva altoandina, cordillera Urbina F (1986) Amazonia, Naturaleza y Cultura. Bogota DC: Banco de
oriental de Colombia. Colombia Forestal, 8(17), 43-59. Occidente.
Rubiano LJ (1988) Líquenes como indicadores de contaminación en el Vargas N, Restrepo S (2019). Macrohongos (Basidiomycota y Ascomycota)
Complejo Industrial de Betania y la Termoeléctrica de Zipaquira, de los ecosistemas exocársticos de el Peñón (Andes), Santander,
Cundinamarca. Acta Biológica Colombiana, 1(4), 95-125. Colombia. In: Lasso et al., (eds). Biodiversidad subterránea y epigea
Rubiano LJ (1987) Delimitación en áreas de isocontaminación en Cali y de los sistemas cársticos de El Peñón (Andes). Santander: Humboldt.
Medellín utilizando líquenes como bioindicadores. Pérez-Arbelaezia, Vasco-Palacios AM (2021) Etnomicología. Boletín Semillero de Investigación
1 (4-5), 7-41. en Etnobiología Universidad del Cauca, 1, 61-54.
Sánchez de Prager M (2007) Las endomicorrizas: Expresión Bioedáfica de Vasco-Palacios AM, Franco-Molano AE (2013) Diversity of Colombian
importancia en el trópico (No. Doc. 27034) CO-BAC, Bogotá. macrofungi (Ascomycota - Basidiomycota). Mycotaxon, 121, 499.
Sánchez de Prager M, Posada R, Velásquez D, Narváez M (2010) Vasco-Palacios AM, Bahram M, Boekhout T, Tedersoo L (2020) Carbon
Metodologías básicas para el trabajo con micorriza arbuscular y content and pH as important drivers of fungal community structure in
hongos formadores de micorriza arbuscular. Universidad Nacional de three Amazon forests. Plant and Soil, 450(1), 111-131. doi: 10.1007/
Colombia. Palmira. s11104-019-04218-3
Sanjuan T (1999) La Diversidad del género Cordyceps en hormigas del Vasco-Palacios AM, Hernandez J, Peñuela MC, Boekhout T (2018)
bosque húmedo tropical de Colombia. Undergraduation thesis. Ectomycorrhizal fungi diversity in a white sand forest in western
Universidad Nacional de Colombia. Santafé de Bogotá. Amazonia. Fungal Ecology, 31, 9-18. doi: 10.1016/j.funeco.2017.10.003
Sanjuan T, Kepler RM, Spatafora JW, Tabima J, Vasco-Palacios AM, Vasco-Palacios AM, Suaza SC, Castanõ-Betancur M, Franco-Molano AE
Restrepo S (2015) Five new species of entomopathogenic fungi from (2008) Conocimiento etnoecólogico de los hongos entre los indígenas
the Amazon and evolution of neotropical Ophiocordyceps. Fungal Uitoto, Muinane y Andoke de la Amazonía Colombiana. Acta Amazónica,
Biology, 119(10), 901-916. doi: 10.1016/j.funbio.2015.06.010 38, 17-30.
Schenck NC, Spain JL, Sieverding E, Howeler RH (1984) Several new and Veerkamp J, Gams W (1983) Los Hongos De Colombia—VII Some new
unreported vesicular-arbuscular mycorrhizal fungi (Endogonaceae) species of soil fungi from Colombia. Caldasia, 13(65), 709-717.
from Colombia. Mycologia, 76, 685-699. Velandia C, Galindo L, Mateus K (2008) Micolatría en la iconografía
Schultes RE, Bright A (1979) Ancient gold pectorals from Colombia: prehispánica de América del Sur. International Journal of South
Mushroom effigies? Botanical Museum Leaflets Harvard University, 27 American Archaeology, 3, 6-13.
(5/6), 113-141. Villalobos S, Mengual M, Henao-Mejía LG (2017) Uso de los Hongos, Podaxis
Schultes RE, Hofmann A, Ratsch C (2000) Plantas de los dioses: orígenes pistillaris, Inonotus rickii y Phellorinia herculeana (basidiomycetes), por
del uso de los alucinógenos. Fondo de Cultura Económica. la Etniaa Wayuu en la Alta Guajira Colombiana. Etnobiologia, 15(1),
Serrano M (1943) Blastomicosis. Revista Medicina y Cirugía, VII(11), 393- 64-73.
422. Wolf JH (1993) Diversity patterns and biomass of epiphytic bryophytes and
Sierra MA, Danko DC, Sandoval TA, Pishchany G, Moncada B, Kolter R, lichens along an altitudinal gradient in the northern Andes. Annals of
Mason CE, Zambrano MM (2020) The microbiomes of seven lichen the Missouri Botanical Garden, 80(4), 928-960. doi: 10.2307/2399938
genera reveal host specificity, a reduced core community and potential Wolf JH (1994) Factors controlling the distribution of vascular and non-
as a source of antimicrobials. Frontiers in Microbiology, 11, 398. doi: vascular epiphytes in the northern Andes. Vegetation, 112(1), 15-28.
10.3389/fmicb.2020. 00398 Wolf JH (1995) Non-vascular epiphyte diversity patterns in the canopy
Sieverding E (1984) Aspectos básicos de la investigación de MVA. In: of an upper montane rain forest (2,550-3,670 m), central cordillera,
Investigaciones sobre micorrizas en Colombia. Universidad Nacional Colombia. Selbyana, 16(2), 185-195.
de Colombia. Yilmaz N, López-Quintero CA, Vasco-Palacios AM, Frisvad JC, Theelen B,
Sieverding E (1989a) La micorriza: Un componente biotecnológico en la Boekhout T, Teelen B, Boekhout T, Samson R, Houbraken J (2016).
producción vegetal. Revista Colombia Ciencia y Tecnología, 7, 9-11. Four novel Talaromyces species isolated from leaf litter from Colombian
Sieverding E (1989b) Ecology of VAM fungi in tropical ecosystems. Amazon rain forests. Mycological Progress, 15(10), 1041-1056.
Agriculture, Ecosystems and Environment, 29, 369-390.
Sieverding E, Howeler RH (1985) Influence of spores of VA mycorrhizal
fungi on cassava yield response to phosphorus fertilisation. Plant and
Soil, 88, 213-221.
Rhytidhysteron columbiense
[Robert Lücking]
44 CATALOGUE OF FUNGI OF COLOMBIA
Chapter 3
Diversity of Fungi of Colombia
Ester Gaya1, Viviana Motato-Vásquez2,3 & Robert Lücking4*
1
Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3DS, UK.
2
Grupo de Investigación en Biología de Plantas y Microorganismos (BPM), Departamento de Biología, Facultad de Ciencias Naturales y Exactas, Universidad
del Valle, Calle 13 # 100-00, Cali, Valle del Cauca, Colombia.
3
Calima, Fundación para la Investigación de la Biodiversidad y Conservación en el Trópico, Cali, Colombia.
4
Botanischer Garten und Botanisches Museum, Freie Universität Berlin, Königin-Luise-Straße 6–8, 14195 Berlin, Germany.
ABSTRACT
This chapter provides a brief overview of the diversity of fungi of Colombia, both in comparison with other organisms and
in a global context, including an assessment of the current state of knowledge and an estimate of the actual species
richness. The current checklist of fungi of Colombia, including lichenised and fungus-like organisms unrelated to the true
Fungi, comprises 7,241 species. Assessments of biodiversity-rich countries are usually based on plants and vertebrates,
but here we provide an expanded perspective, including fungi and selected invertebrates. The documented diversity of fungi
of Colombia is lower than that of plants, vertebrates, and insects, which we attribute to the lack of rigorous taxonomic
studies and systematic inventories. Colombia ranks second in plant and vertebrate diversity in the world, trailing only Brazil.
However, in terms of known fungal diversity, Colombia is not among the top ten countries and even far behind smaller
temperate countries, such as France, Italy, the UK, Germany, and Japan, again highlighting the need for much additional
work. Estimates of the existing species richness of fungi of Colombia oscillate between 27,430 and 380,000 species,
depending on the extrapolation method. Although these numbers may seem exaggerated, currently reported numbers
for the United States already exceed 45,000 and estimates for Mexico predict up to 260,000 species. To catalogue the
diversity of the Colombian funga fully, a thorough assessment is needed, including molecular studies of presumably known
taxa that may include morphologically cryptic species and effective detection methods for ecologically hidden fungi.
RESUMEN
Este capítulo proporciona una breve introducción a la diversidad de hongos de colombia, tanto en comparación con otros
organismos como en un contexto global, incluyendo una evaluación del estado actual del conocimiento y una estimación
de la riqueza real de las especies. El listado actual de los hongos de colombia comprende 7.241 especies, incluyendo
especies liquenizadas y organismos similares a los hongos pero no relacionados con los hongos verdaderos, tales como
los oomicetes y los mixomicetes. Si bien las evaluaciones de los países ricos en biodiversidad generalmente se basan
en plantas y vertebrados, aquí brindamos una perspectiva ampliada, la cual incluye los hongos e también invertebrados
seleccionados. La diversidad documentada de hongos de colombia es menor que la de plantas (26,150 especies),
vertebrados (7,607) e insectos (68,000), lo que atribuimos a la falta de estudios taxonómicos rigurosos e inventarios
sistemáticos. Colombia ocupa el segundo lugar en diversidad de plantas y vertebrados en el mundo, solo detrás de Brasil.
Sin embargo, en términos de diversidad fúngica conocida, Colombia no se encuentra entre los diez primeros países
del mundo e incluso muy por detrás de algunos países templados más pequeños, como Francia (24,840 especies),
Italia (22,700), el Reino Unido (20,000), Alemania (15,295) y Japón (14,000), lo que también destaca la necesidad de
mucho trabajo adicional. Las estimaciones de la riqueza de especies existentes de hongos de colombia oscilan entre
27,430 y 380,000 especies, según el método de extrapolación. Si bien estos números pueden parecer exagerados, los
números reportados actualmente para los Estados Unidos ya superan las 45,000 y las estimaciones para México predicen
hasta 260,000 especies. Para catalogar completamente la diversidad de los hongos de colombia, se necesita inventarios
exhaustivos, los cuales incluyen estudios moleculares de taxones supuestamente conocidos que pueden incluir especies
morfológicamente crípticas, tanto como métodos efectivos de detección de hongos ecológicamente escondidos, a travez
del metabarcoding ambiental.
INTRODUCTION & Thines, 2017; Burki et al., 2019; James et al., 2020).
Fungi represent one of the three major eukaryotic life forms These include the Oomycota (egg fungi or water moulds),
on our planet, besides plants and animals (Burki et al., Hyphochytridiomycota, and Labyrinthulomycetes (slime nets),
2019). Mycology, the study of fungi, was traditionally treated which form separate lineages within the Heterokontophyta
as a branch of botany, but fungi are more closely related (part of the SAR supergroup and related to brown algae and
to animals than plants (Figure 1). To further complicate diatoms), as well as the Phytomyxea within the Rhizaria (also
matters, fungi are not a natural group, with fungal life forms part of the SAR supergroup). Other fungus-like organisms are
convergently evolving several times among eukaryotes. We represented by the Myxogastria or Myxomycetes (plasmodial
can even find fungus-like bacteria in the Actinomycetales slime moulds) within the Amoebozoa, the Acrasida (cellular
(Ventura et al., 2007). Although nearly 99% of all fungi slime moulds) within the Discoba, and the Fonticulida (another
are representatives of the kingdom Fungi, fungus-like group of cellular slime moulds) within the Cristidiscoidea,
organisms are also found in various other clades (Beakes a clade basally related to the Fungi and together with the
FIGURE 1. Simplified Tree of Life highlighting the position of the true Fungi (dark brown) and other fungus-like organisms (light brown) relative
to animals (Animalia) (blue) and true plants (Viridiplantae) (dark green) and plant-like organisms (light green) (compiled after Burki et al.,
2019 and James et al., 2020, as well as various other sources for specific clades). Unranked major groups above kingdom are indicated
to the right.
latter forming the Holomycota (Figure 1). The true fungi are In terms of known species, Fungi currently rank third
named Fungi (as a scientific name written in upper case and amongst the three large kingdoms, with 150,000 species
italics) to distinguish them from other fungus-like organisms, (Lücking et al., 2021a), after Animalia (animals), with 1.66
whereas the non-scientific term “fungi” also encompasses million known species (Zhang, 2013), and Viridiplantae
all unrelated fungus-like organisms (as a vernacular name (plants), with at least 350,000 known species (Christenhusz
written in lower case and standard font; Thines et al., 2020). & Byng, 2016; The Plant List, 2021). Fungi are clearly
The convergent evolution of the fungal lifestyle within the least well-known of these three kingdoms, given that
the eukaryotes is comparable to the situation in plants estimates predict a much higher number of species,
(Burki et al., 2019; James et al., 2020): the true plants between 1.5 and 6.3 million (Blackwell, 2011; Hawksworth
are grouped in the kingdom Plantae (corresponding to the & Lücking, 2017; Baldrian et al., 2021). These estimates
unranked Viridiplantae), but other multicellular plant-like would place Fungi as the second-largest kingdom after
organisms are found in the related red (Rhodophyta) and animals, including estimates for unknown arthropods (see
brown algae (Phaeophyceae), the latter classified within below), but far ahead of plants. Predictive estimates of
the Heterokontophyta and thus more closely related to the global fungal diversity are based on diverse considerations,
Oomycota, Hyphochytridiomycota, and Labyrinthulomycetes including a mean global fungi:plant species ratio of between
(Figure 1). Fungi are sessile (like plants) and heterotrophic 6:1 and 10:1, but also taking into account accumulating
organisms that are incapable of photosynthesis (like evidence of hidden diversity among presumably known
animals), with various strategies to obtain carbohydrates. species, with a mean ratio of over 10:1, as well as a wealth
These strategies range from the decomposition of dead of data emerging from environmental sequencing studies
organic material (saprotrophs) to attacking living organisms (Baldrian et al., 2021; Lücking et al., 2021a). Cataloguing
(pathogens, parasites, or carnivores) to forming different this unknown diversity is the major challenge for mycology
types of mutualistic symbioses with phototrophic plants, in terms of time and resources and how to deal with fungi
algae, and cyanobacteria (mycorrhizae and lichens) (Dix & only known from sequence data (Lücking et al., 2021a).
Webster, 1995; Mueller et al., 2004; Piepenbring, 2015; Much of this unknown fungal diversity is expected to be
Watkinson et al., 2016). Endophytic and endolichenic fungi found in tropical regions, such as Colombia.
are ubiquitous as asymptomatic inhabitants of plants, plant-
like organisms, and lichens (Rodriguez et al., 2009; U’Ren et DIVERSITY OF FUNGI OF COLOMBIA IN THE TREE OF LIFE
al., 2012). They are often classified as symbionts (e.g., in the
FUNGuild database; Nguyen et al., 2016), but their biological The ColFungi project (Gaya et al., 2021; https://colfungi.
roles are diverse and often unknown, and they should only org) represents the first compilation of data on all groups
be considered symbionts if there is actual evidence for this of fungi of Colombia into a single resource, including
(Arnold & Lewis, 2005; Bolívar-Anillo et al., 2016; Bastias nomenclatural and taxonomic checks. This project resulted
et al., 2021). Indeed, the diversity of potential life forms of in a total of 7,241 accepted species of fungi for Colombia,
endophytic and endolichenic fungi has been dubbed the encompassing a high diversity of fungal life forms, including
“endophytic continuum” (Schulz & Boyle, 2005). mushrooms and lichen-formers, plant- and animal-
Although mostly invisible (“ecologically hidden”), fungi pathogenic microfungi, moulds, yeasts, among many others
are crucial components of most ecosystems on earth, being (Figure 2). By itself, the number of 7,241 species is difficult
particularly involved in nutrient cycling (Dix & Webster, 1995; to evaluate, but when put into a taxonomic and geographic
Watkinson et al., 2016). In addition, they have played, and context, it illustrates the current state of knowledge of the
continue to play, important roles in ancient and modern Colombian funga.
cultures (Guzmán et al., 2003; Dugan, 2008). These Country- or region-based comparisons of biodiversity
include their uses as food, in industrial, biotechnological, are often presented in the context of global conservation
and pharmaceutical applications, and in biomonitoring and approaches, but these listings are usually limited to plants
mycoremediation (Nimis et al., 2002; Rhodes, 2014; Hyde et and vertebrates (mammals, birds, reptiles, amphibians,
al., 2019; see also Chapter 13). and fish; e.g., Butler, 2016, 2019), whereas fungi are rarely
Fungi also hold several records among living organisms. included in such comparisons. Colombia is currently ranked
For instance, species of Armillaria (honey mushrooms) form second among the ten most biodiversity-rich countries in
several of the largest known single organisms on earth, plant and vertebrate species richness, trailing Brazil but
with biomasses corresponding to up to three blue whales ahead of Indonesia, China, Mexico, Peru, Australia, India,
(Smith et al., 1992; Schmitt & Tatum, 2008; Anderson et al., Ecuador, and Venezuela (Butler, 2016, 2019). According to
2018). The widespread Schizophyllum commune (split gill) these listings, Colombia has 51,220 plant species, 2,053
and Coprinopsis cinerea (gray shag), both model fungi, have fish, 1,878 birds, 771 amphibians, 601 reptiles, and 442
been shown to develop more than 20,000 mating types mammals, and updated numbers for all vertebrate groups
or “sexes” (Kothe, 1999; Casselton & Kües, 2007). The are consistently higher (Table 1). Notably, with 7,241 known
lichenised fungus Buellia frigida in dry valleys of Antarctica species, fungi would rank second in this enumeration
forms thalli that are among the oldest documented for Colombia.
organisms on earth, in part estimated at over 6,500 years Still, this ranking would be misleading: plants and fungi
old (Green et al., 2012). essentially represent entire kingdoms, whereas, in terms of
A B C D E
F G H I
J K L M
N O P Q
R S T U V
W X Y Z
FIGURE 2. Examples of fungi of Colombia representing a diversity of lifestyles and phenotypes of spore-bearing structures. A Campanella
caesia. B Cordyceps nidus. C Laetiporus sulphureus. D Candida sp. E Auricularia mesenterica. F Tolypocladium capitatum. G Laccaria
laccata. H Boletus sp. I Yoshimuriella peltigera. J Scleroderma flavidum. K Panaeolus semiovatus. L Cyathus striatus. M Phillipsia
domingensis. N Rhytidhysteron columbiense. O Cerioporus scutellatus. P Hericium erinaceus. Q Hydnopolyporus fimbriatus. R Leotia lubrica.
S Geastrum pectinatum. T Cora elephas. U Rhodotorula sp. V Calostoma cinnabarinum. W Trametes versicolor. X Collybia plectophylla.
Y Chlorociboria aeruginascens. Z Phallus indusiatus. Photographs: A, B, C, F, G, I, J, N, O, Q, T, W, Y: Robert Lücking. D, U: Mauricio Ramírez.
E, L, R, S: Aída Vasco-Palacios. H, V: Bibiana Moncada. K, M, P, Z: Ana Esperanza Franco-Molano. X: Ana Cristina Bolaños.
TABLE 1. Updated totals for the number of known species of selected groups of organisms in Colombia. The global totals exclude fossil taxa.
Butler Proportion of
Updated
Group (2016, Global total updated figures Reference(s) for updated figures
figures
2019) vs global total
Animals — — 1,530,000 — Zhang (2013); this paper
Vertebrates 5,745 7,607 57,102 13.2% This paper
Mejia & Acero (2002); Maldonado-Ocampo et al. (2008);
Fish 2,053 3,700 32,000 11.6%
Nelson et al. (2016); DoNascimento et al. (2018)
Amphibia 771 836 8,348 10.0% Acosta-Galvis (2019); Frost (2021)
Reptiles 601 634 11,440 5.5% Uetz et al. (2021)
Birds 1,878 1,909 9,159 20.8% Barrowclough et al. (2016); Avendaño et al. (2017)
Mammals 442 528 6,451 8.2% Burgin et al. (2018); Mammal Diversity Database (2021)
Invertebrates
Insects — 68,000 1,000,000 6.8% This paper
Papilionoidea — 3,642 18,768 19.4% Garwood et al. (2021)
Formicidae — 1,100 13,809 8.0% Fernández et al. (2019); Bolton (2020)
Staphylinidae — 800 48,000 1.7% Newton et al. (2005)
Cicadellidae — 679 22,000 3.1% Freytag & Sharkey (2002)
Odonata — 335 6,000 5.6% Pérez-Gutiérrez & Palacino-Rodríguez (2011)
Blattaria — 133 4,330 3.1% Vélez (2008)
Arachnida — 1,546 57,126 2.7% Perafán et al. (2013)
Bernal et al. (2016a, b, 2019); Diazgranados et al. (2021);
Plants 51,220 26,150 350,000 7.5%
The Plant List (2021)
Bernal et al. (2016a, b, 2019); Diazgranados et al. (2021);
Vascular plants — 24,500 316,000 7.8%
The Plant List (2021)
Bernal et al. (2016a, b, 2019); Diazgranados et al. (2021);
Bryophytes — 1,650 34,500 4.8%
The Plant List (2021)
Fungi — 7,241 150,000 4.8% Gaya et al. (2021); Lücking et al. (2021a); this paper
Ascomycota — 4,564 93,632 4.9% Gaya et al. (2021); Kirk (pers. comm. July 2021); this paper
Basidiomycota — 2,318 50,460 4.7% Gaya et al. (2021); Kirk (pers. comm. July 2021); this paper
species richness, vertebrates are only a small fraction of globally (Bolton, 2020). The most comprehensive and most
the animal kingdom, the diversity of which is concentrated recent checklist of Colombian butterflies (Papilionoidea)
among arthropods (Stork, 2018). Indeed, the known insect compiled data on 3,642 species out of a global number of
diversity of Colombia alone should easily surpass that of 18,768 (Garwood et al., 2021). According to Stork (2018),
vertebrates and likely even plants. For instance, Newton et about one million insect species have been named, which
al., (2005) listed nearly 800 species only of Staphylinidae means that globally, Staphylinidae corresponds to 4.8%,
(rove beetles) for the country, out of a total of 48,000 known Papilionoidea 1.9%, Formicidae 1.4%, Odonata 0.6%, and
globally, and Vélez (2008) enumerated 133 species of Blattaria a little over 0.4% of that number. Extrapolating from
Blattaria (cockroaches), compared to 4,330 worldwide (Table these proportions in the case of Colombia implies that a
1). Pérez-Gutiérrez & Palacino-Rodríguez (2011) reported total of between 17,000 and 194,000 insect species are
335 species of Odonata (dragonflies), out of a global total of likely known from the country, with a median of 43,350 and
around 6,000. For Formicidae (ants), Fernández et al., (2019) a mean of 68,000 (Table 1). Although the exact number has
listed 1,100 species for Colombia, compared to over 13,800 not yet been compiled, this extrapolation is far ahead of the
7,241 fungi and even ahead of the actual known number of a result, in biodiversity-rich countries, such as Colombia,
plant species (see below). even though the number of fungal species is expected to be
Compared to the 51,220 species of plants reported for higher than in extra-tropical areas, the absolute number of
Colombia (Butler, 2016, 2019), the number of 7,241 fungal mycological studies is much lower. For instance, a search
species also appears relatively low, with a fungus:plant in the Web of Science [https://www.webofscience.com/
species ratio of 1:7. However, the figure of 51,220 plant wos/woscc/basic-search], using the string <(ALL=(Fungi))
species goes back to outdated sources, being recently AND ALL=(Country)>, retrieved around 16,000 results
corrected to about 26,150 species in the Catálogo de Plantas between 1900 and 2020 for the U.K. (England, Northern
y Líquenes de Colombia (Bernal et al., 2016a, b, 2019) and Ireland, Scotland, Wales), but only 1,155 for Colombia.
the most recent compilation of ColPlantA (Diazgranados Unfortunately, reliable estimates of known fungal diversity
et al., 2021). Based on the corrected number, Colombia’s are difficult to obtain for most countries due to the lack of
fungus:plant species ratio currently amounts to about comprehensive databases or checklists, exceptions being
1:3.6, still much lower than ratios of between 6:1 and 10:1 lichenised fungi and macrofungi (Feuerer & Hawksworth,
obtained from well-studied extra-tropical areas (Hawksworth 2007; Gaya et al., 2021). Also, the nature and detail
& Lücking, 2017). In terms of global diversity, Colombia of existing compilations may differ markedly between
harbours 4.8% of the 150,000 known fungal species and countries, making comparisons challenging. Nevertheless,
7.5% of the 350,000 known plant species (Table 1). While we attempted to compile existing data to compare Colombia
these percentages are in the same order of magnitude, they with selected countries within and outside the tropics (see
do not consider the much more limited state of knowledge of Gaya et al., 2021; here updated).
global fungal diversity. The highest number of fungi, including all lifestyles,
Fungi and fungus-like organisms are currently arranged has been reported for the United States (Table 2), with
into 64 classes (Species Fungorum 2021), of which 36 have approximately 45,000 species (Perlmutter & Weakley,
been reported from Colombia. The remaining 28 classes are 2018; Bates et al., 2018; Esslinger, 2019). However, this
species-poor, with between one and 321 species globally. impressive number is tentative, as, for non-lichenised
These remaining classes potentially occur in Colombia fungi, current names and synonyms have not been fully
but are challenging to detect, as they largely represent sorted out. After India, with 29,000 species, France
ecologically hidden microfungi. Among the 36 classes remarkably ranks third, with 24,840 species, followed
present in Colombia, 11 are overrepresented compared to by China (23,927) and Italy (22,700) (Roux, 2012; Onofri
global proportions, by a factor of up to 6.5 (Figure 3). The et al., 2013; Medardi, 2006; Venturella et al., 2011;
seven most overrepresented classes are Glomeromycetes Nimis, 2016; Institute of Microbiology, Chinese Academy
(factor 6.5), Coniocybomycetes (3.9), Mortierellomycetes of Sciences, 2018; Gargominy et al., 2020; Wang et al.,
(4.3), Schizosaccharomycetes (4.0), Lecanoromycetes 2020; Wei, 2021). We had originally reported 20,500
(3.8), Candelariomycetes (3.8), and Arthoniomycetes (3.6). species for Brazil (Gaya et al., 2021), but a reanalysis of the
Four of these seven classes have low species numbers in underlying data, including accepted names vs synonyms,
Colombia, between one and 26, so the observed deviations led to a correction, with now 13,950 species (see Table
are not meaningful, but the three remaining classes are 2). This number is still substantially higher than the 5,719
represented by higher species numbers: Lecanoromycetes species reported by Maia et al., (2015), the latter likely
(2,167), Arthoniomycetes (189) and Glomeromycetes (106). an underestimate based on an incomplete list of selected
The substantial overrepresentation of Lecanoromycetes records for particular groups.
and Arthoniomycetes is because these mostly lichen- Among the countries assessed here, Colombia ranks
forming fungi are much better studied than any other seventh for lichenised fungi (see Chapter 6). In comparison,
larger group of fungi in Colombia (Gaya et al., 2021), so the overall number and particularly the number of non-
the observed deviations are based on study bias. On the lichenised fungi is comparatively low, with Colombia currently
other hand, five important classes that are highly diverse taking 13th place. In terms of species density and species
in tropical regions but substantially underrepresented per log10(area), Colombia also ranks comparatively low,
and understudied in Colombia are Dothideomycetes, particularly when compared to its richness in other groups
Sordariomycetes, Leotiomycetes, Pezizomycetes, and of organisms, leading us to the question: how many species
Laboulbeniomycetes (Figure 3). of fungi can we expect in Colombia?
DIVERSITY OF FUNGI OF COLOMBIA IN A GLOBAL CONTEXT HOW MANY SPECIES OF FUNGI ARE THERE IN COLOMBIA?
Knowledge on organismic diversity is unevenly distributed Predictive estimates of species richness are naturally
across the planet, and even the absolute state of speculative, especially for ecologically largely hidden
knowledge is opposed to the estimated diversity in a organisms and in groups where taxon concepts have not
region. This scenario is largely because there are more been fully resolved. Several approaches exist to predict
specialists per area working in regions with fewer species species numbers on the basis of known species and
per area for almost any group of organisms, particularly other data. One possible approach is the assumption that
fungi (Hawksworth, 2001; Piepenbring et al., 2018). As Colombia’s species per log10(area) estimate should be
FIGURE 3. Comparison of the proportional phylogenetic diversity of Fungi and fungus-like organisms at the class level between Colombia
and the global funga. The percentages indicate the number of species per class compared to the corresponding total. Selected classes
that are over- or underrepresented in Colombia are marked in dark grey, and the corresponding percentages are indicated. For detailed
data, see Suppl. File S1.
TABLE 2. Comparison of numbers of known fungal species between selected countries. The number of 16,900 macrofungi for the United
States is an estimate based on Bates et al. (2018). The corrected total for Brazil (13,950; as compared to 20,500 reported in Gaya et al.,
2021) is based on the accepted number of fungal names for Brazil on SpeciesLink (approximately 11,330), minus the number of lichenised
species in that database (approximately 1,690), plus the updated number of lichenised species based on Aptroot (2021).
Density
Country / All Lichenised Macro-
[species / Reference(s)
Territory fungi fungi fungi
log10(km2)
United Perlmutter & Weakley (2018); Bates et al. (2018); Esslinger
45,000 4,341 16,900 6,578
States (2019)
Sarbhoy et al. (1996); Singh & Sinha (2010); Manoharachary &
India 27,000 2,714 — 4,450
Nagaraju (2016); Sinha et al. (2018)
France 24,840 2,917 — 4,276 Roux (2012); Gargominy et al. (2020)
Institute of Microbiology, Chinese Academy of Sciences (2018);
China 23,927 3,050 7,138 3,427
Wang et al. 2020; Wei (2021)
Onofri et al. (2013); Medardi (2006); Venturella et al. (2011);
Italy 22,700 2,704 4,198 4,143
Nimis (2016); Checklist of Italian Fungi (2021)
United Legon & Henrici (2005); Smith et al. (2009); Dines & McCarthy
20,000 2,000 3,100 3,343
Kingdom (2014); Clubbe et al. (2020)
Germany 15,295 1,946 6,120 2,754 Wirth et al. (2011); DGfM (2021)
Japan 14,000 1,906 4,160 2,510 Hosoya (2006); Katumoto (2009); Ohmura & Kashiwadani (2018)
Lewinsohn & Prado (2005); Maia et al. (2015); Aptroot (2021);
Brazil 13,950 4,310 2,300 2,013
SpeciesLink (2021)
Australia 12,870 4,003 4,000 1,869 May et al. (2004); McCarthy (2020)
Canada 12,800 2,262 4,500 1,829 Perlmutter & Weakley (2018); Bates et al. (2018); Esslinger (2019)
Venezuela 8,300 1,801 — 1,392 Marcano et al. (1996); Itturiaga (2000); Hernández-M. (2021)
Guzmán (1998a, b); Cifuentes (2008); Aguirre-Acosta et al.
Mexico 7,632 2,722 2,900 1,212
(2014); Herrera-Campos et al. (2014); Bates et al. (2018)
Vasco-Palacios & Franco-Molano (2013); Gaya et al. (2021);
Colombia 7,241 2,559 1,239 1,210
Lücking et al. (2021b); this paper
Hywel-Jones & Boonpratuang (2001); Chandrasrikul et al. (2011);
Thailand 7,300 1,292 2,100 1,278
Buaruang et al. (2017); Hyde et al. (2018); Index Fungorum (2021)
Malaysia 3,804 — — 689 Chua et al. (2012)
Costa
3,500 1,740 2,000 743 Halling & Mueller (2005); Mueller et al. (2007)
Rica
Piepenbring (2004); Rodriguez de Flakus et al. (2016); Melgarejo-
Bolivia 2,234 1,353 500 370
Estrada et al. (2020a, b)
Panama 1,807 325 300 370 Piepenbring (2007)
comparable to that of a well-studied temperate country, (see Chapter 6), this would result in approximately 35,000
such as the United States. Colombia encompasses an area species of non-lichenised fungi. Since about 5,000 species
of 1.143 million km2, compared to 9.834 million km2 for of non-lichenised fungi are known from Colombia (Gaya
the United States. The latter has 6,578 fungal species per et al., 2021), nearly 30,000 still need to be catalogued.
log10(km2), so with log10(1,143,000) = 6.058, one would However, the actual number could be even higher because
expect 6,578 x 6.058 = 39,850 species for Colombia tropical ecosystems on average support higher small-scale
(Table 3). This figure is about 5.5 times higher than the species richness than temperate areas, which means
currently known number. Given that lichen-forming fungi that patterns found in the United States or other largely
have been estimated at nearly 5,000 species for Colombia temperate countries cannot be directly transferred.
TABLE 3. Various predictions of the actual species richness of the globally known fungi as a basis, the estimate would range
Colombian funga using different approaches. between 105,600 and 182,400 species, still much higher
than the estimate based on log10(area) extrapolation (Table
Prediction Approach 3). Such estimates may seem unrealistic, but figures in the
same order of magnitude, namely (100,000–)200,000(–
Adjusted according to a proportion of well-
27,430 260,000) species of fungi, have been proposed for Mexico
studied class Lecanoromycetes
(Guzmán 1998a, b; Aguirre-Acosta et al., 2014).
Species versus log10(area) relationship The fungus:plant species ratio is another commonly
39,850
using the USA as a benchmark used measure for predictive species richness estimates
(Martin, 1951; Hawksworth, 1991; Hawksworth & Lücking,
Colombian proportion of global estimated
2017). Empirical data for this measure vary widely, ranging
richness, using known fungi as a benchmark
105,600 from 1.8:1 to 19.1:1, depending on the ecosystem and
(4.8%) and 2.2–3.8 million estimated
–182,400 sampling methods. Thus, while plants within a given area
fungal species globally (Hawksworth &
can be sampled quite accurately, fungal sampling restricted
Lücking, 2017)
to macroscopically visible fungi and within a limited time
156,900 frame will naturally and often grossly underestimate fungal
Fungi:plant ratio between 6:1 and 10:1
–261,500 diversity, as evidenced by a study performed in Panama
(Piepenbring et al., 2012). The ratio also depends on the
Colombian proportion of global estimated
group of fungi. For example, rust fungi (Pucciniomycetes)
richness, using known plants as a
165,000 show lower empirical fungus:plant species ratios between
benchmark (7.5%) and 2.2–3.8 million
–285,000 1:10 and 1:100, with more realistic ratios ranging between
estimated fungal species globally
1:4 and 1:20, resulting in estimates for the diversity of
(Hawksworth & Lücking, 2017)
rust fungi alone in Colombia of between 2,500 and nearly
Colombian proportion of global estimated 13,000 species (Salazar-Yepes & Alves de Carvalho Júnior,
richness, using known amphibia as a 2013). However, these approaches have largely not been
220,000 tested using molecular approaches, both in terms of taxon
benchmark (10.0%) and 2.2–3.8 million
–380,000 concepts in rust fungi and the detection of species not
estimated fungal species globally
(Hawksworth & Lücking, 2017) producing spore-bearing structures. In addition, fungus:plant
species ratios vary with latitude, higher ratios in temperate
areas going along with substantially lower plant diversity,
which requires a downwards correction when extrapolating
A similar prediction is achieved when considering the globally (Tedersoo et al., 2014). Estimates may be further
phylogenetic diversity of fungi of Colombia at the class level misguided by outdated numbers for plant diversity, such
(Figure 3). As outlined above, the largest overrepresented as the earlier - reported exaggerated figure of 51,220 plant
class is Lecanoromycetes (30.3% in Colombia versus 7.9% species for Colombia (see above). Studies that include more
globally), which includes most of the lichenised fungi, a sophisticated sampling methods, such as long-term plots
group particularly well-studied in Colombia. If we assume and/or metabarcoding, show overall fungus:plant species
that the global proportion of Lecanoromycetes is relatively ratios of between 6:1 and 10:1. However, such studies
constant across terrestrial ecosystems dominated by woody have been mostly restricted to temperate ecosystems
plants, then Colombia would be expected to harbour at least (Hawksworth & Lücking, 2017). Given the number of 26,150
27,430 species of fungi to render the current number of plant species currently known from Colombia, applying the
2,167 Lecanoromycetes at a proportion of 7.9%. fungus:plant species ratio approach would thus result in
Another approach is to look at proportional richness 156,900–261,500 species of fungi for Colombia, a figure
per organism group in Colombia relative to the known comparable to that derived from the proportional richness
global diversity of the group. The observed values range approach (Table 3).
between 4.3% for insects and 20.8% for birds, with others Although a reliable estimate of fungal diversity in
in between, e.g., 7.5% for plants, 8.2% for mammals, and Colombia remains speculative, we can safely assume that
10.0% for amphibia (Table 1). Relying on well-studied groups the predicted figure is perhaps an order of magnitude higher
and taking into account how species in particular groups can than the currently number of known species. Apart from the
successfully disperse, that is, how sizable average species sheer size of that number, the more daunting perspective is
ranges probably are, for fungi, one could assume values the effort required to document this largely unknown diversity,
similar to those of plants or amphibia (or butterflies), that requiring a large task force of experts in the different fungal
is, 7.5%–10.0% (–19.4%). The global estimate of between groups and consequent application of modern methods of
2.2 and 3.8 million species (Hawksworth & Lücking, 2017) biodiversity research. Unfortunately, molecular approaches
would give an estimate of (165,000–)220,000–285,000(– to catalogue fungi are still limited in Colombia. We performed
380,000) species for Colombia when applying the 7.5%– a structured search in the NCBI GenBank [https://www.ncbi.
10.0% range (the 19.4% range probably being too high). Even nlm.nih.gov/genbank], using the string <Fungi[organism]
taking the currently observed 4.8% for known Colombian vs AND (5.8S[title] OR ITS1[title] OR ITS2[title] OR “internal
A B
FIGURE 4. Number of NCBI GenBank accessions obtained using the fungal ITS barcoding marker (A) and number of biosamples from
metabarcoding studies using various markers (B), adjusted for area size [relative to log10(area)] for countries across the Americas.
transcribed spacer”[title]) AND country>, for all countries a global context, the inventory of fungi of Colombia is still
in the Americas, to assess how many accessions of the at its onset in terms of expected numbers. Although a good
fungal ITS barcoding marker exist for each country, and then balance of taxonomic groups has been studied to date in
divided the number by log10(area), to filter the area effect. Colombia, including a wide array of macro- and microfungi
The United States ranks two orders of magnitude ahead of and fungi representing different lifestyles, the development
Colombia, whereas Canada, Mexico, Costa Rica, Ecuador, of mycology is not at a level comparable to that in temperate
Brazil, and Argentina rank one order of magnitude ahead countries, in which non-professional mycology also plays a
(Figure 4A). We further queried the NCBI Sequence Read critical role in performing inventory work. We believe that
Archive [https://www.ncbi.nlm.nih.gov/sra] using the country mycology in Colombia can only develop further if the focus
name and recording the number of resulting biosamples, also is not only on applied approaches, but more resources are
dividing them by log10(area) to obtain an estimate of existing provided for basic scientific studies, allowing the adoption of
metabarcoding studies. Again, the United States ranges modern technologies, such as molecular (meta-)barcoding
two orders of magnitude ahead of Colombia and most other at a broader scale. In parallel, fostering taxonomic specialist
countries except Canada (Figure 4B). groups for the various taxa is necessary, particularly for
Considering that it took about 200 years to arrive at a list taxa that remain grossly understudied. The formation of
of just 7,241 species of fungi known from Colombia using associations, such as the Asociación Colombiana de Micología
mostly traditional inventory methods, efforts have to be (ASCOLMI) and the Grupo Colombiano de Liquenología (GCOL)
increased by about two orders of magnitude to catalogue an can be considered an excellent initiative to achieve this goal.
additional at least 30,000 species within a reasonable time
frame of 15–20 years. For this to happen, there needs to
be a paradigm shift in the political and societal support for Acknowledgements
this kind of work in Colombia, and across Latin America and Paul Kirk is warmly thanked for putting data from Index Fungorum
other tropical regions in general. and Species Fungorum at our disposal and for assistance with
obtaining data on the checklist of fungi from China. Kevin Hyde,
CONCLUSIONS Gareth Jones, and Gary Perlmutter assisted with obtaining
references regarding fungal diversity in Thailand, Malaysia, and
The first assembly of a critical checklist of the Colombian North America. Meike Piepenbring and Mauricio Salazar-Yepes
funga is a milestone in the state of knowledge of fungi in provided thoughtful reviews that helped to improve this chapter,
the country. However, compared to other organisms and in and Rafael Felipe de Almeida assisted in linguistic editing.
Gaya E, Vasco-Palacios AM, Vargas-Estupiñan N, Lücking R, Carretero Institute of Microbiology, Chinese Academy of Sciences (2018) Checklist
J, Sanjuan T, Moncada B, Allkin B, Bolaños-Rojas AC, Castellanos- of Fungi in China. Available from: http://124.16.146.175:8080/
Castro C, Coca LF, Corrales A, Cossu T, Davis L, dSouza J, Dufat A, checklist/checklist.html
Franco-Molano, AE, Garcia F, Gómez-Montoya N, González-Cuellar FE, Iturriaga T (2000) Estado Actual del Conocimiento de la Micobiota en
Hammond D, Herrera A, Jaramillo-Ciro MM, Lasso-Benavides C, Mira Venezuela (No. 2). Caracas: Ministerio del Ambiente y de los Recursos
MP, Morley J, Motato-Vásquez V, Niño-Fernandez Y, Ortiz-Moreno ML, Naturales Renovables.
Peña-Cañón ER, Ramirez-Castrillón M, Rojas T, Ruff J, Simijaca D, James TY, Stajich JE, Hittinger CT, Rokas A (2020) Toward a fully resolved
Sipman HJM, Soto-Medina E, Torres G, Torres-Andrade PA, Ulian T, fungal tree of life. Annual Review of Microbiology, 74, 291-313. doi:
White K, Diazgranados M (2021) ColFungi: Colombian Resources for 10.1146/annurev-micro-022020-051835
Fungi Made Accessible. Richmond: Royal Botanic Gardens, Kew. Katumoto K (2009) List of Fungi Recorded in Japan. Tokyo: The Kanto Branch
Green TA, Brabyn L, Beard C, Sancho LG (2012) Extremely low lichen of the Mycological Society of Japan.
growth rates in Taylor Valley, Dry Valleys, continental Antarctica. Polar Kothe E (1999) Mating types and pheromone recognition in the
Biology, 35(4), 535-541. doi: 10.1007/s00300-011-1098-7 homobasidiomycete Schizophyllum commune. Fungal Genetics and
Guzmán GN (1998a) Análisis cualitativo y cuantitativo de la diversidad de Biology, 27(2-3), 146-152.
los hongos en México (Ensayo sobre el inventario fúngico del país). In: Legon NW, Henrici A (2005) Checklist of the British and Irish Basidiomycota.
Halffter G (ed). La Diversidad Biológica de Iberoamérica II. Pp. 111-175. Richmond: Royal Botanic Gardens, Kew. Available at: http://www.
[Acta Zoológica Mexicana, Nueva Serie Vol. Especial]. Xalapa: CYTED basidiochecklist.info
e Instituto de Ecología. Lewinsohn TM, Prado PI (2005) How many species are there in Brazil?
Guzmán GN (1998b) Inventorying the fungi of Mexico. Biodiversity and Conservation Biology, 19(3), 619-624.
Conservation, 7, 369-384. Lücking R, Aime MC, Robberts B, Miller AN, Aoki T, Ariyawansa HA, Cardinali
Guzmán GN, Gómez-Pompa A, Allen MF, Fedick SL, Jiménez-Osorio JJ G, Crous PW, Druzhinina IS, Geiser DM, Hawksworth DL, Hyde KD,
(2003) Fungi in the Maya Culture: Past, Present and Future. In: Fedick Irinyi L, Jeewon R, Johnston PR, Kirk PM, Malosso E, May TW, Meyer
S, Allen M, Jiménez-Osornio J, Gomez-Pompa A (eds). The Lowland W, Nilsson HR, Öpik M, Robert V, Stadler M, Thines M, Vu D, Yurkov
Maya Area. Three Millennia at the Human-Wildland Interface. New York: AM, Zhang N, Schoch CL (2021a) Fungal taxonomy and sequence-based
Food Products Press. Pp. 315-325. nomenclature. Nature Microbiology, 6, 540-548. doi: 10.1038/s41564-
Halling RE, Mueller GM (2005) Common Mushrooms of the Talamanca 021-00888-x
Mountains, Costa Rica. Bronk: New York Botanical Garden. Lücking R, Moncada B, Soto-Medina E, Simijaca D, Sipman HJ (2021b).
Hawksworth DL (1991) The fungal dimension of biodiversity: magnitude, Actualización nomenclatural y taxonómica del Catálogo de Líquenes
significance, and conservation. Mycological Research, 95(6), 641-655. de Colombia. Revista de la Academia Colombiana de Ciencias Exactas,
Hawksworth DL (2001) The magnitude of fungal diversity: the 1.5 million Físicas y Naturales, 45(174), 147-189. doi: 10.18257/raccefyn.1266
species estimate revisited. Mycological Research, 105(12), 1422- Maia LC, Carvalho AA, Cavalcanti LH, Gugliotta AM, Drechsler-Santos
1432. ER, Santiago ALM, Cáceres ME, Gibertoni T, Aptroot A, Giachini AJ,
Hawksworth DL, Lücking R (2017) Fungal diversity revisited: 2.2 to 3.8 Soares AMS, Silva ACG, Magnago AC, Goto BT, Lira CRS, Motoya CAS,
million species. Microbiology Spectrum, 5(4), FUNK-0052-2016. doi: Pires-Zotarelli CLA, daSilva DKA, Soares DJ, Rezende DHC, Luz EDM,
10.1128/microbiolspec.FUNK-0052-2016 Gumboski EL, Wartchow F, Karstedt F, Freire FM, Coutinho FP, Melo
Hernández-M JA (2021) A new look at Venezuelan lichen diversity. GSN, Sotão HM, Baseia IG, Pereira J, Oliveira JJS, Souza JF, Bezerra
International Association for Lichenology 9th Symposium (IAL 9). JL, Araujo LS, Pfenning H, Gusmão LFP, Neves MA, Capelari M, Jaeger
Brazil (online). IAL9 Program and Abstract Book, 77. MCW, Palacios MP, Menolli N, Medeiros PS, Friedrich RCS, Chikowski R,
Herrera-Campos MA, Lücking R, Pérez-Pérez RE, Miranda-González R, Pires RM, Melo RF, Silveira RMB, Urrea-Valencia S, Costez VG, Silva VF
Sánchez N, Barcenas-Peña A, Carrizosa A, Zambrano A, Ryan BD, Nash (2015) Diversity of Brazilian Fungi. Rodriguesia 66(4), 1033-1045. doi:
III TH (2014) Biodiversidad de líquenes en México. Revista Mexicana 10.1590/2175-7860201566407
de Biodiversidad, 85, S82-S99. doi: 10.7550/rmb.37003 Maldonado-Ocampo JA, Vari RP, Usma JS (2008) Checklist of the freshwater
Hosoya T (2006) History and the current status of fungal inventory and fishes of Colombia. Biota Colombiana, 9(2), 143-237.
databasing in Japan. In: Tomida I (ed). Proceedings of the 7th and 8th Mammal Diversity Database (2021) Mammal Diversity Database (Version
symposia on Collection Building and Natural History Studies in Asia and 1.5) Zenodo. Available at: http://doi.org/10.5281/zenodo.4139818
the Pacific Rim. [National Museum Monographs 34]. Tokyo: National Manoharachary C, Nagaraju D (2016) New additions to the fungi of India.
Science Museum. Pp. 51-61. Indian Phytopathology, 69, 93-96.
Hyde KD, Norphanphoun C, Chen J, Dissanayake AJ, Doilom M, Hongsanan Marcano V, Morales–Méndez A, Sipman H, Calderon L (1996) A first
S, Jayawardena RS, Jeewon R, Perera RH, Thongbai B, Wanasinghe checklist of the lichen-forming fungi of the Venezuelan Andes. Tropical
DN, Wisitrassameewong K, Tibpromma S, Stadler M (2018) Thailand’s Bryology, 12, 193-235.
amazing diversity: up to 96% of fungi in northern Thailand may be Martin GW (1951) The numbers of fungi. Proceedings of the Iowa Academy of
novel. Fungal Diversity, 93, 215-239. doi: 10.1007/s13225-018- Science, 58(1), 175-178.
0415-7 May TW, Milne J, Wood AE, Shingles S, Jones RH, Neish P (2004) Interactive
Hyde KD, Xu JC, Lumyong S, Rapior S, Jeewon R, Lumyong S, Niego AGT, Catalogue of Australian Fungi, Version 3.0. Australian Biological
Abeywickrama PD, Aluthmuhandiram JVS, Brahamanage RS, Brooks Resources Study, Canberra/Royal Botanic Gardens Melbourne.
S, Chaiyasen A, Chethana KWT, Chomnunti P, Chepkirui C, Chuankid McCarthy PM (2020). Checklist of the Lichens of Australia and its Island
B, de Silva NI, Doilom M, Faulds C, Gentekaki E, Gopalan V, Kakumyan Territories. Australian Biological Resources Study, Canberra. Version
P, Harishchandra D, Hemachandran H, Hongsanan S, Karunarathna 1 March 2020. Available at: http://www.anbg.gov.au/abrs/lichenlist/
A, Karunarathna SC, Khan S, Kumla J, Jayawardena RS, Liu JK, Liu introduction.html
N, Luangharn T, Macabeo APG, Marasinghe DS, Meeks D, Mortimer Medardi G (2006) Ascomiceti d’Italia. A.M.B. Centro Studi Micologici,
PE, Mueller P, Nadir S, Nataraja KN, Nontachaiyapoom S, O’Brien M, Associazione Micologica Bresadola, Trento.
Penkhrue W, Phukhamsakda C, Ramanan US, Rathnayaka AR, Sadaba Melgarejo-Estrada E, Suarez ME, Rocabado D, Maillard O, Lechner BE
RB, Sandargo B, Samarakoon BC, Tennakoon DS, Siva R, Sriprom (2020a) Checklist of Bolivian Agaricales: 1. Species with dark and pink
W, Suryanarayanan TS, Sujarit K, Suwannarach N, Suwunwong T, spore prints. Mycotaxon, 134(4), 739. doi: 10.5248/134.739
Thongbai B, Thongklang N, Wei D, Wijesinghe SN, Winiski J, Yan J, Melgarejo-Estrada E, Rocabado D, Suárez ME, Maillard O, Lechner BE
Yasanthika E, Stadler M (2019) The amazing potential of fungi, 50 (2020b) Checklist of Bolivian Agaricales. 2: Species with white or pale
ways we can exploit fungi industrially. Fungal Diversity, 97, 1-136. doi: spore prints. Mycotaxon, 135(1), 233. doi: 10.5248/135.233
10.1007/s13225-019-00430-9 Mejia LS, Acero P (2002) Libro Rojo de Peces Marinos de Colombia. Bogotá:
Hywel-Jones NL, Boonpratuang T (2001) Updated Preliminary Checklist of INVEMAR, Instituto de Ciencias Naturales, Universidad Nacional de
Fungi Recorded from Thailand. BIOTEC Yothi-Mycology Laboratory. Colombia, Ministerio de Medio Ambiente.
Index Fungorum (2021) Index Fungorum. Available from: http://www. Mueller GM, Foster M, Bills G (2004) Biodiversity of Fungi. Inventory and
indexfungorum.org/names/Names.asp Monitoring Methods. Burlington, MA: Academic Press (Elsevier)
Mueller GM, Schmit JP, Leacock PR, Buyck B, Cifuentes J, Desjardin DE, Schmitt CL, Tatum ML (2008) The Malheur National Forest: Location of the
Halling RE, Hjortstam K, Iturriaga T, Larsson KH, Lodge DJ (2007) World’s Largest Living Organism (the Humongous Fungus). Washington,
Global diversity and distribution of macrofungi. Biodiversity and DC: Forest Service, US Department of Agriculture.
Conservation, 16(1), 37-48. doi: 10.1007/s10531-006-9108-8 Singh KP, Sinha GP (2010) Indian Lichens: An Annotated Checklist. Kolkata:
Do Nascimento C, Herrera-Collazos EE, Maldonado-Ocampo JA (2018) Botanical Survey of India.
Lista de Especies de Peces de Agua Dulce de Colombia / Checklist Sinha GP, Nayaka S, Joseph S (2018) Additions to the checklist of Indian
of the Freshwater Fishes of Colombia. V2.10. Bogotá: Asociación lichens after 2010. Cryptogam Biodiversity and Assessment, Special
Colombiana de Ictiólgos. Dataset/Checklist. Available at: http://doi. Volume, 2018, 197-206. doi: 10.21756/cab.esp16
org/10.15472/numrso Smith CW, Aptroot A, Coppins BJ, Fletcher A, Gilbert OL, James PW,
Nelson JS, Grande TC, Wilson MV (2016) Fishes of the World. Hoboken, Wolseley PA (2009) The Lichens of Great Britain and Ireland. London:
NJ: John Wiley & Sons. British Lichen Society.
Newton AF, Gutiérrez-Chacón C, Chandler DS (2005) Checklist of the Smith ML, Bruhn JM, Anderson JB (1992) The fungus Armillaria bulbosa
Staphylinidae (Coleoptera) of Colombia. Biota Colombiana, 6(1), 1-72. is among the largest and oldest living organisms. Nature, 356(6368),
Nguyen NH, Song Z, Bates ST, Branco S, Tedersoo L, Menke J, Schilling 428-431. doi: 10.1038/356428a0
JS, Kennedy PG (2016) FUNGuild: an open annotation tool for parsing Species Fungorum (2021) Species Fungorum. Available at: http://www.
fungal community datasets by ecological guild. Fungal Ecology, 20, speciesfungorum.org
241-248. doi: 10.1016/j.funeco.2015.06.006 SpeciesLink (2021) SpeciesLink. Available at: https://specieslink.net
Nimis PL (2016) The Lichens of Italy. A Second Annotated Catalogue. Stork NE (2018) How many species of insects and other terrestrial
Trieste: EUT Edizioni Università di Trieste. arthropods are there on earth? Annual Review of Entomology, 63, 31-
Nimis PL, Scheidegger C, Wolseley PA (2002) Monitoring with Lichens – 45. doi: 10.1146/annurev-ento-020117-043348
Monitoring Lichens. Dordrecht: Springer Tedersoo L, Bahram M, Põlme S, Kõljalg U, Yorou NS, Wijesundera R,
Ohmura Y, Kashiwadani H (2018) Checklist of Lichens and Allied Fungi of Ruiz LV, Vasco-Palacios AM, Thu PQ, Suija A, Smith ME, Sharp C,
Japan. Tokyo: National Museum of Nature and Science. Saluveer E, Saitta A, Rosas M, Riit T, Ratkowsky D, Pritsch K, Põldmaa
Onofri S, Bernicchia A, Filipello-Marchisio V, Padovan F, Perini C, Ripa C, K, Piepenbring M, Phosri C, Peterson M, Parts K, Pärtel K, Otsing E,
Savino E, Venturella G, Vizzini A, Zotti M, Zucconi L (2013) Checklist Nouhra E, Njouonkou AL, Nilsson RH, Morgado LN, Mayor J, May TW,
of the Macrobasidiomycetes of Italy / Checklist dei Macrobasidiomiceti Majuakim L, Lodge DJ, Lee SS, Larsson KH, Kohout P, Hosaka K,
d’Italia. Available at: http://dryades.units.it/macrobasidiomiceti Hiiesalu K, Henkel TW, Harend H, Guo L, Greslebin A, Grelet G, Geml
Perafán C, Sabogal A, Moreno-González JA, García-Rincón A, Luna- J, Gates G, Dunstan W, Dunk C, Drenkhan R, Dearnaley J, De Kesel
Sarmiento D, Romero-Ortíz C, Flórez E (2013) Diagnóstico del A, Dang T, Chen X, Buegger F, Brearley FQ, Bonito G, Anslan S, Abell
S, Abarenkov K (2014) Global diversity and geography of soil fungi.
estado actual de la fauna de arácnidos y de su gestión en Colombia.
Science, 346(6213), 1078. doi: 10.1126/science.1256688
Simposio 8: Aracnología. In: Rueda-Ramírez D, Torrado-León E,
The Plant List (2021) The Plant List. Available at: http://www.theplantlist.
Becerra EH (eds). Memorias Congreso Colombiano de Entomología.
org
40 Congreso Socolen. Socolen, Bogota, DC: Sociedad Colombiana de
Thines M, Aoki T, Crous PW, Hyde KD, Lücking R, Malosso E, May TW,
Entomología. Pp. 275-335.
Miller AN, Redhead SA, Yurkov AM, Hawksworth DL (2020) Setting
Pérez-Gutiérrez LA, Palacino-Rodríguez F (2011). Updated checklist of
scientific names at all taxonomic ranks in italics facilitates their quick
the Odonata known from Colombia. Odonatologica, 40(3), 1-203.
recognition in scientific papers. IMA Fungus, 11(1), 1-5. doi: 10.1186/
Perlmutter GB, Weakley AS (2018) USDA Plants Database Lichen Update,
s43008-020-00048-6
Phase II. Greensboro, NC: NRCS – East National Technology Support
Uetz P, Freed P, Aguilar R, Hošek J (2021) The Reptile Database. Available
Center
at: http://www.reptile-database.org
Piepenbring M (2004) 3.2.1. Fungi. In: Ibisch PL, Mérida G (eds).
U’Ren JM, Lutzoni F, Miadlikowska J, Laetsch AD, Arnold AE (2012) Host
Biodiversity: the Richness of Bolivia, State of Knowledge and
and geographic structure of endophytic and endolichenic fungi at a
Conservation. Santa Cruz de la Sierra: Editorial FAN. Pp. 89-94. continental scale. American Journal of Botany, 99(5), 898-914. doi:
Piepenbring M (2007) Inventorying the fungi of Panama. Biodiversity and 10.3732/ajb.1100459
Conservation, 16(1), 73-84. Vasco-Palacios AM, Franco-Molano AE (2013) Diversity of Colombian
Piepenbring M (2015) Introduction to Mycology in the Tropics [Introducción macrofungi (Ascomycota–Basidiomycota). Mycotaxon, 121(1), 100-158.
a la Micología en los Trópicos]. St. Paul: APS Press. Vélez, A. (2008). Checklist of Colombian cockroaches (Dictyoptera,
Piepenbring M, Hofmann TA, Unterseher M, Kost G (2012) Species Blattaria). Biota Colombiana, 9(1), 21-38.
richness of plants and fungi in western Panama: towards a fungal Ventura M, Canchaya C, Tauch A, Chandra G, Fitzgerald GF, Chater KF,
inventory in the tropics. Biodiversity and Conservation, 21(9), 2181- van Sinderen D (2007) Genomics of Actinobacteria: tracing the
2193. doi: 10.1007/s10531-011-0213-y evolutionary history of an ancient phylum. Microbiology and Molecular
Piepenbring M, Lotz-Winter H, Hofmann TA (2018) Incentives and Biology Reviews, 71(3), 495-548.
challenges for mycologists in the tropics. Biosystematics and Ecology Venturella G, Altobelli E, Bernicchia A, Di Piazza S, Donnini D, Gargano ML,
Series, 34, 481-514. Gorjòn SP, Granito VM, Lantieri A, Lunghini D, Montemartini A (2011)
Rhodes CJ (2014) Mycoremediation (bioremediation with fungi) – growing Fungal biodiversity and in situ conservation in Italy. Plant Biosystems-
mushrooms to clean the earth. Chemical Speciation & Bioavailability, An International Journal Dealing with all Aspects of Plant Biology, 145(4),
26(3), 196-198. doi: 10.3184/095422914X14047407349335 950-957.
Rodriguez RJ, White Jr JF, Arnold AE, Redman ARA (2009) Fungal Watkinson SC, Boddy L, Money NP (2016) The Fungi. Third Edition. Oxford,
endophytes: diversity and functional roles. New Phytologist, 182(2), UK: Academic Press (Elsevier).
314-330. doi: 10.1111/j.1469-8137.2009.02773.x Wang K, Zhao M, Su J, Yang L, Deng H, Wang Y, Wu H, Li Y, Wu H, Wei X, Wei
Rodriguez de Flakus P, Kukwa M, Etayo J, Lücking R, Meneses RI, Plata T, Cai L, Yao Y (2020) The use of Checklist of Fungi in China database
ER, Stanton D, Truong C, Vargas R, Flakus A (2016) Preliminary in the red list assessment of macrofungi in China. Biodiversity Science,
Catalogue of Lichens and Lichenicolous Fungi from Bolivia. Available 28(1), 74-98. doi: 10.17520/biods.2019163
at: https://bio.botany.pl/lichens-bolivia/en,strona,catalogue,5.htm Wei JC (2021) The Enumeration of Lichenized Fungi in China. Beijing: China
Roux C (2012) Liste des lichens et champignons lichénicoles de France. Forestry Publishing House.
Bulletin de la Société Linnéenne de Provence, N Spécial 16, 1-220. Wirth V, Hauck M, Brackel WV, Cezanne R, de Bruyn U, Dürhammer O,
Salazar-Yepes M, Alves de Carvalho Júnior A (2013) Uredinales (rust Eichler M, Gnüchtel A, Litterski B, Otte V, Schiefelbein U (2011)
fungi) biota of the Parque Nacional do Itatiaia, Brazil: an analysis of Checklist of Lichens and Lichenicolous Fungi in Germany, Version 2.
composition, species diversity and altitudinal distribution. Caldasia Göttingen: Georg August University.
35(1), 165-176. Zhang ZQ (2013) Animal biodiversity: An update of classification
Sarbhoy AK, Agarwal DK, Varshney JL (1996) Fungi of India 1982-1992. and diversity in 2013. Zootaxa, 3703(1), 5-11. doi: 10.11646/
New Delhi: CBS Publishers and Distributors. zootaxa.3703.1.3
Panaeolus semiovatus
[Ana Esperanza Franco-Molano]
58 CATALOGUE OF FUNGI OF COLOMBIA
Chapter 4
Diversity of Basidiomycota in Colombia
Ana Esperanza Franco-Molano 1*, Melissa Palacio 1,2 & Nataly Gómez-Montoya 1
1
Laboratorio de Taxonomía y Ecología de Hongos -TEHO, Instituto de Biología, Facultad de Ciencias Exactas y Naturales, Universidad de Antioquia - UdeA,
Calle 70 No. 52-21, Medellín, Antioquia, Colombia.
2
Laboratório de Micologia, Departamento de Botânica, Instituto de Biociências, Universidade Federal do Rio Grande do Sul, Av. Bento Gonçalves, 9500,
Prédio 43.433, Campus do Vale, Agronomia, 91501-970, Porto Alegre, Rio Grande do Sul, Brasil.
ABSTRACT
Colombia is one of the most biodiverse countries on the planet due to its geographical location and its great diversity in
ecosystems and climates, which give rise to the high richness and diversity of its funga. Its funga will probably prove to
be one of the most diverse in South America. In the present study, we analysed the dataset for Basidiomycota provided
by the ColFungi project and the Herbarium of the University of Antioquia (HUA). Statistical analyses show a high diversity
of Basidiomycota, especially for Agaricomycetes in oak and lowland forests. Also, evidence of the lack of studies for some
departments of Colombia, such as the Guaviare, is provided. Regarding the conservation of fungi, it is observed that only a
few studies have been carried out for Basidiomycota macrofungi in Colombia, since only 22 species have been assessed to
date for their extinction threat.
RESUMEN
Colombia es uno de los países más biodiversos en el planeta debido a su ubicación geográfica y su gran diversidad de
ecosistemas y climas, por lo que promete una alta riqueza y diversidad de hongos, y probablemente se constituya en uno de
los países más diversos de Sudamérica respecto a su funga. En el presente trabajo se hace la recopilación de datos para
Basidiomycota, suministrados por el KEW y el Herbario de la Universidad de Antioquia (HUA). Análisis estadísticos demuestran
alta diversidad en cuanto a Agaricomycetes, principalmente para los ecosistemas de roble y tierras bajas, así como también
la falta de estudios para algunos departamentos de Colombia, como Guaviare. En cuanto a la conservación, se observa que,
en el país, se han realizado pocos estudios sobre macrohongos de Basidiomycota, ya que hasta el momento solo 22 especies
han sido evaluadas.
He et al., 2019). Currently, Basidiomycota comprises of 341 million years (He et al., 2019). Tremellomycetes
four subphyla: Pucciniomycotina with nine classes, include saprotrophic, animal and fungi parasites, or can be
Ustilaginomycotina with four classes, Wallemiomycotina commensals with fungi and lichens (Millanes et al., 2011;
with a single class, and Agaricomycotina with three classes Weiß et al., 2014). The class includes asexual yeasts and
(He et al., 2019). fungi with conspicuous and gelatinous basidiomas such as
Pucciniomycotina has more than 8,000 described Tremella (Weiß et al., 2014). Finally, the Agaricomycetes
species and represents a group that diverged at least have an evolutionary history dating to at least 298
406 million years ago (He et al., 2019). This subphylum is million years ago and include 30,143 described species
characterised by predominant asexual or anamorphic stages distributed in 19 orders (He et al., 2019). Recent studies
in most lineages. In fact, only these stages are known in suggest that resupinate basidiomas that are fully attached
some classes, such as Tritirachiomycetes. Pucciniomycotina to the substrate provide the most likely ancestral trait for
species play different roles in ecosystems, but they are Agaricomycetes (Sánchez-García et al., 2020). Furthermore,
better known as phytopathogens and parasites of animals the pileate-stipitate morphology (e.g., agaricoid and boletoid)
or other fungi (Aime et al., 2014; Malysheva et al., 2021). is correlated with high rates of diversification, suggesting
Ustilaginomycotina, with more than 1,800 described species, that this morphology represents a key character in the
has a minimum age of divergence of 430 million years (He et success of Agaricomycetes (Varga et al., 2019; Sánchez-
al., 2019). This subphylum comprises plant parasitic fungi, García et al., 2020).
such as Jamesdicksonia irregularis, and other economically Ecologically speaking, species of Basidiomycota are an
important fungi, such as Ustilago maydis, a model organism essential part of natural ecosystems, with many recycling or
used to study the interaction between plants and their degrading organic matter and shaping communities of plants
specific parasites (Begerow et al., 2014). Wallemiomycotina through parasitic and symbiotic interactions. Saprotrophic
is a recently proposed group of unicellular fungi (Zhao et al., fungi are those that degrade organic matter such as dead wood
2017), that includes xerophilic, osmophilic, and halophilic and litter (Posada et al., 2012). They play an essential role
fungi (Zajc & Gunde-Cimerman, 2018). in forest ecosystems as they are one of the largest recycling
Within Basidiomycota, Agaricomycotina has the largest groups (Robledo & Urcelay, 2009). On the other hand, many
number of described species, around 30,800, and its species are essential in conserving ecosystems by forming
evolutionary history dates to at least 406 million years a mycorrhizal symbiosis with trees and shrubs (Tedersoo et
ago (He et al., 2019). Agaricomycotina are divided into al., 2018). Some species can associate with termites and
three classes mainly characterised by the presence of leafcutter ants, generating a complex symbiosis in which
macroscopic structures called basidiomas, which are insects feed on the cultivated fungi. Other species within this
used for sexual reproduction (He et al., 2019; Kirk et al., phylum are parasitic and can cause mild or potentially lethal
2008). Basidiomas vary from very small structures, like infections in animals and humans. In the case of plants, rusts
those in Mycena spinosissima, to very large such as those and smuts represent the most critical pathogens (Watkinson
in Termitomyces titanicus. The basidiomas also present et al., 2016). Around 41,270 Basidiomycota species have
different morphological patterns named: agaricoids, been recorded globally, making this phylum the second most
boletoids, polyporoids, resupinates, clavaroids, coralloid, speciose after Ascomycota (He et al., 2019).
gasteroids, and secotioids (Cepero de García et al., 2012; From a biogeographic point of view, some members
Varga et al., 2019). These types of basidiomas have emerged of Basidiomycota show remarkable patterns of origin and
through multiple evolutionary convergences in the history of distribution directly related to their habitat types, being
Basidiomycota (Hibbett & Binder, 2002; Sánchez-García et influenced by geological history, vegetation, whether native
al., 2020). According to He et al., (2019), this subphylum or introduced, climate, and anthropogenic impacts (Halling
currently comprises three classes: Dacrymycetes, & Mueller, 2002). In Colombia, geological events have
Tremellomycetes, and Agaricomycetes. played essential roles in determining distribution. These
Dacrymycetes comprise 146 described species events are well documented, by Halling & Ovrebo (1987),
distributed into two orders, with an estimated minimum Halling (1996), Halling & Mueller (2002), and Mueller et
divergence time of 298 million years ago (He et al., 2019). al., (2006), among others (see Chapter 10). In Colombia,
It is also the sister group of Agaricomycetes (McLaughlin et ectomycorrhizal fungi, which may have diversified later in the
al., 2016; Zamora & Ekman, 2020). This class comprises Jurassic (208 – 146 million years ago), present a limited
species characterised by having sterigmatized or bifurcated distribution that corresponds with the distribution of their
holobasidia (except for Unilacryma unispora), and by the hosts, and are more abundant and diverse at 2,200–
presence of continuous parenthesomes that cover the 3,200 m altitude where most of the oak forests (Quercus
dolipore septa (Zamora & Ekman, 2020). Dacrymycetes humboldtii) are found (Avella-Muñoz & Rangel-Churio, 2014).
includes saprotrophic fungi that produce gelatinous, waxy, Colombian ectomycorrhizal fungi from oak forests show
fleshy, or cartilaginous basidiomas with yellow to orange affinities with northern temperate taxa, migrating from
colouration due to their carotenoid pigments (Zamora & north to south along with those forest communities (Halling,
Ekman, 2020). 1996, Chapter 10). Another important Fagaceae associated
Tremellomycetes comprise nearly 500 accepted species with ectomycorrhizal fungi is Colombobalanus excelsa, a
distributed in five orders, with an evolutionary history species endemic to the Colombian Andes of which there
are eight populations located between 1,800 and 2,200 TABLE 1. List of fungal species categorised by the IUCN for
altitude (Davila et al., 2012). However, very few mycological Colombia. LC- least concern. CR- critically endangered. VU-
explorations have been carried out in this type of forest, and vulnerable. NT- near threatened. DD- data deficient.
few results have been published. Another ectomycorrhizal-
forming tree from which some agarics have been described is IUCN
Population
Pseudomonotes tropenbosii (Dipterocarpaceae), an endemic Species Red List
trend
species, described from the Colombian Amazon at 300 m category
altitude (Londoño et al., 1995). Some non-native plants, Amanita fuligineodisca Decreasing VU
such as Pinus, Eucalyptus, and Cupressus trees, are also able
to form these symbiotic associations. These species were Amanita sepultipes Decreasing VU
introduced into the country in the 1950s and 1960s bringing
their ectomycorrhizal fungi hidden in their roots (Vargas et Austroboletus amazonicus Unknown CR
al., 2019). Nevertheless, these exotic fungi compete with
native fungal species, invading other ecosystems such as Binderoboletus segoi Unknown DD
the oak forest (Vargas et al., 2019). The saprotrophic agarics
Cantharellus guyanensis Unknown LC
registered for Colombia are cosmopolitan or have restricted
distribution giving good biogeographical information (Halling Clavulina kunmudlutsa Unknown DD
& Mueller, 2002; see Chapter 10).
Regarding the conservation of fungi in Colombia, it is Clavulina tepurumenga Unknown DD
essential to note that the IUCN has been working on the
conservation assessment of species of macrofungi only Coprinus comatus Stable LC
in recent years (Dahlberg & Mueller 2011; IUCN, 2021).
Worldwide, 328 Basidiomycota species (IUCN, 2021) have Cortinarius aurantiobrunneus Decreasing NT
been evaluated and placed in the following categories:
Cymatoderma sclerotioides Decreasing NT
Critically threatened (12 species), Endangered (57 species),
and Vulnerable (110 species). The remaining species have Fomitiporia bambusarum Increasing LC
been evaluated as least concern, deficient data, or almost
threatened. Specifically for Colombia, there are few studies Gloiocephala quercetorum Decreasing NT
[e.g., Amanita sepultipes (Vargas-Estupiñan et al., 2020)
and Austroboletus amazonicus (Vasco-Palacios et al., 2020)] Hericium erinaceus Decreasing LC
(Table 1, Chapter 14) on fungal conservation as fungi are
not mentioned in Colombian environmental legislation, Lactifluus hallingii Decreasing VU
and their socio-political interest is not of much relevance Leccinum andinum Decreasing NT
(pers. observ.). The only group of fungi contemplated
within Resolution 213 of 1977 of the National Institute of Lycoperdon perlatum Stable LC
Renewable Natural Resources and Environment (INDERENA)
are lichens, which are mentioned in the resolution as non- Phylloporus fibulatus Decreasing VU
vascular epiphytes (Resolution 213, 1977). The goal of
this chapter is to present the current state of knowledge of Suillus luteus Stable LC
the Basidiomycota in Colombia in terms of their diversity,
Tylopilus bulbosus Decreasing NT
ecology, and conservation.
Tylopilus obscurus Decreasing NT
MATERIALS AND METHODS
Two fungi databases for fungal occurrences in Colombia
were used: the ColFungi database comprising more than
7,000 species, and the Herbarium of the University of
Antioquia (2020) with more than 13,000 records from which RESULTS AND DISCUSSION
only Basidiomycota records were considered. Statistical Currently, there are 2,386 species of Basidiomycota
analyses and graphics were made using R studio version recorded for Colombia (including mushrooms, stinkhorns,
1.3.959 (RStudio Team, 2020) and Microsoft Excel (2020). puffballs, earth stars, shelf fungi, gelatinous fungi, smuts,
The data used for the analyses were divided by ecosystem rusts and yeasts, among others) (Gaya et al., 2021). These
into six groups as follows: oak forest, coniferous forest species are classified within three of the four subphyla
(mainly dominated by Pinus patula and Cupressus lusitanica), recognised by Tedersoo et al., (2018) and He et al., (2019):
mixed forest, Amazon Forest (in the departments of Agaricomycotina (1,642 species), Pucciniomycotina (579
Amazonas and Caquetá), lowland forest (below 1,400 masl, species), and Ustilaginomycotina (84 species) (Figure
excluding previous ecosystems), and other ecosystems (for 1). To date, there are no records for Wallemiomycotina
urban areas or or areas lacking information). in Colombia.
FIGURE 2. Number of records grouped by class. Abundance by class is shown using the HUA database.
FIGURE 3. Richness of Basidiomycota macrofungi and their distribution by departments of Colombia (modified from Gómez-Montoya et al.,
2021). Amazonas: ama; Antioquia: ant; Arauca: ara; Atlántico: atl; Bolivar: bol; Boyacá: by; Caldas: cl; Caquetá: cq; Casanare: cs; Cauca:
cau; Cesar: ce; Chocó: cho; Córdoba: co; Cundinamarca: cun; El Archipiélago de San Andrés, Providencia y Santa Catalina: asps; Guainía:
gn; Guaviare: gv; Huila: hu; La Guajira: gua; Magdalena: ma; Meta: met; Nariño: na; Norte de santander: ns; Parque Nacional Natural Los
Nevados: PNNN.
The high diversity of Agaricomycetes present in these seen in mycological explorations in easily accessible areas
ecosystems corroborates Mueller & Halling (1995), who and ecosystems and/or where mycologists or people with
documented a high level of diversity of ectomycorrhizal fungi mycological training are located. The need to carry out
and a high degree of endemism, between 30–100%, for inventories and/or monitoring is reaffirmed in places such
these ecosystems in Costa Rica and Colombia. Likewise, as the department of Guaviare, which has no records of
López-Quintero et al., (2012) showed great diversity in Basidiomycota, or in the departments of Atlántico, Arauca,
terms of funga for the Amazon lowland forests, where Córdoba, Casanare, Guainía, La Guajira, Sucre, Vichada,
saprotrophic fungi show high diversity. The Archipelago of San Andrés, Providencia, and Santa
The results here presented are an example of the Catalina, with only one to eight records (Gómez-Montoya et
efforts made to know the funga of Colombia, especially al., 2021). It is also important to emphasise the need for
the Basidiomycota. In recent decades, progress has been more studies on other ecosystems currently categorised as
FIGURE 4. Number of records grouped by ecosystems. Abundance by class is shown using the HUA database.
Medina E, Torres G, Torres-Andrade PA, Ulian T, White K, Diazgranados Mueller GM, Halling RE (1995) Evidence for high biodiversity of Agaricales
M (2021) ColFungi: Colombian resources for Fungi Made Accessible. (Fungi) in neotropical montane Quercus forests. In: Churchill SP,
Richmond: Royal Botanic Gardens, Kew. 36p. Balslev H, Forero E (eds). Biodiversity and Conservation of Neotropical
Gentry AH (1978) Floristic knowledge and needs in Pacific Tropical Montane Forests. New York: New York Botanical Garden. Pp. 303-312.
America. Brittonia 30: 134-156. Mueller GM, Halling RE, Carranza J, Mata M, Schmit JP (2006) Saprotrophic
Gómez-Montoya N, Ríos-Sarmiento C, Zora-Vergara B, Benjumea- and ectomycorrhizal macrofungi of Costa Rican oak forests. In:
Aristizabal C, Santa-Santa DJ, Zuluaga-Moreno M, Franco-Molano Kappelle M (ed). Ecology and Conservation of Neotropical Montane Oak
AE (2021) Diversidad de macrohongos (Basidiomycota) de Colombia: Forests. Ecological Studies 185. Berlin: Springer. Pp. 55-68.
listado de especies. Actualidades Biológicas, in press. Posada RH, Madriñan S, Rivera EL (2012) Relationships between the
Halling RE (1996) Boletaceae (Agaricales): latitudinal biodiversity and litter colonisation by saprotrophic and arbuscular mycorrhizal fungi
biological interactions in Costa Rica and Colombia. Revista de Biología with depth in a tropical forest. Fungal Biology, 116(7), 747-755. doi:
Tropical, 44 (Suppl. 4), 11-114. 10.1016/j.funbio.2012.04.003
Halling RE, Mueller GM (2002) Agarics and Boletes of Neotropical Oakwoods. Resolución No. 213 DE 1977. Available at: https://cvc.gov.co/sites/
In: Watling RJ, Frankland JC, Ainsworth AM, Isaac S, Robinson CH default/ files/Sistema_Gestion_de_Calidad/Procesos%20y%20
(eds). Tropical Mycology. London: CABI Publishing. Pp. 1-10. procedimientos%20Vigente/Normatividad_Gnl/Resolucion%20
Halling RE, Ovrebo CL (1987) A new species of Rozites from oak forests 213%20de%201977-Feb-01.pdf
of Colombia, with notes on biogeography. Mycologia, 79(5), 674–678. Robledo G, Urcelay C (2009) Hongos de la madera en árboles nativos del
Hawksworth DL (1991) The fungal dimension of biodiversity: magnitude, centro de Argentina. Cordoba: Ed. Universidad Nacional de Cordoba.
significance, and conservation. Mycological Research, 95(6), 641- 224p.
655. RStudio Team (2020) RStudio: integrated development for R. Available at:
Hawksworth DL, Lücking R (2017) Fungal diversity revisited: 2.2 to 3.8 http://www.rstudio.com/
million species. Microbiology Spectrum, 5(4), FUNK-0052-2016. doi: Sánchez-García M, Ryberg M, Khan FK, Varga T, Nagy LG, Hibbett DS
10.1128/microbiolspec.FUNK-0052-2016 (2020) Fruiting body form, not nutritional mode, is the major driver of
He MQ, Zhao RL, Hyde KD, Begerow D, Kemler M, Yurkov A, McKenzie diversification in mushroom-forming fungi. Proceedings of the National
EH, Raspe O, Kakishima M, Sánchez-Ramírez S, Vellinga EC, Halling Academy of Sciences U S A , 117(51), 32528-32534. doi: 10.1073/
R, Papp V, Zmitrovich IV, Buyck B, Ertz D, Wijayawardene NN, Cui BK, pnas.1922539117
Schoutteten N, Liu XZ, Li TH, Yao YJ, Zhu XY, Liu AQ, Li GJ, Zhang Tedersoo L, Sánchez-Ramírez S, Kõljalg U, Bahram M, Döring M, Schigel D,
MZ, Ling ZL, Cao B, Antonín V, Boekhout T, Barbosa da Silva BD, May T, Ryberg M, Abarenkov K (2018) High-level classification of the
De Crop E, Decock C, Dima B, Dutta AK, Fell JW, Geml J, Ghobad- fungi and a tool for evolutionary ecological analyses. Fungal diversity,
Nejhad M, Giachini AJ, Gibertoni TB, Gorjon SP, Haelewaters D, He 90(1), 135-159. doi: 10.1007/s13225-018-0401-0
SH, Hodkinson BP, Horak E, Hoshino T, Justo A, Lim YW, Menolli N, Universidad de Antioquia (2020) Herbario Universidad de Antioquia (HUA),
Mesic A, Moncalvo JM, Mueller GM, Nagy LG, Nilsson RH, Noordeloos Coleccción de Hongos en línea. Available at: http://www2.udea.edu.
M, Nuytinck J, Orihara T, Ratchadawan C, Rajchenberg M, Silva-Filho co/ herbario/paginas/consultas/consultarEjemplares.iface
A, Sulzbacher MA, Tkalcec Z, Valenzuela R, Verbeken A, Vizzini A, Vargas N, Gonçalves SC, Franco-Molano AE, Restrepo S, Pringle A (2019)
Wartchow F, Wei TZ, Weiß M, Zhao CL, Kirk PM (2019) Notes, outline In Colombia the Eurasian fungus Amanita muscaria is expanding its
and divergence times of Basidiomycota. Fungal Diversity, 99(1), 105- range into native, tropical Quercus humboldtii forests. Mycologia,
367. doi: 10.1007/s13225-019-00435-4 111(5), 758-771. doi: 10.1080/00275514.2019.1636608
Hibbett DS, Binder M (2002) Evolution of complex fruiting - body Vargas-Estupiñan N, Benjumea C, Corrales A, Gomez-Montoya N, Peña-
morphologies in homobasidiomycetes. Proceedings of the Royal Cañón R, Vasco-Palacios AM (2020) Amanita sepultipes (amended
Society of London. Series B: Biological Sciences, 269(1504), 1963- version of 2020 assessment). The IUCN Red List of Threatened Species,
1969. doi: 10.1098/rspb.2002.2123 2020, e.T174797136A179540675. doi: 10.2305/IUCN.UK.2020-3.
IUCN (2021) The IUCN Red List of Threatened Species, version 2021–1. RLTS.T174797 136A179540675
Available at: https://www.iucnredlist.org Vasco-Palacios AM, Calle A, Drechsler-Santos ER, Kossmann T, da Cunha
Kirk PM, Cannon P, Minter D, Stalpers J (2008) Ainsworth & Bisby’s KM, TEHO Group (2020) Austroboletus amazonicus (amended version
Dictionary of the Fungi. 10th ed. CAB International, Walling ford, UK. of 2020 assessment). The IUCN Red List of Threatened Species, 2020,
Kwon-Chung KJ, Fraser JA, Doering TL, Wang ZA, Janbon G, Idnurm A, e.T75098759A179542191. doi: 10.2305/IUCN.UK.2020-3.RLTS.
Bahn YS (2014) Cryptococcus neoformans and Cryptococcus gattii, the T75098759 A179542191.en
etiologic agents of cryptococcosis. Cold Spring Harbour Perspectives Watkinson S, Boddy L, Money N (2016) The Fungi. Third ed. London:
in Medicine, 4(7), a019760. doi: 10.1101/cshperspect.a019760 Academic Press. 453p.
Londoño AC, Alvarez E, Forero E, Morton CM (1995) A new genus and Weiß, M, Bauer R, Sampaio JP, Oberwinkler F (2014) Tremellomycetes
species of Dipterocarpaceae from the Neotropics. I. Introduction, and related groups. In: McLaughlin DJ, Spatafora JW (eds). The
taxonomy, ecology, and distribution. Brittonia, 47(3), 225-236. Mycota VII Part A. Systematics and Evolution. Berlin: Springer.
López-Quintero CA, Straatsma G, Franco-Molano AE, Boekhout T (2012) Pp. 331-355.
Macrofungal diversity in Colombian Amazon forests varies with World Atlas (2021) Countries of the World. Available at: https://www.
regions and regimes of disturbance. Biodiversity and Conservation, worldatlas.com/
21(9), 2221-2243. doi: 10.1007/s10531-012-0280-8 Zajc J, Gunde-Cimerman N (2018) The genus Wallemia-from contamination
Malysheva V, Schoutteten N, Verbeken A, Spirin V (2021) Identity and of food to health threat. Microorganisms, 6(2), 46. doi: 10.3390/
typification of Achroomyces effusus (Pucciniomycotina, Basidiomycota). microorganisms6020046
Mycological Progress, 20(4), 413-417. doi: 10.1007/s11557-021- Zamora JC, Ekman S (2020) Phylogeny and character evolution in
01671-2 the Dacrymycetes, and systematics of Unilacrymaceae and
McLaughlin DJ, Healy RA, Kumar TA, McLaughlin EG, Shirouzu T, Binder Dacryonaemataceae fam. nov. Persoonia.: Molecular Phylogeny and
M (2016) Cultural and cytological characterisation of Dacryopinax Evolution of Fungi, 44, 161-205. doi: 10.3767/persoonia.2020.44.07
primogenitus, a new species in the Dacrymycetes with a fully Zhao RL, Li GJ, Sánchez-Ramírez S, Stata M, Yang ZL, Wu G, Dai YC, He
sequenced genome. Mycologia, 108(2), 457-468. doi: 10.3852/15- SH, Cui BK, Zhou JL, Wu F (2017) A six-gene phylogenetic overview
229 of Basidiomycota and allied phyla with estimated divergence times of
Millanes AM, Diederich P, Ekman S, Wedin M (2011) Phylogeny and higher taxa and a phyloproteomics perspective. Fungal Diversity, 84,
character evolution in the jelly fungi (Tremellomycetes, Basidiomycota, 43-74. doi: 10.1007/s13225-017-0381-5
Fungi). Molecular Phylogenetics and Evolution, 61(1), 12-28. doi:
10.1016/j.ympev.2011.05.014
Moore RT (1980) Taxonomic proposals for the classification of marine
yeasts and other yeast-like fungi including the smuts. Botanica
Marina, 23 (1980), 361-373.
Cordyceps nidus
[Robert Lücking]
66 CATALOGUE OF FUNGI OF COLOMBIA
Chapter 5
Diversity of Non-Lichenised Macro-Ascomycota of Colombia
Tatiana Sanjuan1* & Kent Brothers2
Grupo Micologos de Colombia, Bogota DC, Colombia.
1
ABSTRACT
Ascomycota is the most diverse phylum of the kingdom Fungi, comprising 92,725 species described to date. They are
characterised by producing spores in a sac that resembles a row of peas, a so-called ascus, in higher Ascomycota formed
in structures termed ascomata. In most Ascomycota, the ascomata are primarily microscopic structures, reaching a few
millimeters in diameter, but some Ascomycota produce macro-ascocarps comparable in size to those of higher Basidiomycota.
Unfortunately, there is no straightforward definition of what would constitute macro-Ascomycota, as size differences are often
gradual and workers define "macro" and "micro" in this phylum differently. Globally the larger non-lichenized Ascomycota
are found primarily within the classes Geoglossomycetes, Leotiomycetes, Orbiliomycetes, Pezizomycetes, Sordariomycetes,
and Xylobotryomycetes. These classes encompass a total of 36,462 species, corresponding to 39% of the currently known
Ascomycota, although not all their members can be considered macro-Ascomycota and there are a few other classes with small
numbers of larger species which are not discussed here. The purpose of this chapter is to elaborate on the state of knowledge
of the diversity of the fungi within these classes in Colombia, based on records in the ColFungi portal and with additional
records from local herbaria, literature, and private collections. In the past ten years, Colombia increased the number of known
species of macro-Ascomycota from 181 to 1,114. Regarding the number of records, Colombia has 6% of the total reported for
the Neotropical region, compared to Brazil with 41%. Notably, the macro-Ascomycota deposited in herbaria outside Colombia
comprises mostly Leotiomycetes, whereas Sordariomycetes are the best represented class in Colombian collections. Rather
than the relative abundance of the various fungal groups, this difference reflects the interest of mycologists who collected
these specimens. A representative inventory for the true diversity of macro-Ascomycota in Colombia is still needed.
RESUMEN
Los Ascomycota son el filo más diverso de hongos, con 92.725 especies. Estos hongos caracterizados por su producción de
esporas en sacos pueden ser saprofíticos patógenos de animales, plantas y otros hongos, y simbiontes, en asociación con
algas, como los líquenes, o trabajando en mutualismo con plantas, como en las ectomicorrizas y los endófitos. De manera
notable, mientras en el mundo, los líquenes corresponden al 13% del total de las especies de ascomicetes registradas, en
Colombia corresponden al 52% de las especies de anotadas en ColFungi, sesgo derivado de su estudio tradicional asociado
a la botánica. En cambio, los macro-Ascomycota no liquenizados, según lo definido aquí, representan un 24% de los registros
de ascomicetes en Colombia, en comparación con 39% en el mundo. Los ascomicetes que tienen su ascoma o estructura
reproductiva asexual a simple vista se ubican en el subfilo Pezizomycotina, en las clases Geoglossomycetes, Leotiomycetes,
Orbiliomycetes, Pezizomycetes, Sordariomycetes y Xylobotryomycetes, además de algunos miembros del orden Eurotiales dentro
de la clase Eurotiomycetes. Para 2013, el listado de macro-Ascomycota de Colombia alcanzaba 181 especies, mientras que
en esta revisión se ha recopilado información de al menos 1,114 especies de esas clases, tomando los datos de ColFungi,
así como de literatura gris, herbarios nacionales y colecciones privadas. En cuanto a las colecciones de especímenes
preservados, Colombia, en comparación con otros países de la región neotropical, se ubica por debajo de Brasil, que posee el
41% de las colecciones, seguido de Costa Rica, con 21% y México con 18%. Colombia y Venezuela, países vecinos con áreas
similares, poseen una proporción del 6% y 8% respectivamente. Cuando se analiza la distribución de las clases de macro-
ascomicetes no liquenizados en los diferentes países de la región, Colombia se destaca por su mayor número de registros de
discomicetes de la clase Leotiomycetes. Esto se debe a los estudios esporádicos de investigadores extranjeros con enfoque
en sus especialidades taxonómicas en la década de los 70 y 80. Sin embargo, cambia cuando se analizan los herbarios
locales no conectados con las bases internacionales, donde se nota que los especialistas nacionales se han enfocado en
la clase Sordariomycetes. Las 1,114 especies de macro-ascomicetes de Colombia no liquenizados registradas a la fecha
superan las 463 especies de Panamá y las 1.483 especies de Ecuador, porque este número incluye además líquenes.
Finalmente, la distribución de los macro-ascomicetes no liquenizados dentro del territorio nacional según su ecología se
da de manera heterogénea de acuerdo con el gradiente altitudinal que va de la cordillera de los Andes hacia la cuenca
Amazónica. Las especies ectomicorrizicas se distribuyen en los bosques nublados de roble, mientras los discomicetes
pueden abarcar desde las alturas de los páramos hasta las selvas bajas. Los Xylariales, en cambio, muestran su mayor
expresión de diversidad y tamaños en las tierras bajas, aprovechando la gran biomasa de lignina y celulosa de los bosques
tropicales. Cabe destacar que los hongos artrópodo-patógenos del orden Hypocreales se distribuyen por el país siguiendo los
patrones de diversidad de los artrópodos que sirven como hospederos, los cuales son más diversos en los bosques húmedos
tropicales bajos; en consecuencia, alli se presenta la mayor riqueza de, especies con 15 géneros y al menos 82 especies. La
presente revisión plantea el desafío de analizar los datos de macro-ascomicetes no liquenizados desde un mayor número de
fuentes secundarias para así estimar de una manera más amplia la enorme diversidad de estos hongos y ubicar los vacíos
de información y las áreas aun sin explorar.
several renowned mycologists. Among these publications, worldwide. Among the lichenised Ascomycota (primarily
Carpenter & Dumont (1978) expanded the distribution of the classes Arthoniomycetes, Coniocybomycetes, Lecanoro-
Bisporella discendens, found in many localities in the Andean mycetes, Lichinomycetes), Colombia accounts for 19%
region, and Veerkamp & Gams (1983) described new species of those worldwide. However, they constitute 52% of the
of soil fungi, Trichoderma inhamatum and Morteriella ornata. Ascomycota species reported for Colombia. So while
Studies of Colombian Ascomycota by local mycologists Colombia's lichen biota is well known, the same cannot
have, for the most part, been restricted in scope, most be said for non-lichenised Ascomycota. In particular, for
of them focusing on micro-Ascomycota with relevance to classes comprising mostly micro-Ascomycota, such as
phytopathology (Buritica, 1999). In the 1990s, Tobón (1991) Dothidiomycetes, Eurotiomycetes, and Laboulbeniomycetes,
published a study on the Ascomycota of the Andean region Colombian species represent only 2.4% of those registered
from the department of Antioquia, including 11 species of in the world.
discomycetes. Local mycologists continue to publish studies In comparison, the classes that include non-
on Ascomycota for Colombia, with some of them describing lichenised macro-Ascomycota as here defined, namely
new species, such as Cookeina colombiana (Raymundo et al., Geoglossomycetes, Leotiomycetes, Orbiliomycetes, Pezizo-
2020), others encompassing broader systematic studies, mycetes, Sordariomycetes, and Xylobotriomycetes, are
such as that for the genus Cordyceps s.lat. (Sanjuan et al., represented by 36,462 species worldwide and 1,114
2014; 2015). species in Colombia. They represent only 3% of the world
species, and inside of Colombia they comprise 24%, not as
many as the lichens, but more than the micro-Ascomycota.
COLOMBIAN ASCOMYCOTA IN NUMBERS
Overall, the Ascomycota in Colombia are not well sampled
To address the "state of knowledge" of the diversity of and studied. They might also not be as conspicuous and
ascomycetes in Colombia, it is useful to compare species useful as Basidiomycota for local communities in Colombia.
counts of the various taxa in Colombia to those for the Apart from lichens, incentives to study Ascomycota have
world and for similar countries. Referring to Table 1, we see mostly arisen from their medicinal importance or their
that Ascomycota reported for Colombia comprise 5% of the effects on crops (Sipman, 1986; Garces et al., 1999).
92,725 species known worldwide. For yeasts and dimorphic Indeed, throughout humanity’s history, we have mostly
yeasts (subphyla Saccharomycotina and Taphrinomycotina) studied what benefits or harms us, and so the amount of
Colombia registers 9% and 2%, respectively, of species knowledge reflects our anthropocentric point of view.
TABLE 1. Comparison of the number of species of Ascomycota in the world (Catalogue of Life, 2021) with species reported from Colombia.
World Colombian Colombian
Subphylum Class Life form
species species percentage
1,180 106
Saccharomycotina Saccharomycetes True yeast 9.0%
(1.3%) (2%)
Pneumocystomycetes 164 3
Taphrinomycotina Dimorphic yeast 1.8%
Schizosaccharomycetes (0.2%) (0.1%)
Non-lichen non-
Dothideomycetes Eurotiomycetes 37,238 906
yeast micro- 2.4%
Pezizomycotina Laboulbeniomycetes (40%) (20%)
ascomycetes
73
Incertae sedis
(2%)
FIGURE 1. Collections of classes containing non-lichenised macro-Ascomycetes in Neotropical countries based on GBFI (2021).
Xylobotryomycetes is omitted due to their low number of species. Numbers inside the circles represent the number of preserved specimens.
Shades of grey for each country reflect the corresponding total number of records.
STUDIES OF NON-LICHENISED MACRO-ASCOMYCOTA IN THE to entomopathogenic fungi within Hypocreales and lignicolous
NEOTROPICS fungi from Xylariales. There is a notable difference in
When analysing the composition of Ascomycota classes in taxonomic coverage based on GBIF versus HUA records, likely
collections from each country (Figure 1), in most countries because the GBIF records are biased towards the collections
Sordariomycetes are the most abundant by far. This class made by Kent Dumont in the NY herbarium, who focused on
includes families such as Xylariaceae and Cordycipitaceae, Helotiales (Leotiomycetes) (https://mycoportal.org). In fungi
which are very common in tropical lowlands (Soto-Medina & collections, the abundance of a taxon is often explained by
Bolaños-Rojas, 2013; Sanjuan, 2015). Brazil is currently the the presence of a specialist who worked on it. The number
country with the highest number of digitized records in both of non-lichenised Ascomycota for Colombia (1,114 species)
absolute numbers and proportions (GBIF, 2021). By contrast, is greater than for Panama (483; Pipenbring, 2007) and
in Colombia, Leotiomycetes are more representative, including Ecuador; the latter has a total of 1,483 Ascomycota reported
Leotiaceae, a family with ectomycorrhizal relationships but that number includes lichens (Læssøe & Petersen, 2008;
with oak (Quercus humboldtii) and Trigonobalanus excelsa Batallas et al., 2020). This underlines Colombia's high fungal
(Sierra et al., 2011). Colombia represents the southernmost diversity, matching that of other organisms, such as birds,
distribution of these forest associations originating from the orchids, and amphibians (Álvarez Hincapié et al., 2021).
Northern Hemisphere (Gonzalez et al., 2006). Additionally,
this ecosystem is located in the highlands, where most REPRESENTATIVE GROUPS OF NON-LICHENISED MACRO-
mycological research has been done in Colombia. ASCOMYCOTA IN COLOMBIA
On the other hand, in the Antioquia University Herbarium To date, we have seen the abundance and diversity of
(HUA), Sordariomycetes are the most representative class, Ascomycota, especially the non-lichenised ones, at global
with 381 species recorded to date for Colombia, mostly due and regional scales. However, given Colombia's diversity of
ecosystems, the distribution of macro-Ascomycota is not 3,300 meters, corresponding to cloud forests and Andean
homogeneous across the country, responding to factors such paramos, classified under Holdridge's life zone system
as rainfall, temperature, and altitude (Álvarez Hincapie et al., (Holdridge, 1967) as moist montane forests (MMF) and sub-
2021). The most representative groups of macro-Ascomycota alpine rain paramo (SARP). The Pezizales Cookeina speciosa,
in Colombia can be grouped according to the shape of their Cookeina tricholoma (Figure 2d), and Phillipsia domingensis,
ascoma and/or their role in the ecosystem, relative to the are more frequently collected in lowlands, e.g., in the
most representative biomes of Colombia (Figure 2). Magdalena Valley and in the Amazon foothills and basin,
which correspond to tropical rainforests (RFT). However, a
CUPULATE ASCOMYCOTA new species of Cookeina was discovered recently, namely
The terms "cup fungi" or "discomycetes" are informally used C. colombiana, found in tropical dry forests (DFT) in the
for Ascomycota with discoid, more or less sessile ascomata, Caribbean region (Raymundo et al., 2020). Thus, although
representing the classes Leotiomycetes (including Helotiales discomycetes have been collected mainly throughout the
and some Rhytismatales), Orbiliomycetes (including Orbiliales), Andean, Amazon, and Pacific regions, the Caribbean and
and Pezizomycetes (including Pezizales). Discomycetes have Orinoquia would benefit from more field surveys, as revealed
148 species reported for Colombia, with Lachnum (Helotiales, by the newly discovered species of Cookeina.
Lachnaceae) being the most diverse and abundant, with 30
species and 150 records. Calycina claroflava (Helotiales, HYPOGEOUS ASCOMYCOTA
Helotiaceae) is the most frequently collected species, with Spore-sequestering fungi have evolved several times from
71 specimens in total. As part of the "Hongos de Colombia" different lineages of Ascomycota (Bonito & Smith, 2016).
project, Carpenter & Dumont (1978) recorded more than In Colombia, truffle-like fungi represent Pezizomycetes
70 specimens of C. claroflava, under its synonym Bisporella (Pezizales) and Eurotiomycetes (Eurotiales). These fungi are
discendens. The most extensive collection of discomycetes ectomycorrhizal, associated with oak (Quercus humboldtii)
has been made in the Andean region, between 1,500 and and Trigonobalanus excelsa trees. The false truffle,
FIGURE 2. Representative groups of non-lichenised macro-Ascomycota in Colombia. A Xylaria telfarii, from Tolima in the Magdalena valley.
B False truffle, Elaphomyces muricatus, parasitised by Tolypocladium capitatum, from Boyaca in the Andean region. C Xylobotryum sp.,
from Amazonas in the Amazon basin. D Cookeina speciosa and C. tricholoma, from Antioquia in the Magdalena Valley. E Leotia lubrica and
L. chlorocephala, from Huila in the Andean region. F Annulohypoxylon sp. from Andean cloud forest in Cundinamarca. G Ophiocordyceps
albacongiuae, from Antioquia in the Magdalena valley. (Photographs A, B, D and E by Tatiana Sanjuan; C, F and G by Kent Brothers).
Elaphomyces muricatus (Eurotiales, Elaphomycetaceae) spathulate or strongly stipitate ascomata are found in
has been reported from the department of Cundinamarca species of Geoglossomycetes (Geoglossaceae). Each of
by Guzmán & Varela (1978). However, there are collections these groups is characterized by a specific trophic mode
identified to genus from the departments of Antioquia, Boyacá, (Beug et al., 2014).
Huila, and Tolima, most of them parasitised by species of The genus Leotia (jelly babies; Figure 2e) includes three
Tolypocladium (Hypocreales, Ophiocordycipitaceae) (Herbario species in Colombia. Leotia lubrica is the most common,
de la Universidad de Antioquia, 2021) (Figure 2b). found above 2,400 meters as ectomycorrhiza of oak (Quercus
Another false truffle, with specimens housed in HUA, humboltii) in moist montane forests (MMF). The saddle
was collected in the departments of Antioquia and Tolima, fungus, Helvella, is found in the same region, although not
namely Hydnotrya tulasnei (Pezizales, Discinaceae), known only in oak forests but sometimes also in grasslands close
for its good flavour when young. A genuine truffle, Tuber to cloud forests (MWF) in the Andean region. In Colombia,
melanosporum (Pezizales, Tuberaceae), was reported three species of Helvella have been recorded, with Helvella
in 1986 from Antioquia. Given that this name covers a lacunosa being the most frequently collected.
European species of black Tuber, the Colombian specimen Morels are found in anthropized ecosystems in Colombia.
may have been identified erroneously, assuming that the The first formal report of morels was made from Cundinamarca
most common European species may also be present in by Pinzón-Osario & Pinzón-Osario (2017), although Matis had
Colombia. Another record of this genus was reported for already illustrated Colombian morels for the first time in 1783
Cundinamarca in 2019 by the specialist in hypogeous from the Royal Botanical Expedition (Figure 3). No information
about the location was recorded in that study. Given the lack
fungi, Matthew Smith (https://mycoportal.org/portal/
of historical records, it is difficult to know whether morels
collections/individual/index.php?occid=10515278), but no
were originally native or introduced to Colombia by Europeans.
DNA sequences are available for it yet. Truffles, in general,
According to Sanchez (2019), three infra-generic clades are
are undersampled in Colombia and would greatly benefit
present in Colombia: subsect. Elata, sect. Esculenta and sect.
from timely studies, given that oak forests in Colombia are
Rufobrunnea. Standardised cultures were made from a strain
classified as vulnerable by the IUCN due to deforestation for
of Morchella rufobrunnea found in Colombia (Sanchez, 2019).
agriculture and animal breeding activities. Morchella elata and M. gracilis have also been collected in
the countryside of Colombia in the departments of Cesar,
STIPITATE ASCOMYCOTA Cundinamarca, and Quindío.
Some macro-Ascomycota have ascomata supported by a Earth tongues from the class Geoglossomycetes
fleshy stalk (e.g., Pezizales, Helvellaceae), sometimes hollow comprise four species in Colombia: Geoglossum fallax, G.
and filled with jelly and with viscid discs (e.g., Leotiales, nigritum, Trichoglossum hirsutum, and T. walteri. They all
Leotiaceae), or with an alveolate, conical disc supported grow associated with mosses or debris in montane cloud
by a spongy stalk (Pezizales, Morchellaceae). Clavate to forests (MWF) with high precipitation. Also in these cases,
the names for Colombian material have been adopted from
Northern Hemisphere species, and no DNA sequences from
Colombian specimens are available to date. Thus, it would
not be surprising to discover that some species of this group
from Colombia are new to science.
STROMATIC ASCOMYCOTA
The class Sordariomycetes includes Ascomycota producing
perithecioid ascomata in well-developed, often conspicuous
stromata (Kirk et al., 2008). Two orders contain macro-
Ascomycota, namely Xylariales, producing carbonaceous
stroma on woody substrata, and Hypocreales, producing
often brightly coloured stromata on plants, animals, and
other fungi (Beug, 2014).
Xylariales in Colombia comprises nine families, 24 genera,
and 124 species, compared to 16 families, 223 genera, and
2,911 species worldwide (https://colfungi.org; Catalogue of
Life, 2021). This order is distributed throughout the country,
but with more species and collections from lowland tropical
rainforests. A revision of Sordariomycetes from Colombia
for the bamboo Guadua angustifolia evidenced nine new
FIGURE 3. Morels drawn by Matis in 1783, one of the painters records for the country, including Diatrype bermudensis
of the botanical expedition of the viceroyalty of New Granada (Diatrypaceae) (Soto -Agudelo et al., 2016). As expected,
coordinated by Jose Celestino Mutis (courtesy of the Royal Xylaria is the most diverse genus in Colombia with 49
Botanical Garden of Madrid). species, more than Panama with 33 species, but fewer
TABLE 2. Species of Hypocreales described for first time from Colombia and recent new records for the country and their distribution.
than Ecuador with 60 species (Pipenbring, 2007; Læssøe approach is necessary to understand the systematics of
& Petersen, 2008). Xylaria metaeformis and X. willsii were this order, which in other parts of the world has revealed
the first species of the genus reported from Colombia, from important antimicrobial properties (Hyde et al., 2019).
wood collected in Bogotá in the 1860s. The most recent Hypocreales comprises eight families, ten genera, and
record is X. bambusicola, collected in Quindio in the lowlands 415 species in Colombia, compared to 14 families, 434
of the Andes region (Soto -Agudelo et al., 2016). Xylariales are genera, and 5,313 species worldwide (https://colfungi.
represented by numerous collections in the HUA herbarium, org; Catalogue of Life 2021). This order has a broad range
with 488 specimens from the Amazon basin and the Choco of trophic modes, as animal, fungal, or plant pathogens,
biogeographic zone, but the Orinoquia region is poorly endophytes, or saprotrophs. As part of the "Hongos de
represented. A study of the distribution of X. telfarii (Figure Colombia" project, Dumont et al. (1976) collected 39
2a) and related species in Colombia confirmed that Xylariales species of Nectriaceae, five of which were new to science
occur predominantly under certain precipitation regimes, with (Table 2) (Schroers, 2001). The latter family, together with
no dependence on temperature (Miñana, 2018). That study Bionectriaceae and Xylariaceae, often have brightly coloured
also revealed sampling gaps for these species throughout stromata that develop on various types of wood (Beug et al.,
Colombia, which could be extended to all genera of this 2014). Both families were reported on Guadua angustifolia
order. Xylariales show high diversity in Colombia, but there in the department of Quindio in Colombia, with four new
are no studies yet based on DNA sequences. A polyphasic records for this country (Table 2) (Soto -Agudelo et al., 2016).
Hypocreales also contain one of the kingdom's most molecular tools. Basic research into these kinds of fungi in
notable and conspicuous fungi, the arthropod pathogens Colombia would likely benefit the country, as they appear to
colloquially called "zombie fungi", distributed among the have considerable biotechnological potential.
Clavicipitaceae, Cordycipitaceae, and Ophiocordycipitaceae
(Sung et al., 2007). In Colombia, 92 arthropod pathogens FUTURE PERSPECTIVES FOR NON-LICHENISED MACRO-
have been reported, and another seven fungal pathogens. ASCOMYCOTA OF COLOMBIA
Most species have been found in lowland tropical forests. Ascomycota of Colombia have mostly been studied as part
Around agricultural areas, such as those for coffee crops, of sporadic visits by foreign mycologists, and more recently
the genera Beauveria, Lecanicillium, and Metarhizium are also by trained local mycologists. Until now, the "Hongos
more frequently collected. However, two species of Beauveria de Colombia" project of Dumont and collaborators in the
were described from undisturbed forests in the Amazon and 1970s continues to be the most complete study to date
Putumayo departments, namely B. diapheromeriphila and on the macro-Ascomycota of Colombia, although these
B. acridophila (Table 2), which represent rare examples of workers favoured sampling areas with easy access at the
sexual forms of this cosmopolitan genus (Sanjuan et al., time, such as those near roads or farms, in an opportunistic
2014). By contrast, Akanthomyces sabanensis was described manner. There have been few inventories in pristine forests
from an urban street of high-level city of Bogotá (Table 2). in areas such as the Orinoquía region or on islands, such
This entomopathogen parasitises scale insects feeding as San Andres and Providencia in the Caribbean or Gorgona
on the emblematic tree Ficus soatensis, its distribution in the Pacific, where many gaps persist. Novel sampling
corresponding to areas of low pollution within the city (Chirivi methodologies are required that include all life forms and
et al., 2015). trophic modes of Ascomycota in a temporal framework to
The distribution of arthropod pathogens follows that assess the importance of these organisms for tropical
of their hosts, which are highly diverse and abundant, ecosystems in Colombia. Finally, it is necessary to train
especially in the Amazon basin. The orders Coleoptera new mycologists with an integrative approach that includes
(weevils and beetles) and Hymenoptera (ants and wasps) traditional taxonomy and molecular phylogenetics.
are the most parasitised in Colombia, with seven and
13 species, respectively, in concordance with the high
abundance and richness of both groups. However, the most Acknowledgements
abundant species in all altitudinal zones, Cordyceps tenuipes
(synonym: Isaria tenuipes), attacks pupae of Lepidoptera We thank Thomas Læssøe and Rosy Batallas for making the
Ecuador data available for this study, and Samuel Guerrero for his
(Sanjuan, 2015). This species is found in lowland tropical
help with the English writing. This review was possible due to the
forests (TRF) as well as in montane moist forests (MMF) financial support of the Grupo Micologos Colombia (NPO).
and shares habitats with Tolypocladium capitatum (synonym:
Cordyceps capitatum). It attacks the hypogeous ascomycete
Elaphomyces muricatus, an ectomycorrhizal associate of oak References
belonging to the Eurotiales.
Aguirre-C J (1985) Algas, líquenes, hongos, hepaticas y musgos / Ilustran
Considerable attention is given to these arthropod este Tomo con 51 láminas en color y 1 monocroma Francisco Javier Matis,
pathogens, due to their ability to manipulate their host's Antonio Cortes, Salvador Rizo y otros pintores de la Flora de Bogotá,
behaviour to their benefit, especially the zombie ants, cuyas laminas no fueron firmadas. Identificó las láminas, interpretó las
which have been widely documented (Araújo et al., 2018). descripciones de los botánicos de la Expedición Botánica y redactó los
textos Jaime Aguirre Ceballos, colaboró en la identificación de láminas y
In particular, Ophiocordyceps unilateralis manipulates ants en la redacción de parte de los textos de los Hongos Francisco de Diego
of the genus Camponotus (Formicinae, Camponotini), forcing Calonge. Ediciones Cultura Hispánica, Madrid. 83p.
them to die in particular locations (Andersen et al., 2009). Álvarez Hincapié CF, Maldonado-Ocampo J, Sanjuan T, Álvarez-Dávila E,
This behaviour was already observed by the Inga indigenous Ajiaco RE, Buitrago L, González MA, Herrera-R GA, Ortíz Gallego R,
Plata C, Polanco A, Rosselli L, Uribe U, Alterio H, Barrero EY, Laverde-
people in the Putumayo region of Colombia, where it forms RO (2021) Estado de la biodiversidad en Colombia. In: Gómez-S
part of a myth from the indigenous cosmology (Sanjuan, R, Chaves ME, Ramírez W, Santamaría M, Andrade G, Solano C,
1999). Three new ant pathogens have been reported from Aranguren S (eds). Evaluación Nacional de Biodiversidad y Servicios
Colombia, one of which, Ophiocordyceps albacongiuae (Figure Ecosistémicos de Colombia. Bogotá DC: Instituto de Investigación de
Recursos Biológicos Alexander von Humboldt, Programa de Naciones
2g), was discovered in Río Claro canyon, a tropical forest in the Unidas para el Desarrollo y el Centro Mundial de Monitoreo para la
karst limestone formations of the Magdalena valley. Another Conservación del Programa de las Naciones Unidas para el Medio
33 species of arthropod pathogens have been reported Ambiente, Ministerio Federal de Medio Ambiente, Conservación de la
from the same location, showing a high diversity in a small Naturaleza y Seguridad Nuclear de la República Federal de Alemania.
In Press.
area with species common to the Amazon basin (Posada & Andersen SB, Gerritsma S, Yusah KM, Mayntz D, Hywel-Jones NL, Billen J,
Sanjuan, 2018). Unfortunately, this location is threatened Hughes DP (2009) The life of a dead ant: the expression of an adaptive
by a limestone mining operation that has accelerated after extended phenotype. The American Naturalist, 174(3), 424-433. doi:
the signing of the peace accord in Colombia. Overall, each 10.1086/603640
Araújo JPM, Evans HC, Kepler R, Hughes DP (2018) Zombie-ant fungi
year a new species of arthropod pathogens is discovered across continents: 15 new species and new combinations within
globally, and to date, 677 species have been described. Ophiocordyceps. I. Myrmecophilous hirsutelloid species. Studies in
However, the discovery of cryptic species requires the use of Mycology, 90, 119-160. doi: 10.1016/j.simyco.2017.12.002
Batallas R, Moya F, Navas D (2020) Listado de la colección de hongos Sanjuan T, Tabima J, Restrepo S, Læssøe T, Spatafora JW, Vasco-Palacios
(Ascomycota y Basidiomycota) del Herbario Nacional del Ecuador. (2014) Entomopathogens of Amazonian stick insects and locusts are
(QCNE) del Instituto Nacional de Biodiversidad (INABIO). Revista members of the Beauveria species complex (Cordyceps sensu stricto).
Avances para Ciencia e Ingenieria, 12(20), 38-71. doi: 10.18272/aci. Mycologia, 106(2), 260-275. doi: 10.3852/106.2.260
v12i1.1755 Sanjuan T (2015) Estudio filogenético de la interacción Cordyceps – insectos
Beug M, Bessette AE, Bessette AR (2014) Ascomycota Fungi of North en el noroeste amazónico, un análisis morfológico, molecular y ecológico.
America: A Mushroom Reference Guide, Vol. 69. Austin: University of Doctoral Dissertation. Universidad de Antioquia. Medellín. 126p.
Texas Press. 488p. Sierra Toro JD, Arias González JA, Sánchez Sáenz M (2011) Preliminary
Bonito GM, Smith ME (2016) General systematic position of the truffles: Report of Macrofungi (Ascomycota and Basidiomycetes) in the
evolutionary theories. In: Zambonelli A, Iotti M, Murat C (eds). True Romeral's Montain Humid Forest (Angelopolis, Antioquia, Colombia).
Truffle (Tuber spp.) in the World. Berlin: Springer. Pp. 3-18. Revista Facultad Nacional de Agronomía Medellín, 64(2), 6159-6174.
Buritica P (1999) Iconografía fitopatológica colombiana. Revista de Sipman HJM (1986) Three new lichens from Colombia. Willdenowia, 16,
la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 279-284.
23(86), 81-95. Soto-Agudelo R, Valenzuela R, Gómez-Marín G, Raymundo T (2016)
Carpenter SE, Dumont KP (1978) Hongos de Colombia IV. Bisporella Ascomicetos de los guaduales en la reserva natural La Montaña del
triceptata and its allies in Colombia. Caldasia, 58(12), 339-348. Ocaso departamento del Quindío Colombia. Revista de la Academia
Catalogue of Life (2021) Fungi. Available at: https://www.catalogueoflife. Colombiana de Ciencias Exactas Físicas y Naturales, 40(156), 438-
org/ 449. doi: 10.18257/ raccefyn.352
Cepero de García MC, Cardenas M, Franco-Molano AE, Restrepo S, Soto-Medina E, Bolaños-Rojas AC (2013) Xylariaceae en un bosque
Vargas N (2012) Biología de hongos. Bogotá DC: Ediciones Uniandes- de niebla del Valle del Cauca (Colombia). Revista de la Academia
Universidad de los Andes. 520p. Colombiana de Ciencias Exactas, Físicas y Naturales, 37(144), 343-
Chardón CE, Toro RA (1930) Mycological explorations of Colombia. The 351.
Journal of Agriculture of the University of Puerto Rico, 14(4), 195-369. Sung GH, Hywel-Jones NL, Sung JM, Luangsa-Ard JJ, Shrestha B,
doi: 10.46429/jaupr.v14i4.14223 Spatafora JW (2007) Phylogenetic classification of Cordyceps and the
Chirivi-Salomon JS, Danies G, Restrepo S, Sanjuan T (2015) Lecanicillium clavicipitaceous fungi. Studies in Mycology, 57, 5-59. doi: 10.3114/
sabanense sp. nov. (Cordycipitaceae) a new fungal entomopathogen of sim.2007.57.01
coccids. Phytotaxa, 234(1): 63-74. doi: 10.11646/phytotaxa.234.1.4 Tobón LE (1991) Ascomycota de Colombia: Discomicetes del departamento
Garces E, Orozco M, Zapata AC (1999) Fitopatología en flores de de Antioquia. Caldasia, 16(78), 327-336.
Colombia. Acta Biológica Colombiana, 4(2), 5-26. Veerkamp J, Gams W (1983) Hongos de Colombia VIII. Some new species
GBIF - Global Biodiversity Information Facility (2021) GBIF Occurrences. of soil fungi from Colombia. Caldasia, 65(13), 709-717.
Available at: https://www.gbif.org/ Voglmayr H, Fournier J, Jaklitsch WM (2019) Two new classes of
González C, Jarvis A, Palacio JD (2006) Biogeography of the Colombian Ascomycota: Xylobotryomycetes and Candelariomycetes. Persoonia, 42,
Oak, Quercus humboldtii Bonpl: Geographical Distribution and Their 36-49. doi: 10.3767/persoonia.2019.42.02
Climatic Adaptation. Popayán: International Centre for Tropical
Agriculture (CIAT)/Museo de Historia Natural, Universidad del Cauca.
Herbario de la Universidad de Antioquia (2021) Base de ejemplares.
Available at: http://www2.udea.edu.co/herbario/paginas/consultas/
consultarEjemplares.iface
Holdridge LR (1967) Life Zone Ecology. San José: Tropical Science Center.
Hyde KD, Xu J, Rapior S, Jeewon R, Lumyong S, Niego AGT, Stadler M
(2019) The amazing potential of fungi: 50 ways we can exploit fungi
industrially. Fungal Diversity, 97(1), 1-136. doi: 10.1007/s13225-
019-00430-9
Kirk PM, Canon PF, Minter DW, Staplers JA (2008) Dictionary of the Fungi,
10th Edition. CABI.
Læssøe T, Petersen J (2008) Equatorial fungi - mycological biodiversity in
Ecuador. Svampe, 58, 1-52.
Miñana S (2018) Xylaria telfairii en Colombia: variación morfológica,
distribución geográfica y comentarios sobre su estatus taxonómico.
Undergraduate Thesis. Bogotá: Universidad Nacional de Colombia.
27p.
Piepenbring M (2007) Inventoring the fungi of Panama. Biodiversity and
Conservation, 16, 73-84. doi: 10.1007/s10531-006-9051-8
Pinzón-Osario CA, Pinzón-Osario J (2017) Primer registro de la familia
Morchellaceae (Ascomycota, Pezizales) para Colombia. Revista
Peruana de Biología, 24, 105-110. doi: 10.15381/rpb.v24i1.12481
Posada A, Sanjuan T (2018) Cañon del Rio Claro Reserva Natural is a
refuge of species of entomopathogenic fungi Cordyceps s.l. in the
Magdalena river basin, Colombia. Conference paper 11 International
Mycological Congress. San Juan de Puerto Rico.
Raymundo T, Montes-Fuentes G, Valenzuela R (2020) Cookeina
colombiana (Sarcoscyphaceae, Ascomycota), una especie nueva del
departamento Córdoba, Colombia. Acta Botánica Mexicana, 128,
e1763. doi: 10.21829/abm 128.2021.1763
Sanchez JA (2019) Caracterización del hongo Morchella sp. para
Cundinamarca (Colombia) y estandarización de las condiciones
Nutricionales para su Producción sobre Sustratos Agrícolas.
Undergraduate Thesis. Universidad Nacional de Colombia.
Bogotá.153p.
Sanjuan T (1999) La diversidad del género Cordyceps en hormigas
del bosque húmedo tropical de Colombia. Undergraduate Thesis.
Universidad Nacional de Colombia. Bogotá. 126p.
Phaeographis galeanoae
[Robert Lücking]
76 CATALOGUE OF FUNGI OF COLOMBIA
Chapter 6
Diversity, Ecogeography, and Importance of Lichens of Colombia
Bibiana Moncada1,2,8, Luis Fernando Coca3,8, David Díaz-Escandón4,8, Margarita Jaramillo-Ciro5,8, Diego Simijaca-Salcedo6,8, Edier Soto-
Medina7,8 & Robert Lücking2,8*
1
Licenciatura en Biología, Universidad Distrital Francisco José de Caldas, Cra. 4 No. 26D-54, Torre de Laboratorios, Herbario, Bogotá DC, Colombia.
2
Botanischer Garten und Botanisches Museum, Freie Universität Berlin, Königin-Luise-Straße 6–8, 14195 Berlin, Germany.
3
Herbario Universidad de Caldas (FAUC), Edificio Bicentenario, Calle 65 No. 26-10, Manizales, A.A. 275, Caldas, Colombia.
4
Department of Biological Sciences CW405, University of Alberta, Edmonton, Alberta, Canada.
5
Modeling and Computational Simulation Research Group (Grupo de Investigación en Modelamiento y Simulación Computacional – GIMSC), School of
Engineering, San Buenaventura University, Carrera 56C No. 51-110, Medellín, Colombia.
6
Centro de Ciencias Básicas, Universidad Autónoma de Aguascalientes, Aguascalientes, México.
7
Grupo de Ecología y Diversidad Vegetal, Departamento de Biología, Facultad de Ciencias Naturales, Universidad del Valle, Cali, Colombia.
8
Grupo Colombiano de Liquenología (GCOL).
ABSTRACT
Lichenised fungi constitute a substantial portion of the known Colombian fungi, with 2,670 out of 7,241 species. This fairly
high number is not because lichens represent a particularly diverse group of fungi but because they are relatively well-
studied in the country compared to non-lichenised fungi. Lichens have traditionally been defined as a symbiosis between a
fungus (mycobiont), an alga and/or a cyanobacterium (photobionts). A modern definition also incorporates components of the
lichen microbiome, particularly other fungi, and bacteria. However, the scientific names given to lichens strictly refer to the
primary mycobiont. Globally, Colombia ranks among the top ten countries in terms of known lichen diversity. Most top-ranking
countries are outside the tropics, so this supports Colombia’s position among the three most biodiverse tropical countries
worldwide. The total number for Colombia is estimated at 5,000 species, almost twice the number of species currently known.
Unrecognised species are predicted to be found in understudied regions, understudied groups, and in genera where broad
species concepts may include substantial hidden diversity. Diverse ecological studies assess environmental factors, such
as altitudinal range, topography, and habitat diversity, as drivers of lichen biodiversity. On the basis of these findings, the
impact of land-use changes and environmental pollution on lichen communities can be quantified, providing the foundation
for using lichens as bioindicators to monitor ecosystem health and to perform environmental impact studies. However, such
applications first require a systematic inventory of the lichens of Colombia.
RESUMEN
Los hongos liquenizados constituyen una parte sustancial de la funga Colombiana, con 2.670 de las 7.241 especies
conocidas. Este número bastante alto no se debe a que los líquenes representen un grupo de hongos particularmente
diverso, sino a que están relativamente bien estudiados en el país en comparación con los hongos no liquenizados. Los
líquenes se han definido tradicionalmente como una simbiosis entre un hongo (micobionte) y un alga y/o una cianobacteria
(fotobiontes). Una definición moderna también incorpora componentes del microbioma del liquen, en particular otros
hongos y bacterias. Sin embargo, los nombres científicos dados a los líquenes se refieren estrictamente al micobionto
primario. A nivel mundial, Colombia se encuentra entre los diez países con mayor diversidad de líquenes conocida. El pais
de mayor diversidad liquénica conocido es Estados Unidos (4,341 especies), seguido por Brazil (4,310), Australia (4,003),
China (3,050), Francia (2,917), México (2,722), India (2,714) e Italia (2,704). Sin embargo, la mayoría de los países con alta
diversidad liquénica están fuera de los trópicos, lo que respalda la posición de Colombia entre los tres países tropicales
con mayor biodiversidad en todo el mundo. El número total de especies liquenizadas para Colombia se estima en 5,000,
casi el doble del número de especies conocidas actualmente. Se predice que las especies no reconocidas se encontrarán
en regiones poco estudiadas, en grupos taxonómicos poco estudiados y en géneros donde los conceptos tradicionales de
especies pueden incluir una alta diversidad críptica. Históricamente, Colombia es uno de los países más importantes en
cuanto a estudios sobre líquenes neotropicales, junto a Brasil, Cuba y México. Los líquenes de Colombia se documentaron
por primera vez como parte de la Expedición Botánica del Nuevo Reino de Granada y colecciones realizadas por Humboldt
y Bonpland; sin embargo, las colecciones históricas de líquenes más importantes de Colombia son las de Lindig, en su
mayoría estudiadas por Nylander. Consecuentemente, más de 400 tipos se encuentran en los herbarios H y PC, siendo
una referencia histórica esencial para los líquenes (neo) tropicales. Diversos estudios ecológicos han evaluado los factores
ambientales, como el rango altitudinal, la topografía y la diversidad de hábitats, como impulsores de la biodiversidad
de líquenes en Colombia. Con base en estos hallazgos, se puede cuantificar el impacto de los cambios en el uso de
la tierra y la contaminación ambiental en las comunidades de líquenes, siendo la base para el uso de líquenes como
bioindicadores para monitorear la salud de ecosistemas y realizar estudios de impacto ambiental. Sin embargo, tales
aplicaciones requieren un inventario sistemático más profundo de líquenes de Colombia.
INTRODUCTION cases, whereas the vernacular name can be used for the
Lichens have long been treated as a separate group of entire lichen. For example, in the case of the widespread
organisms, even after discovering their symbiotic nature Xanthoria parietina (yellow wall lichen), a species also found
(Lücking et al., 2021). Biologically, lichens are just one in Colombia, the scientific name (X. parietina) applies to the
example of the diverse nutritional types that have evolved mycobiont alone. By contrast, the common name (yellow wall
among the Fungi, including saprotrophs, pathogens, lichen) addresses the entire lichen (Lücking et al., 2021).
mycorrhizas, and even carnivores (Watkinson et al., 2016). Lichens are integral components of terrestrial ecosystems
However, in lichens, the vegetative hyphae form a persistent, and sometimes are the dominant or exclusive life form
macroscopically conspicuous thallus (Honegger, 2012; (e.g., in the Antarctic continent). Even aquatic and marine
Figure 1). By contrast, non-lichenised fungi are characterised lichens have been known for almost 50 years (Seaward,
by ecologically hidden mycelia that are only usually visible 1977; Sanders et al., 2004; Feuerer & Hawksworth, 2007;
due to their spore-bearing structures. Mia˛dlikowska et al., 2014). Lichens are generally believed
After its discovery, the lichen symbiosis has continuously to escape the temperate-tropical diversity gradient, being
been defined as bipartite or tripartite associations between more conspicuous in temperate to arctic regions (Lücking
a fungal partner (mycobiont) and one or two photosynthetic et al., 2011). However, this is a matter of scale: at larger
partners (photobionts; Lücking et al., 2021). Known lichen scales (regional and landscape level), the species richness
mycobionts are only found in the two major phyla of the Fungi, of temperate and tropical regions is comparable, whereas
representing either Ascomycota (99%) or Basidiomycota (1%) at smaller scales (local and habitat level), species richness
(Lücking et al., 2017). At the same time, known photobionts appears to be greater in the tropics (Lücking et al., 2011).
encompass two domains and three kingdoms: Chlorophyta Notably, the highest species richness is found in tropical
and Heterokonta among the eukaryotes and Cyanobacteria lowland rainforests. Nonetheless, lichens are not conspicuous
among the prokaryotes (Friedl & Büdel, 2008; Saini et al., in these ecosystems, and their associated biomass is low.
2019). Recent studies using advanced molecular approaches Yet, a single square kilometre (100 ha) of tropical rainforest
have challenged the bipartite or tripartite nature of the lichen can harbour up to 600 species, including several hundred
symbiosis, postulating that other fungi, in particular yeast- species of foliicolous lichens growing on living leaves (Lücking
like Basidiomycota, as well as certain bacteria, together et al., 2011). For comparison, the best-studied temperate
constitute the lichen microbiome as possible integral regions may yield higher species numbers, but across much
components of the lichen symbiosis (Grube et al., 2015; larger areas, such as the Cévennes National Park in France
Spribille et al., 2016; Lendemer et al., 2019; Hawksworth (973 species within 93,700 ha), Glacier Bay National Park in
& Grube, 2020; Sierra et al., 2020; Grimm et al., 2021; Alaska (831 species within 600,000 ha), and Klondike Gold
Tuovinen et al., 2021). Consequently, the definition of the Rush National Historical Park (668 species within 5,300
lichen symbiosis as a bipartite or tripartite association ha), among others (Roux et al., 2008; Spribille et al., 2010,
of one mycobiont and one or two photobionts has been 2020; Lücking et al., 2011). The main difference between
abandoned in favour of a broader definition encompassing these temperate and the hitherto studied tropical sites is
these additional biological components (Hawksworth & that landscape diversity largely drives lichen diversity in
Grube, 2020; Lücking et al., 2021). the former. By contrast, high species richness has been
Owing to their symbiotic nature, the nomenclature applied reported in tropical locations for more or less homogeneous
to lichens has also posed challenges. Before discovery of ecosystems, indicating that these habitats have a higher
the symbiosis, the scientific Latin name applied to the entire species carrying capacity (Lücking et al., 2011).
lichen, including its photobiont(s), at the time termed gonidia
(Lücking et al., 2021). With the realisation that lichens are BIODIVERSITY OF LICHENS OF COLOMBIA
composed of more than one organism, the need arose to Colombia is home to extraordinary biodiversity, given its
adjust the precise application of the scientific name, which geographic position at the conjunction between Central
was then ruled to apply to the mycobiont alone (Lücking and South America. The country is bordered by two oceans,
et al., 2021). This solution seemed straightforward, but comprises an abrupt topography ranging from sea level to
the argument could be raised that the same fungus may 5,775 m altitude, and has diverse ecosystems (Arbeláez-
form different lichens, so-called photosymbiodemes, when Cortés, 2013; Murcia et al., 2013; Rangel-Ch., 2015). This
associated with different photobionts (Figure 1), differing in great biodiversity also extends to lichens with 2,670 species
ecology or distribution (Goward, 2008). The scientific name reported to date from the country (Gaya et al., 2021; this
should be used for the mycobiont only to account for such paper). Colombia is only surpassed in lichen richness by
A B
C D
E F
FIGURE 1. A–B Thallus of Sticta lobarioides and section of a Parmeliaceae showing the different layers formed by the mycobiont and
the photobiont. C–D Cephalodia in Stereocaulon novogranatense (C) and Placopsis rhodocarpa (D). E–F Photosymbiodeme in Sticta aff.
subscrobiculata, with cyanobacterial lobes emerging from green-algal lobes (E) and vice versa (F). (Photographs by Robert Lücking).
TABLE 1. Species numbers of lichenised fungi reported from selected regions and countries worldwide.
Species / Species /
Country / territory Number Reference(s)
1,000 km2 log2(km2)
United States 4,341 0.44 187 Perlmutter & Weakley (2018); Esslinger (2019)
India 2,714 0.83 125 Singh & Sinha (2010); Sinha et al. (2018)
United Kingdom 2,000 8.25 112 Smith et al. (2009); Coppins (2021)
FIGURE 2. A.Part of the “Lindig trail” in the “El Delirio Ecological Reserve”; B leading from the Southeastern outskirts of the municipality
of Bogotá DC at 2,800 m altitude up to the paramo region of El Verjón at 3,400 m altitude, along the Río Fucha; C this area is the type
locality for several species collected by Lindig, among them Sticta peltigerella, which typically grows on mossy boulders in streams and is
an excellent indicator of water quality. (Photographs by Robert Lücking).
the USA (4,341 species), Brazil (4,310), Australia (4,003), Colombia’s most important historical lichen collections
China (3,050), France (2,917), Mexico (2,722), India (2,714), (see Chapter 2). The famous “Lindig trail”, presumably
and Italy (2,704) (Singh & Sinha, 2010; Herrera-Campos et the type locality of many new species described from
al., 2014; Nimis, 2016; Perlmutter & Weakley, 2018; Sinha Bogotá DC, is still preserved in the “El Delirio” Reserve
et al., 2018; Esslinger, 2019; McCarthy, 2020; Aptroot, (Figure 2). Unfortunately, the exact localities of the collected
2021; Wei, 2021; Table 1). These countries are much larger specimens are unknown, as most only indicate “Bogotá”.
than Colombia (USA, Canada, Brazil, Australia, China, and Most of Lindig’s collections were studied in comprehensive
India) or represent much better studied temperate countries treatments by Nylander (1859, 1863a–c, 1864, 1867).
(France and Italy). Among the ten richest countries in relative Over 400 types of names applying to (neo-) tropical species
species density, Colombia ranks fourth with 2.34 species are now housed in the H and PC herbaria. This number is
per 1,000 km2, trailing Austria (28.0), Italy (8.97) and France comparable to names based on type material from Cuba
(4.53), but ahead of Mexico (1.38), India (0.83), Brazil (0.52), and only surpassed by the historical role of Brazil (Marcelli,
Australia (0.52), the USA (0.44), and China (0.32). When 1998). Therefore, the Colombian lichen biota is an essential
considering a logarithmic relationship of species richness historical reference for (neo-)tropical lichens. In modern
with area size, Colombia ranks seventh among the ten most times, Harrie Sipman and collaborators (see Chapter 2) made
species-rich countries [133 species per log2(area)], after the substantial contributions to the knowledge of Colombian
USA (187), Brazil (187), Australia (175), France (151), Italy lichens, with comprehensive checklists enumerating 1,553
(149), and Austria (144), but ahead of China (131), Mexico and more recently 1,674 species (Sipman et al., 2008;
(130), and India (125). Some small countries or territories Sipman & Aguirre, 2016).
with comparatively high species numbers surpass the high Although the currently known 2,670 species already
species density of Italy, ranking far ahead of Colombia as represent a high number, the true diversity of the lichens of
well, such as Puerto Rico (85.8 species per 1,000 km2), Colombia is probably much higher. Three reasons account
Costa Rica (34.0), and Cuba (9.4). Still, no further country for this assumption: 1. understudied and unexplored regions
included in this analysis has a higher species per log2(area) that harbour unknown species; 2. understudied taxonomic
count than Colombia. groups; and 3. refined species concepts in groups that
Overall, the high biodiversity of lichens of Colombia were considered to be well-known. GBIF (https://www.gbif.
reflects that of other groups of organisms for the country, org/occurrence/search?country=CO&taxon_key=180) has
such as plants, vertebrates, and insects (see Chapter 3), and over 55,000 occurrence records for Lecanoromycetes in
further supports the status of Colombia as one of the world’s Colombia, which is by far the largest class of lichenised
17 megadiverse countries (Mittermeier, 1988; Mittermeier Ascomycota. Most of these records are georeferenced, but
et al., 1999; Arbeláez-Cortés, 2013). It also corroborates only 38% correspond to preserved specimens, whereas 62%
Colombia’s position as the second most biodiverse tropical represent human observations, mostly through environmental
country, after Brazil (Butler, 2016, 2019). Most other top- impact assessments. The resulting map shows a clear bias
ranked countries and regions in terms of known lichen of existing collections to the Andean region. By contrast, the
diversity are largely or entirely outside the tropics. Caribbean, Pacific (Chocó biogeographic region), Orinoco,
Historically, Colombia is one of the most important and Amazon regions are grossly undersampled (Figure 3).
countries as regards studies on neotropical lichens, Such pattern is also seen for other organisms (Arbeláez-
alongside Brazil, Cuba, and Mexico. Lichens of Colombia Cortés, 2013). Given that tropical lichens are particularly
were first documented as part of the Expedición Botánica del diverse at lower altitudes and dry forests often harbour
Nuevo Reino de Granada, organised by José Celestino Mutis, unique taxa, many additional species can be expected from
with the collaboration of Francisco José de Caldas, between these four undersampled regions, which harbour large areas
1783 and 1816, with three colour plates representing the of lowland rainforests, savannas, and dry forests (Rincón-
species Baeomyces imbricatus (≡ Phyllobaeis imbricata), Espitia et al., 2011; Lücking et al., 2019; Soto-Medina et al.,
Cladonia didyma, and Stereocaulon ramulosum (Aguirre-C., 2021). A particular hotspot is expected in the geographically
1985; see Chapter 2). Halfway through this monumental isolated Sierra Nevada de Santa Marta, where only a few
endeavour, which unfortunately yielded few lichens and non- lichen studies have been carried out so far (Mägdefrau &
lichenised fungi, in 1801 Mutis and Caldas crossed pathways Winkler, 1967; Nowak & Winkler, 1970; Sipman, 1986;
with Alexander von Humboldt and Aime Bonpland in Bogotá Ramírez-Roncallo, 2018).
during the travels of the latter two through the northern Understudied taxonomic groups are particularly
Andes (Colombia, Ecuador, and Peru). The collections made represented by large genera that are either poorly collected
by Humboldt and Bonpland were treated by Hooker (1822), or lack thorough taxonomic studies because they have been
who reported 73 species, including 12 presumably new to deemed too difficult to study. Two examples are the genera
science, among them Baeomyces imbricatus (≡ Phyllobaeis Graphis (and its recent segregate Allographa) and Usnea.
imbricata), Lecidea parmelioides (= Coccocarpia erythroxyli), In a revision of fungarium specimens, Motta & Amórtegui
Pyrenula marginata (= P. mamillana), Sticta humboldtii, and S. (2018) unveiled 15 new species and 67 new records of
pallida (≡ Lobariella pallida). Graphis and Allographa for Colombia, some of which have
Alexander (Alexandre) Lindig, a German botanist who been published (Motta et al., 2019), raising the number
resided in Bogotá DC between 1859 and 1863, gathered of species known from Colombia by 149%. A molecular
FIGURE. 3. Grid maps of georeferenced records of Lecanoromycetes in Colombia available from GBIF (https://www.gbif.org/occurrence/
map?country=CO&taxon_key=180), showing records based on preserved specimens (left map) versus all records, i.e., also including
human observations (right map). Size of dots is proportional to number of records per grid. Note that human observation records (mostly
through environmental impact assessments) mitigate sampling gaps particularly in northwestern Colombia.
barcoding study of the genus Usnea suggested that its on average double the number of species in many genera,
true species richness for Colombia may be up to 2.5 times adding another 1,500 species. Thus, a revised estimate,
greater than is currently known (Moncada et al., 2020). considering under- and unexplored regions, understudied
Some examples of refined species concepts are found in groups, and hidden diversity would arrive at a number close
the genera Lobariella and Sticta (Ascomycota) and the genus to or above 5,000 species.
Cora (Basidiomycota). Before molecular studies, the three All major taxonomic groups and biotypes of lichens
genera were presumed to be well-known in Colombia, with are represented in Colombia. However, compared to
four species of Lobariella, 42 of Sticta, and a single species temperate lichen biotas, such as that of the United
of Cora (Sipman et al., 2008). Since then, these numbers Kingdom, crustose groups are overrepresented, particularly
have risen, based on molecular phylogenetic revisions, those primarily confined to tropical regions, such as
to 24, 78, and 49 species, respectively, increasing their Graphidales, Pyrenulales, and Trypetheliales, which feature
species richness by more than 200%. This increase is trentepohlioid photobionts adapted to tropical climates. By
particularly dramatic for the genus Cora, a well-known contrast, orders such as Acarosporales, Baeomycetales,
element of the Colombian Andean region. Lecideales, Pertusariales, Teloschistales, and Verrucariales,
Lücking et al. (2009) estimated the number of lichenised are underrepresented (Figure 4). Additional orders, such as
species in Colombia at 3,600. However, at that point, the Arthoniales, Caliciales, Lecanorales, and Peltigerales, exhibit
magnitude of the hidden diversity in presumably well-known a similar richness in tropical and temperate regions but
groups had not yet been realised. We anticipate that an encompass different species. A particularly diverse group
estimated 3,600 species would perhaps cover additional of tropical lichens are foliicolous taxa, which grow on the
species found in understudied regions or taxonomic groups. living leaves of vascular plants (Lücking, 2008; Mateus et
Yet, hidden diversity in presumably well-known lineages would al., 2012). The Colombian paramos is mentioned in terms
FIGURE 4. Comparison between Colombia and the United Kingdom in terms of the composition of lichenised fungi by order, representing a
tropical versus temperate region. Data from Gaya et al. (2021) and this chapter for Colombia, and Smith et al. (2009) and Coppins (2021)
for the UK.
of ecosystem diversity, featuring a high lichen biodiversity The division of Colombia into ecoregions (IDEAM et al.,
(Sipman, 2002; Cleef, 2008; Madriñán et al., 2013), with 2017) provides an excellent background for investigating
similar habitats only found in parts of Venezuela and tropical lichens beyond their taxonomy and systematics.
northern Ecuador. By contrast, the lichen biota of the Ecological, ecogeographical, and biogeographical studies
Colombian Amazon is broadly shared with neighbouring of Colombian lichens go back to the early 1970s, covering
countries, such as Venezuela, the Guianas, Brazil, Ecuador, topics such as distribution patterns, landscape ecology,
and Peru (Lücking et al., in prep.). altitudinal gradients, and habitat preferences (Sipman,
1986, 1989, 1998; Ahti, 1992; Wolf, 1993a; Kessler,
ECOGEOGRAPHY OF LICHENS OF COLOMBIA 2000; Aguirre-C. & Sipman, 2004; Pinzón & Linares,
2006; Aguirre-C., 2008; Aguirre-C. & Rangel-Ch, 2008;
Compared to temperate regions in North America and
Pérez-Quintero & Watteijne-Cerón, 2009; Moncada et al.,
Europe, studies on the ecology and distribution of Colombian
lichens are still sparse. Regarding the distribution patterns 2014). Other studies focused on community and population
of Colombian lichens, to date, there is no quantitative structure, substrate specificity, and niche differentiation
analysis, so here we provide a first approach based on (Nowak & Winkler, 1970, 1975; Wolf, 1993b, c, 1994,
the assessment of 2,678 species (Figure 5). According 1995; Soto-Medina et al., 2012, 2015; Zárate-Arias et al.,
to this assessment, most species (35%) are neotropical, 2019). More recent studies have investigated functional
followed by subcosmopolitan (16%), circumpacific (15%), traits of tropical lichens (Chilito-López et al., 2016; Soto-
and pantropical (15%) species. Only a small portion of Medina et al., 2019) or their ecophysiology (Pulido-Herrera
8.5% is currently considered endemic. However, this & Ramos-Montaño, 2016), whereas some classic studies
assessment is preliminary, as molecular studies have analysed lichen-related nitrogen availability and nutrient flux
shown that presumably widespread taxa often represent in tropical forest ecosystems (Forman, 1975; Veneklaas,
species complexes. Thus, we expect the actual proportion 1991). Interactions between lichens and invertebrates,
of widespread, cross-continental species to be lower, specifically regarding lichen-related camouflage, are a
favouring a higher number of neotropical and endemic taxa popular topic in entomological studies (Rivera et al., 2011;
(see Chapter 10). Furthermore, species currently classified Cadena-Castañeda, 2013; Londoño et al., 2017; Lisi et al.,
as circumpacific (i.e., known from the Neotropics and the 2019). The availability of advanced molecular methods in
Eastern Paleotropics) are largely artefactual due to a lack Colombian laboratories has also allowed new investigations
of data from tropical Africa where these species may occur. into the microbiome of paramo lichens, revealing patterns
of host specificity (Sierra et al., 2020). Additionally, several These diverse studies have provided insight into which
studies have addressed the impact of land use change on environmental factors drive lichen biodiversity and species
lichen community diversity and composition (Ardila-Rios et richness, demonstrating how the diversity and composition
al., 2015; Pulido-Herrera & Ramos-Montaño, 2016; Ramírez- of lichen communities depend on macroecological features
Morán et al., 2016; Simijaca et al., 2018). driven by altitude and other factors. Moncada et al. (2014)
Colombia served as the target region for one of the demonstrated that (sub-)Andean Forests and Páramo areas
most important broad-scale ecological studies on tropical harbour the highest richness of Sticta species, with the
ecosystems, the ECOANDES project (Van der Hammen Eastern Cordillera as one of the centres of diversity for the
et al., 1983). ECOANDES was an extraordinary example genus. Soto-Medina et al. (2012) found subtle phorophyte
of collaborative work between Colombian and Dutch preferences of lichens, with a stronger dependency on
researchers during the early 1980s (see Chapter 2), leaving a microclimatic factors. Chilito-López et al. (2016) also found
lasting impact in the development of Colombian biodiversity pronounced microhabitat preferences among lichenised
research and resulting in various studies on lichenised fungi taxa correlated with functional parameters, such as growth
(Sipman, 1986, 1989, 1998; Kessler, 2000; Aguirre-C., form. Unfortunately, a comprehensive database in terms of
2008). The study area encompassed all four major mountain functional parameters does not yet exist for the lichens of
systems of Colombia: the Sierra Nevada de Santa Marta, Colombia or tropical lichens in general, but this would be
the Western Cordillera (Tatamá), the Central Cordillera (Los a desirable achievement for the near future to allow more
Nevados), and the Eastern Cordillera (Sumapaz). Besides thorough assessments in trait-based functional ecosystem
more inventory-oriented lichen studies, the ECOANDES analyses (St. Martin & Mallik 2017).
project also produced the seminal dissertation by Jan Wolf Impact studies suggest that the replacement of
on epiphytic canopy communities on Colombian montane natural forests with tree monocultures natural forests
forests, including numerous new data on lichens (Wolf, influence influences lichen communities. Thus, Eucalyptus
1993a-c, 1994, 1995). Starting with the ECOANDES project, plantations in Colombian montane Quercus forests have a
Harrie Sipman became a pioneer in modern lichen research strong dediversification and homogenisation effect, with
in Colombia and a mentor of Jaime Aguirre-Ceballos, who planted trees featuring highly similar lichen communities,
continued the studies on the ecogeography of Colombian leading to low plot-level species richness (Ardila-Rios et al.,
lichens (Aguirre-C. & Rangel, 2008). 2015). Notably, replacing native Quercus forests with Pinus
plantations does not seem to affect diversity metrics but
causes partial species turnover (Simijaca et al., 2018). Pulido-
Herrera & Ramos-Montaño (2016) found a marked edge effect
on lichen community parameters and chlorophyll content
in fragments of Polylepis forests in the páramo zone, and
Ramírez-Morán et al. (2016) associated functional biotypes
with the conservation status of Andean Forest remnants.
most of these studies were based on lichen community topic have been performed in Colombia (Perico-Franco et al.,
ecology parameters, employing the Index of Atmospheric 2015; Valencia-Islas et al., 2020). Unfortunately, most studies
Purity (IAP), which links the frequency and abundance of do not go beyond reporting the results from standardised
individual taxa to sample species composition, providing bioassays, and potential clinical applications are rarely
a taxon-independent measure for air quality. The IAP was explored (Lücking, 2020). The causes of antibacterial,
also used to determine isocontamination zones in various antitumoral or antioxidative effects are often unclear,
areas of Bogotá DC (Rubiano-Olaya, 2002; Rubiano-Olaya & because the bioactive compounds are not always identified,
Chaparro, 2006; González-Aldana, 2007), Tunja (Simijaca- or substances are tested separately, without considering
Salcedo et al., 2011, 2014), and Quibdó (Mena, 2012), potential interactions with other substances. Another issue
particularly in relation to vehicular traffic (Valois-Cuesta & is inaccurate taxonomy, as experts are rarely consulted,
Mosquera-Palacios, 2014). In Medellín, an updated study and the underlying material is often not documented by
confirmed the usefulness of the IAP across a broader proper voucher material. Fortunately, the aforementioned
range of phorophytes, using air quality monitoring stations Colombian studies are exemplary in this respect, as they
as reference (Correa-Ochoa et al., 2020), and the method cited voucher material and consulted taxonomic experts.
was also employed to assess air quality in suburban areas Although promising, applied approaches using lichens
near Medellín (Quijano-Abril et al., 2021). While broadly depend on accurate assessment of the underlying biodiversity
comparable between regions, the downside of the IAP and hence require rigorous taxonomic studies and inventories
method is that it does not require complete identification using modern methods, including molecular assessments.
of the underlying lichens, as long as they are accurately Therefore, the political support for science should not
recognised as species. Therefore, most of the studies focus on applied aspects alone but should also provide the
mentioned above do not provide complete lists of fully framework for the necessary taxonomic inventories.
identified species, making a proper taxonomic assessment
of urban lichen diversity difficult to obtain. In addition to CONCLUSIONS
biomonitoring using the IAP, a few studies in Colombia have The currently known number of 2,678 species makes
assessed the impact of atmospheric pollutants, including the Colombian lichen funga one of the richest worldwide.
emissions in a sulphur mine in Cauca (Diaz-Escandón et However, the actual number may be almost twice as high,
al., 2016), acid rain (Álvarez-Berrio et al., 2018), and heavy requiring further rigorous inventory works and taxonomic
metals in lichens from Bogotá DC (Rodriguez et al., 2016). revisions, particularly in the understudied regions of the
The potential application of lichens to monitor the impact Caribe, Orinoquía, Pacífico and Amazonía. Unfortunately,
of land use changes are thus far from limited to montane societal and government support for such fundamental
forest ecosystems (Ardila-Rios et al., 2015; Diaz-Escandón studies in Colombia is limited. Instead, there is a strong
et al., 2016; Pulido-Herrera & Ramos-Montaño, 2016; focus on applied aspects, including in the training of young
Ramírez-Morán et al., 2016; Simijaca-Salcedo et al., 2018). students, neglecting the knowledge and tools that are
Unfortunately, standardised protocols for this purpose are necessary to properly assess the underlying taxonomy. Before
not yet available. Ramírez-Morán et al. (2016) assessed the undertaking applied studies, efforts should concentrate on
strategy of using easy-to-identify lichen biotypes instead of providing a complete inventory of the Colombian lichen biota
actual lichen taxa, providing an advantage like that of the so that potential applications can be assessed and explored
IAP. In Colombia, lichens, mosses, and vascular epiphytes more rigorously, taking into account the need for habitat
have been traditionally protected under the Resolución 213 conservation to preserve this diversity. In applied studies,
de 2013 of the Ministerio de Ambiente y Desarollo Sostenible the focus should be on broadly applicable, standardised
(MinAmbiente), initially aiming to prevent the uncontrolled approaches, such as using lichens as biomonitors of air
harvesting and trade of these organisms. For the past pollution and environmental health. Pharmaceutically
decade, this provision has been implemented to monitor the oriented studies should be more systematically structured
environmental impact of projects related to infrastructure to provide new insight into the potential of lichen secondary
or the exploration of mineral resources. Unfortunately, this metabolites, seeking collaborations for follow-up clinical
approach bears a few problems: sampling protocols are not studies when results are promising.
standardised, the underlying taxonomy is often inaccurate,
and there are conflicts of interest if the entity that carries
out the project is also responsible for the impact report. Acknowledgements
A solution could be a standardised protocol using selected
We are thankful to Paul Kirk and Brian Coppins for providing useful
lichen taxa that are known to be sensitive to environmental
data from Index Fungorum and the British Lichen Checklist and the
changes, such as thelotremoid Graphidaceae or lobarioid reviewer Viviana Motato-Vásquez for her suggestions.
Peltigeraceae (Rivas Plata et al., 2008; Ramírez-Morán et al.,
2016), combined with an assessment of the conservation
status of the underlying ecosystem (Etter et al., 2018). References
Another venue in terms of lichen uses is the diverse Aguirre-C J (1985) Flora de la Real expedición Botánica del Nuevo Reino de
composition of secondary substances and their potential Granada, 1783–1808. Tomo II: Algas, Líquenes, Hongos y Hepáticas.
applications (Mitrović et al., 2011). Some studies on this Madrid: Ediciones Cultura Hispánica. 83p.
Aguirre-C J (2008) Los líquenes de la región del Sumapaz (composición Oriental Colombiana, Transecto Sumapaz. Berlin: J. Cramer, Gebrüder
florística, distribución y ecología). In: Van der Hammen T (ed). La Borntraeger. Pp. 565-593.
Cordillera Oriental Colombiana. Transecto Sumapaz. Berlin: J. Cramer, Coppins BJ (2021) Lichen Taxon Dictionary. Available at: https://www.
Borntraeger. Pp. 211-234. britishlichensociety.org.uk/resources/lichen-taxon-database
Aguirre-C J, Rangel-Ch JO (2008) Riqueza y aspectos ecológicos y Correa-Ochoa MA, Vélez-Monsalve LC, Saldarriaga-Molina JC, Jaramillo-
fitogeográficos sobre la flora de líquenes. In: Rangel-Ch JO (ed). Ciro MM (2020) Evaluation of the Index of Atmospheric Purity in an
Colombia. Diversidad biótica VI. Riqueza y Diversidad de los Musgos y American tropical valley through the sampling of corticolous lichens
Líquenes de Colombia. Bogotá DC: Instituto de Ciencias Naturales, in different phorophyte species. Ecological Indicators, 115(10635),
Universidad Nacional de Colombia. Pp. 549-559. 1-11. doi: 10.1016/j.ecolind.2020.106355
Aguirre-C J, Sipman HJM (2004) Diversidad y riqueza de líquenes en el Diaz-Escandón DD, Soto-Medina E, Lücking R, Silverstone-Sopkin PA (2016)
Chocó biogeográfico. In: Rangel-Ch JO (ed). Colombia. Diversidad Corticolous lichens as environmental indicators of natural sulphur
Biótica IV. El Chocó Biogeográfico / Costa Pacífica. Bogotá DC: emissions near the sulphur mine El Vinagre (Cauca, Colombia). The
Instituto de Ciencias Naturales, Universidad Nacional de Colombia & Lichenologist, 48(2), 147-159. doi: 10.1017/S0024282915000535
Conservación Internacional. Pp. 455-474. Esslinger TL (2019) A cumulative checklist for the lichen-forming,
Ahmadjian V (1995) Lichens are more important than you think. BioScience, lichenicolous and allied fungi of the continental United States and
45(3), 124. Canada, version 23. Opuscula Philolichenum, 18, 102-378.
Ahti T (1992) Biogeographic aspects of Cladoniaceae in the páramos. In: Etter A, Andrade A, Saavedra K, Cortés J (2018) Actualización de la Lista
Balslev H, Luteyn JL (eds). Páramo: An Andean Ecosystem under Human Roja de los Ecosistemas Terrestres de Colombia: Herramienta para
Influence. London: Academic Press. Pp. 111-117. la gestión de los ecosistemas. In: Moreno LA, Ruedaand C, Andrade
Ahti T, Mayrhofer H, Schultz M, Tehler A, Fryday AM (2016) First supplement GI (eds). Biodiversidad 2017. Estado y Tendencias de la Biodiversidad
to the lichen checklist of South Africa. Bothalia 46(1), a2065. doi: continental de Colombia. Bogotá DC: Instituto de Investigación de
10.4102/abc.v46i1.2065 Recursos Biológicos Alexander von Humboldt. Pp. 18 (ficha 204).
Álvarez-Berrio JA, Buitrago-Guevara WA, Guzmán-Pinzón M (2018) Feuerer T, Hawksworth DL (2007) Biodiversity of lichens, including a
Determinación del comportamiento ácido de las precipitaciones en worldwide analysis of checklist data based on Takhtajan’s floristic
inmediaciones del Resguardo Indígena de la Serranía El Majuy en regions. Biodiversity and Conservation, 16(1), 85-98. doi: 10.1007/
Cota, Cundinamarca y sus efectos en líquenes de la zona. Luna Azul, s10531-006-9142-6
46, 70-105. doi: 10.17151/luaz.2018.46.6 Forman RT (1975) Canopy lichens with blue‐green algae: a nitrogen
Amrani S, Seaward MRD, Sipman HJM, Feuerer T (2018) Lichenological source in a Colombian rainforest. Ecology, 56(5), 1176-1184. doi:
exploration of Algeria II: checklist of lichenized, lichenicolous and allied 10.2307/1936157
fungi. Herzogia, 31(2), 817-892. doi: 10.13158/heia.31.2.2018.817 Friedl T, Büdel B (2008) Photobionts. In: Nash III TH (ed). Lichen Biology,
Aptroot A (2016) Preliminary checklist of the lichens of Madagascar, with 2nd edition. Cambridge: Cambridge University Press. Pp. 7-26.
two new thelotremoid Graphidaceae and 131 new records. Willdenowia, Fryday AM (2015) A new checklist of lichenised, lichenicolous and allied
46(3), 349-365. doi: 10.3372/wi.46.46304 fungi reported from South Africa. Bothalia, 45(1), 59-122. doi:
Aptroot A (2021) The four lichen floras of Brazil. International Association 10.4102/abc.v45i1.148
for Lichenology 9th Symposium (IAL 9). Brazil. IAL9 Program and Galloway DJ, Quilhot W (1998) Lista patron de los líquenes y hongos
Abstract Book. 53p. liquenícolas de Chile. Gayana Botanica, 55(2), 111-185.
Aptroot A, Lücking R, Sipman HJM, Umaña L, Chaves JL (2008) Gaya E, Vasco-Palacios AM, Vargas-Estupiñan N, Lücking R, Carretero
Pyrenocarpous lichens with bitunicate asci. A first assessment of the J, Sanjuan T, Moncada B, Allkin B, Bolaños-Rojas AC, Castellanos-
lichen biodiversity inventory in Costa Rica. Bibliotheca Lichenologica, Castro C, Coca LF, Corrales A, Cossu T, Davis L, dSouza J, Dufat A,
97, 1-162. Franco-Molano AE, Garcia F, Gómez-Montoya N, González-Cuellar FE,
Arbeláez-Cortés E (2013) Knowledge of Colombian biodiversity: published Hammond D, Herrera A, Jaramillo-Ciro MM, Lasso-Benavides C, Mira
and indexed. Biodiversity and Conservation, 22(12), 2875-2906. doi: MP, Morley J, Motato-Vásquez V, Niño-Fernandez Y, Ortiz-Moreno ML,
10.1007/s10531-013-0560-y Peña-Cañón ER, Ramirez-Castrillón M, Rojas T, Ruff J, Simijaca D,
Ardila-Rios AI, Moncada B, Lücking R (2015) Epiphyte homogenisation Sipman HJM, Soto-Medina E, Torres G, Torres-Andrade PA, Ulian T,
and de-diversification on alien Eucalyptus versus native Quercus forest White K, Diazgranados M (2021) ColFungi: Colombian Resources for
in the Colombian Andes: a case study using lirellate Graphidaceae Fungi Made Accessible. Richmond: Royal Botanic Gardens, Kew. 26p.
lichens. Biodiversity and Conservation, 24(5), 1239-1252. doi: Gonzalez-Aldana WY (2007) Hongos liquenizados como bioindicadores:
10.1007/s10531-014-0855-7 macrozonas de isocontaminación atmosférica de Bogotá D.C.
Buaruang K, Boonpragob K, Mongkolsuk P, Sangvichien E, Vongshewarat (Cundinamarca, Colombia). Pérez Arbelaezia, 17, 137-154.
K, Polyiam W, Rangsiruji A, Saipunkaew W, Naksuwankul K, Goward T (2008) Nameless little things. Evansia, 25, 54-56.
Kalb J, Parnmen S, Kraichak E, Phraphuchamnong P, Meesim S, Grimm M, Grube M, Schiefelbein U, Zühlke D, Bernhardt J, Riedel K (2021)
Luangsuphabool T, Nirongbut P, Poengsungnoen V, Duangphui N, The lichens’ microbiota, still a mystery? Frontiers in Microbiology,
Sodamuk M, Phokaeo S, Molsil M, Aptroot A, Kalb K, Lücking R, 12, 714.
Lumbsch HT (2017) A new checklist of lichenized fungi occurring in Grube M, Cernava T, Soh J, Fuchs S, Aschenbrenner I, Lassek C, Wegner
Thailand. MycoKeys, 23, 1-91. doi: 10.3897/mycokeys.23.12666 U, Becher D, Riedel K, Sensen CW, Berg G (2015) Exploring functional
Butler R (2016) The Top 10 Most Biodiverse Countries. Available at: contexts of symbiotic sustain within lichen-associated bacteria by
https://news.mongabay.com/2016/05/top-10-biodiverse-countries comparative omics. The ISME Journal, 9, 412-424. doi: 10.1038/
Butler R (2019) Countries with the Highest Biodiversity. Available at: ismej.2014.138
https://rainforests.mongabay.com/03highest_biodiversity.htm Hafellner J, Türk R (2016) Die lichenisierten Pilze Österreichs – eine
Cadena-Castañeda OJ (2013) The tribe Dysoniini part II: the genus neue Checkliste der bisher nachgewiesenen Taxa mit Angaben zu
Markia (Orthoptera: Tettigoniidae; Phaneropterinae), new species and Verbreitung und Substratökologie. Stapfia, 104(1), 1-216.
some clarifications. Zootaxa, 3599(6), 501-518. doi: 10.11646/ Hawksworth DL, Grube M (2020) Lichens redefined as complex
zootaxa.3599.6.1 ecosystems. New Phytologist, 227, 1281-1283. doi: 10.1111/
Calvelo S, Liberatore S (2002) Catálogo de los líquenes de la Argentina. nph.16630
Kurtziana, 29(2), 7-170. Hernández-M JA (2021) A new look at Venezuelan lichen diversity.
Chilito-López LG, Soto-Medina EA, Peña AM (2016) Effects of microclimate International Association for Lichenology 9th Symposium (IAL 9),
on species diversity and functional traits of corticolous lichens in the August 1 to 6, 2021, Brazil (online). IAL9 Program and Abstract Book,
Popayan Botanical Garden (Cauca, Colombia). Cryptogamie Mycologie, 77.
37(2), 205-215. doi: 10.7872/crym/v37.iss2.2016.205 Herrera-Campos MA, Lücking R, Pérez-Pérez RE, Miranda-González R,
Cleef AM (2008) Humid cloud superparamo probably acts as a plant Sánchez N, Barcenas-Peña A, Carrizosa A, Zambrano A, Ryan BD,
diversity centre and as a cool refuge: the case of Nevado de Nash III TH (2014) Biodiversidad de líquenes en México. Revista
Sumapaz, Colombia. In: Van der Hammen T (ed). La Cordillera Mexicana de Biodiversidad, 85, S82-S99. doi: 10.7550/rmb.37003
Honegger R (2012) The symbiotic phenotype of lichen-forming Ascomycetes Marcelli MP (1998) History and current knowledge of Brazilian lichenology. In:
and their endo- and epibionts. In: Hock B (ed). Fungal Associations. Marcelli MP, Seaward MRD (eds). Lichenology in Latin America: History,
Berlin: Springer. Pp. 287-339. Current Knowledge and Applications. São Paulo: CETESB. Pp. 25-45.
Hooker WJ (1822) Lichenes, Achar. In: Kunth CS (ed). Synopsis Plantarum, Mateus N, Aguirre-C J, Lücking R (2012) Contribuciones a la biota liquénica
quas in itinere ad plagam aequinoctatem orbis novi, collegerunt Al. de foliícola del Chocó (Colombia). Caldasia, 34(1), 25-32.
Humboldt et Am. Bonpland, Tomo 1. Paris: Levrault. Pp. 14-39. McCarthy PM (2020) Checklist of the Lichens of Australia and its Island
IDEAM — Instituto de Hidrología, Meteorología y Estudios Ambientales, Territories. Australian Biological Resources Study, Canberra. Version 1
Instituto de Investigación de Recursos Biológicos Alexander von March 2020. Available at: http://www.anbg.gov.au/abrs/lichenlist/
Humboldt (Instituto Humboldt), Instituto de Investigaciones Marinas introduction.html
y Costeras José Benito Vives de Andréis (Invemar) and Instituto Mena R (2012) Calidad del aire en Quibdó mediante bioindicadores.
Geográfico Agustín Codazzi (IGAC) (2017) Memoria Técnica. Mapa Bioetnia, 9(2), 215-227. doi: 10.51641/bioetnia.v9i2.92
de Ecosistemas Continentales, Costeros y Marinos de Colombia (MEC), Mercado-Diaz JA, Santiago-Valentin E (2010) Lichenological studies in
Escala 1:100.000. 170p. Puerto Rico: history and current status. Harvard Papers in Botany,
Kessler M (2000) Altitudinal zonation of Andean cryptogam communities. 15(1), 93-101.
Journal of Biogeography, 27(2), 275-282. Mia˛dlikowska J, Richardson D, Magain N, Ball B, Anderson F, Cameron
Lendemer JC, Keepers KG, Tripp EA, Pogoda CS, McCain CM, Kane NC R, Lendemer J, Truong C, Lutzoni F (2014) Phylogenetic placement,
(2019) A taxonomically broad metagenomic survey of 339 species species delimitation, and cyanobiont identity of endangered aquatic
spanning 57 families suggests cystobasidiomycete yeasts are not Peltigera species (lichen‐forming Ascomycota, Lecanoromycetes).
ubiquitous across all lichens. American Journal of Botany, 106, 1090- American Journal of Botany, 101(7), 1141-1156. doi: 10.3732/
1095. doi: 10.1002/ajb2.1339 ajb.1400267
Lisi O, Daza A, Londoño R, Quiroga S, Pilato G (2019) Meplitumenaluna gen. Mitrović T, Stamenković S, Cvetković V, Nikolić M, Tošić S, Stojičić D
nov., sp. nov. an interesting eutardigrade (Hypsibiidae, Itaquasconinae) (2011) Lichens as source of versatile bioactive compounds. Biologica
from the Sierra Nevada de Santa Marta, Colombia. ZooKeys, 865, Nyssana, 2(1), 1-6.
1-20. doi: 10.3897/zookeys.865.30705 Mittermeier RA (1988) Primate diversity and the tropical forest. In: Wilson
Londoño R, Daza A, Lisi O, Quiroga S (2017) New species of waterbear EO (ed). Biodiversity. Washington DC: National Academic Press. Pp.
Minibiotus pentannulatus (Tardigrada: Macrobiotidae) from Colombia. 145-154.
Revista Mexicana de Biodiversidad, 88(4), 807-814. doi: 10.1016/j. Mittermeier RA, Mittermeier CG, Gil PR (1999) Megadiversity: Earth’s
rmb.2017.10.040 biologically wealthiest nations. Chelonian Conservation and Biology,
Lücking R (2008) Foliicolous lichenized fungi. Flora Neotropica Monograph, 3(3), 537.
103, 1–867. Moncada B, Aguirre J, Lücking R (2014) Ecogeografía del género Sticta
Lücking R (2020) Three challenges to contemporaneous taxonomy from a (Ascomycota liquenizados: Lobariaceae) en Colombia. Revista de
licheno-mycological perspective. Megataxa, 1, 78-103. doi: 10.11646/ Biología Tropical, 62(1), 266-281.
MEGATAXA.1.1.16 Moncada B, Sipman HJM, Lücking R (2020) Testing DNA barcoding in
Lücking R, Aptroot A, Chaves JL, Sipman HJM, Umaña-Tenorio L (2007) A Usnea (Parmeliaceae) in Colombia using the internal transcribed
first assessment of the Ticolichen biodiversity inventory in Costa Rica: spacer (ITS). Plant and Fungal Systematics, 65, 358-385. doi:
the genus Coccocarpia (Peltigerales: Coccocarpiaceae). Bibliotheca 10.35535/pfsyst-2020-0026
Lichenologica, 95, 429-457. Motta J, Amórtegui K (2018) Contribución al Género Graphis
Lücking R, Chaves JL, Sipman HJM, Umaña-Tenorio L, Aptroot A (2008) A (Graphidaceae) en Colombia, con Base en las Colecciones del
first assessment of the Ticolichen Biodiversity Inventory in Costa Rica: Herbario Forestal Gilberto Emilio Mahecha Vega (UDBC) Sección
the genus Graphis, with notes on the genus Hemithecium (Ascomycota: Criptógamas de la Universidad Distrital Francisco José de Caldas.
Ostropales: Graphidaceae). Fieldiana (Botany), New Series, 46, 1-131. Undergraduate thesis. Universidad Distrital Francisco José de
Lücking R, Rivas Plata E, Chaves JL, Umaña L, Sipman HJM (2009) How Caldas, Bogotá, Colombia.
many tropical lichens are there... really? Bibliotheca Lichenologica, Motta K, Amórtegui K, Moncada B, Lücking R (2019) New species in the
100, 399-418. genus Graphis with transversally septate ascospores (Ascomycota:
Lücking R, Seavey F, Common RS, Beeching SQ, Breuss O, Buck WR, Ostropales: Graphidaceae) from Colombia. Phytotaxa, 401(4), 257-
Crane L, Hodges M, Hodkinson BP, Lay E, Lendemer JC, McMullin RT, 266. doi: 10.11646/phytotaxa.401.4.3
Mercado-Díaz JA, Nelsen MP, Rivas Plata E, Safranek W, Sanders WB, Murcia C, Kattan GH, Andrade‐Pérez GI (2013) Conserving biodiversity
Schaefer HP Jr, Seavey J (2011) The lichens of Fakahatchee Strand in a complex biological and social setting: The case of Colombia.
Preserve State Park, Florida: Proceedings from the 18th Tuckerman In: Raven PH, Sodhi NS, Gibson L (eds). Conservation Biology: Voices
Workshop. Bulletin of the Florida Museum of Natural History, Biological from the Tropics. Oxford: Wiley Blackwell. Pp. 86-96.
Sciences, 49, 127-186. Nelsen MP, Lücking R, Chaves JL, Sipman HJM, Umaña L, Navarro E (2006)
Lücking R, Hodkinson BP, Leavitt SD (2017) The 2016 classification of A first assessment of the Ticolichen biodiversity inventory in Costa
lichenised fungi in the Ascomycota and Basidiomycota - Approaching one Rica: the genus Haematomma (Lecanorales: Haematommataceae).
thousand genera. The Bryologist, 119, 361-416. doi: 10.1639/0007- The Lichenologist, 38, 251-262. doi: 10.1017/S0024282906005573
2745-119.4.361 Nimis PL (2016) The lichens of Italy. A Second Annotated Catalogue.
Lücking R, Moncada B, Martínez-Habibe MC, Salgado-N BE, Celis M, Trieste: EUT Edizioni Università di Trieste. 740p.
Rojas-Zamora O, Rodríguez-M GM, Brokamp G, Borsch T (2019) Lichen Nowak R, Winkler S (1970) Foliicole Flechten der Sierra Nevada de
diversity in Colombian Caribbean dry forest remnants. Caldasia, 41, Santa Marta (Kolumbien) und ihre gegenseitigen Beziehungen.
194-214. doi: 10.15446/caldasia.v41n1.71060 Österreichische Botanische Zeitschrift, 118, 456-485.
Lücking R, Leavitt SD, Hawksworth DL (2021) Species in lichen‑forming Nowak R, Winkler S (1975) Foliicolous lichens of Choco, Colombia, and
fungi: balancing between conceptual and practical considerations, their substrate abundances. The Lichenologist, 7(1), 53-58.
and between phenotype and phylogenomics. Fungal Diversity, 109, 99- Nylander W (1859) Lichenes Exotici, Lichenes in regionibus exoticis
154. doi: 10.1007/s13225-021-00477-7 quibusdam vigentes exponit synopticis enumerationibus. Annales des
Madriñán S, Cortés AJ, Richardson JE (2013) Páramo is the world’s fastest Sciences Naturelles, 11, 205-264.
evolving and coolest biodiversity hotspot. Frontiers in Genetics, 4, 192. Nylander W (1863a) Lichenographiae Novo-Granatensis Prodromus. Acta
doi: 10.3389/fgene.2013.00192 Societatis Scienciarum Fennicae, 7, 415-504.
Mägdefrau K, Winkler S (1967) Lepidostroma terricolens n. g. n. sp., eine Nylander W (1863b) Lichenes. In: Triana J, Planchon JE (eds). Prodromus
Basidiolichene der Sierra Nevada de Santa Marta (Kolumbien). Mitteilungen Florae Novo-Granatensis ou Énumération des plantes de la Nouvelle-
aus dem Instituto Colombo-Aleman de Investigaciones Cientificas, 1, 11-17. Grénade avec description des espèces Nouvelles. Annales des
Marcano V, Morales-Méndez A, Sipman H, Calderon L (1996) A first Sciences Naturelles, 19, 286-382.
checklist of the lichen-forming fungi of the Venezuelan Andes. Tropical Nylander W (1863c) Lichenes. In: Triana J, Planchon JE (eds). Prodromus
Bryology, 12, 193-235. doi: 10.11646/bde.12.1.19 Florae Novo-Granatensis ou Énumération des plantes de la Nouvelle-
Grénade avec description des espèces nouvelles. Annales des Rivera J, Yagui H, Ehrmann R (2011) Mantids in the mist – taxonomy of
Sciences Naturelles, 20, 228-279. the Andean genus Pseudopogonogaster Beier, 1942, a cloud forest
Nylander W (1864) Circa lichenum Novo-Granatensium novas specialist, with notes on its biogeography and ecology (Mantodea:
explorationes Lindigianas. Flora, 47, 617-620. Espidae: Miopteryginae). Insect Systematics & Evolution, 42(4), 313-
Nylander W (1867) Lichenes, additamentum. In: Triana J, Planchon JE 335. doi: 10.1163/187631211X595056
(eds). Prodromus Florae Novo-Granatensis. Annales des Sciences Rodríguez-A OE, Andrade BW, Díaz LF, Celis C, Ortiz-Ardila A (2016)
Naturelles, 7, 301-354. Quantification of heavy metals in lichens from the upper basin of the
Ohmura Y, Kashiwadani H (2018) Checklist of Lichens and Allied Fungi of river Bogotá. Pharmacology OnLine, 2016(2), 21-27.
Japan. Tokyo: National Museum of Nature and Science. 143p. Rodriguez de Flakus P, Kukwa M, Etayo J, Lücking R, Meneses RI, Plata ER,
Osorio HS (1972) Contribution to the lichen flora of Uruguay. VII. A Stanton D, Truong C, Vargas R, Flakus A (2016) Preliminary Catalogue
preliminary catalogue. Comunicaciones Botánicas del Museo de of Lichens and Lichenicolous Fungi from Bolivia. Available at: https://
Historia Natural de Montevideo, 4(56), 1-46. bio.botany.pl/lichens-bolivia/en,strona,catalogue,5.htm
Osorio HS (1992) Contribucion a la flora liquénica del Uruguay. XXV. Roux C (2012) Liste des lichens et champignons lichénicoles de France.
Líquenes publicados entre 1972 a 1991. Anales del Museo Nacional Bulletin de la Société linnéenne de Provence, Numéro spécial 16, 1-220.
de Historia Natural de Montevideo, Series 2, 8, 43-70. Roux C, Coste C, Bricaud O, Bauvet C, Masson D (2008) Lichens et
Paguirigan JAG, de la Cruz TEE, Santiago KAA, Gerlach A, Aptroot A champignons lichénicoles du parc national des Cévennes (France). 5
(2020) A checklist of lichens known from the Philippines. Current – Vue d’ensemble et conclusion. Bulletin de la Société Linnéenne du
Research in Environmental & Applied Mycology, 10(1), 319-376. doi: Provence, 59, 243-279.
10.5943/cream/10/1/29 Rubiano-Olaya LJ (1987) Delimitación de áreas de isocontaminación en
Pérez-Quintero ÁL, Watteijne-Cerón B (2009) Lichen community Cali y Medellín utilizando líquenes como indicadores. Pérez Arbelaezia,
structure and morphological changes in the genus Sticta (Stictaceae) 1(4–5), 7-41.
associated to an altitude gradient. Acta Biológica Colombiana, 14(3), Rubiano-Olaya LJ (1988) Líquenes como indicadores de contaminación
159-172. en el complejo industrial de Betania y la termoeléctrica de Zipaquirá,
Perico-Franco LS, Soriano-García M, Cerbón MA, González-Sánchez Cundinamarca. Acta Biológica Colombiana, 1(4), 65-125.
I, Valencia-Islas NA (2015) Secondary metabolites and cytotoxic Rubiano-Olaya LJ (2002) Monitoreo de áreas de isocontaminación en la
potential of Lobariella pallida and Stereocaulon strictum var. región de in-fluencia de la Central Termoeléctrica Martín del Corral
compressum, two lichens from Colombian páramo region. UK Journal utilizando líquenes como bíoindicadores. Perez Arbelaezia, 13, 91-104.
of Pharmaceutical and Biosciences, 4, 9-17. doi: 10.20510/ukjpb/3/ Rubiano-Olaya LJ, Chaparro M (2006) Delimitación de áreas de
i4/89463 isocontaminación atmosférica en el campus de la Universidad
Perlmutter GB, Weakley AS (2018) USDA Plants Database Lichen Nacional de Colombia mediante el análisis de bioindicadores (líquenes
Update, Phase II. NRCS - East National Technology Support Center, epifitos). Acta Biológica Colombiana, 11(2), 87-102.
Greensboro, NC. Saini KC, Nayaka S, Bast F (2019) Diversity of lichen photobionts: their
Piepenbring M (2007) Inventorying the fungi of Panama. Biodiversity and coevolution and bioprospecting potential. In: Satyanarayana T, Das
Conservation, 16(1), 73-84. doi: 10.1007/s10531-006-9051-8 SK, Johri BN (eds). Microbial Diversity in Ecosystem Sustainability
Pinzón M, Linares EL (2006) Diversidad de líquenes y briofitos en la region and Biotechnological Applications. Volume 2: Soil & Agroecosystems.
subxerofitica de La Herrera, Mosquera (Cundinamarca-Colombia). I. Singapore: Springer Nature. Pp. 307-323.
Riqueza y estructura. Caldasia, 28(2), 243-257. Sanders WB, Moe RL, Ascaso C (2004) The intertidal marine lichen formed
Pulido-Herrera K, Ramos-Montaño C (2016) Efecto de borde en la by the pyrenomycete fungus Verrucaria tavaresiae (Ascomycotina)
distribución de líquenes y el contenido de clorofilas en fragmentos and the brown alga Petroderma maculiforme (Phaeophyceae): thallus
de Polylepis quadrijuga (Rosaceae) en el páramo de La Rusia (Boyacá- organisation and symbiont interaction. American Journal of Botany,
Colombia). Revista de Biología Tropical, 64(4), 1683-1697. doi: 91(4), 511-522. doi: 10.3732/ajb.91.4.511
10.15517/rbt.v64i4. 22735 Seaward MRD (1977) Lichen Ecology. London: Academic Press. 550p.
Quijano-Abril MA, Ramírez-Ospina DM, Domiguez-Rave MI, Londoño- Seaward MRD (1988) Contribution of lichens to ecosystems. In: Galun M
Valencia J (2021) Líquenes como biosensores para la evaluación (ed). CRC Handbook of Lichenology. Volume II. Boca Raton: CRC Press.
de contaminación atmosférica urbana y sub urbana en un valle de Pp. 107-129.
montaña tropical, Rionegro, Antioquia. Bionatura, 6(1), 1501-1509. Sierra MA, Danko DC, Sandoval TA, Pishchany G, Moncada B, Kolter R,
doi: 10.21931/RB/2021.06.01.10 Mason CE, Zambrano MM (2020) The microbiomes of seven lichen
Ramírez-Morán, N.A., León-Gómez, M. and Lücking, R. (2016). Uso genera reveal host specificity, a reduced core community and potential
de biotipos de líquenes como bioindicadores de perturbación en as source of antimicrobials. Frontiers in Microbiology, 11, 398. doi:
fragmentos de bosque altoandino (Reserva Biológica “Encenillo”, 10.3389/fmicb.2020. 00398
Colombia). Caldasia, 38(1), 31-52. doi: 10.15446/caldasia. Simijaca-Salcedo DF, Morales-Puentes ME, Díaz-Pérez CN (2011)
v38n1.57821 Líquenes y contaminación atmosférica en la Universidad Pedagógica
Ramírez-Roncallo KJ (2018) Líquenes de la vertiente noroccidental de y Tecnológica de Colombia, Tunja, Boyacá, Colombia. Ciencia en
la Sierra Nevada de Santa Marta en Colombia. Bachelor’s thesis. Desarollo, 3(2), 69-88.
Universidad del Magdalena, Santa Marta, Colombia. Simijaca-Salcedo DF, Vargas-Rojas DL, Morales-Puentes ME (2014)
Ramos D (2014) Lista de especies de líquenes y hongos liquenícolas del Uso de organismos vegetales no vasculares como indicadores
Perú. Glalia, 6(2), 1-49. de contaminación atmosférica urbana (Tunja, Boyacá, Colombia).
Rangel-Ch JO (2015) La biodiversidad de Colombia: significado y Acta Biologica Colombiana, 19(2), 221-231. doi: 10.15446/abc.
distribución regional. Revista de la Academia Colombiana de Ciencias v19n2.40681
Exactas, Físicas y Naturales, 39(151), 176-200. doi: 10.18257/ Simijaca-Salcedo DF, Moncada B, Lücking R (2018) Bosque de roble o
raccefyn.136 plantación de coníferas,¿ qué prefieren los líquenes epífitos? Colombia
Rincón-Espitia A, Aguirre-C J, Lücking R (2011) Líquenes corticícolas en Forestal, 21(2), 123-141. doi: 10.14483/2256201x.12575
el Caribe Colombiano. Caldasia, 33, 311-327. Singh KP, Sinha GP (2010) Indian Lichens: An Annotated Checklist. Kolkata:
Rivas Plata E, Lücking R, Aptroot A, Sipman HJM, Umaña L, Chaves JL, Botanical Survey of India. 572p.
Lizano D (2006) A first assessment of the Ticolichen biodiversity Sinha GP, Nayaka S, Joseph S (2018) Additions to the checklist of Indian
inventory in Costa Rica: the genus Coenogonium (Ostropales: lichens after 2010. Cryptogam Biodiversity and Assessment, Special
Coenogoniaceae), with a worldwide key and checklist and a phenotype- Volume, 2018, 197-206. doi: 10.21756/cab.esp16
based cladistic analysis. Fungal Diversity, 23, 255-321. Sipman HJM (1986) Liquenes del transecto Buritaca-La Cumbre, Sierra
Rivas Plata E, Lücking R, Lumbsch HT (2008) When family matters: an Nevada de Santa Marta, Colombia. In: Van der Hammen T, Ruiz
analysis of Thelotremataceae (Lichenized Ascomycota: Ostropales) as PM (eds). La Sierra Nevada de Santa Marta (Colombia). Transecto
bioindicators of ecological continuity in tropical forests. Biodiversity Buritaca–La Cumbre. Berlin: J. Cramer, Gebrüder Borntraeger. Pp.
and Conservation, 17, 1319-1351. doi: 10.1007/s10531-007-9289-9 185-188.
Sipman HJM (1989) Lichen zonation in the Parque Los Nevados transect. lichen from the Andean paramo. Pharmaceutical Sciences, 27(2),
In: Van der Hammen T, Diaz-Piedrahita S, Alvarez VJ (eds). La 281-290. doi: 10.34172/PS.2020. 83
Cordillera Central Colombiana Transecto Parque los Nevados (Segunda Valois-Cuesta H, Mosquera-Palacios Y (2014) Líquenes como
Parte). Berlin: J. Cramer, Gebrüder Borntraeger. Pp. 461-483. bioindicadores de la calidad del aire en la cuidad de Quibdó,
Sipman HJM (1998) Los líquenes del transecto Tatamá. In: Van der Chocó, Colombia. Revista Biodiversidad Neotropical, 4(1), 7-15. doi:
Hammen T, Rangel-Ch JO, Cleef AM (eds). La Cordillera Occidental 10.18636/bioneotropical.v4i1.178
Colombiana. Transecto Tatamá. Berlin: J. Cramer, Gebrüder Van der Hammen T, Pérez-Preciado A, Pinto P (1983) La Cordillera Central
Borntraeger. Pp. 267-273. Colombiana, Transecto Parque Los Nevados (Introduccion y datos
Sipman HJM (2002) The significance of the northern Andes for lichens. iniciales). Vaduz: Cramer. 600p.
The Botanical Review, 68(1), 88-99. Vargas-Castillo R, Sandoval-Leiva P (2020) Lista Sistemática de los
Sipman HJM, Aguirre-C J (2016) Líquenes. In: Bernal R, Gradstein SR, Hongos Liquenizados y Liquenícolas Presentes en Chile. Version 1.6.
Celis M (eds). Catálogo de Plantas y Líquenes de Colombia. Volumen 1. Universidad Metropolitana de Ciencias de la Educación. Checklist
Bogotá DC: Universidad Nacional de Colombia, Facultad de Ciencias, Dataset. doi: 10.15468/jxynx2
Instituto de Ciencias Naturales. Pp. 159-281. Veneklaas EJ (1991) Litterfall and nutrient fluxes in two montane tropical
Sipman HJM, Harris RC (1989) Lichens. In: Lieth H, Werger MJA (eds). rain forests, Colombia. Journal of Tropical Ecology, 7(3), 319-336.
Tropical Rain Forest Ecosystems: Biogeographical and Ecological doi: 10.1017/ S0266467400005587
Studies. Amsterdam: Elsevier. Pp. 303-309. Watkinson SC, Boddy L, Money NP (2016) The Fungi. 3rd edition. Oxford:
Sipman HJM, Hekking W, Aguirre-C J (2008) Checklist of Lichenized and Academic Press, Elsevier. 496p.
Lichenicolous Fungi from Colombia. Bogotá DC: Instituto de Ciencias Wei JC (2021) The Enumeration of Lichenized Fungi in China. Beijing:
Naturales, Universidad Nacional de Colombia. 242p. China Forestry Publishing House. 606p.
Sipman HJM, Lücking R, Aptroot A, Chaves JL, Kalb K, Umaña-Tenorio Wirth V, Hauck M, Brackel WV, Cezanne R, de Bruyn U, Dürhammer O,
L (2012) A first assessment of the Ticolichen biodiversity inventory Eichler M, Gnüchtel A, Litterski B, Otte V, Schiefelbein U (2011)
in Costa Rica and adjacent areas: the thelotremoid Graphidaceae Checklist of Lichens and Lichenicolous Fungi in Germany, Version 2.
(Ascomycota: Ostropales). Phytotaxa, 55, 1-214. Göttingen: Georg August University.
Smith CW, Aptroot A, Coppins BJ, Fletcher A, Gilbert OL, James PW, Wolf JHD (1993a) Diversity patterns and biomass of epiphytic bryophytes
Wolseley PA (2009) The Lichens of Great Britain and Ireland. London: and lichens along an altitudinal gradient in the northern Andes.
British Lichen Society. 1046p. Annals of the Missouri Botanical Garden, 80(4), 928-960. doi:
Soto-Medina E, Lücking R, Bolaños-Rojas A (2012) Especificidad de 10.2307/2399938
forófito y preferencias microambientales de los líquenes cortícolas Wolf JHD (1993b) Epiphyte communities of tropical montane rain
en cinco forófitos del bosque premontano de finca Zíngara, Cali, forests in the northern Andes I. Lower montane communities.
Colombia. Revista de Biología Tropical, 60(2), 843-856. Phytocoenologia, 22(1), 1-52.
Soto-Medina E, Londoño-Lemos V, Díaz-Escandón D (2015) Epiphytes Wolf JHD (1993c) Epiphyte communities of tropical montane rain forests in
from a forest type transition zone in the Choco biogeographic region, the northern Andes II. Upper montane communities. Phytocoenologia,
Valle del Cauca, Colombia. Revista de Biología Tropical, 63(4), 915- 22(1), 53-103.
926. doi: 10.15517/rbt.v63i4.16733 Wolf JHD (1994) Factors controlling the distribution of vascular and non-
Soto-Medina E, Lücking R, Silverstone-Sopkin PA, Torres AM (2019) vascular epiphytes in the northern Andes. Vegetatio, 112(1), 15-28.
Changes in functional and taxonomic diversity and composition Wolf JHD (1995) Non-vascular epiphyte diversity patterns in the canopy
of corticolous lichens in an altitudinal gradient in Colombia. of an upper montane rain forest (2,550-3,670 m), central cordillera,
Cryptogamie Mycologie, 40(6), 97-115. doi: 10.5252/cryptogamie- Colombia. Selbyana, 16(2), 185-195.
mycologie2019v40a6 Zárate-Arias N, Moreno-Palacios M, Torres-Benítez A (2019) Diversidad,
Soto-Medina E, Castaño-Naranjo A, Granobles J, Aptroot A (2021) especificidad de forófito y preferencias microambientales de líquenes
Una nueva especie y nuevos registros de líquenes en Colombia y cortícolas de un bosque subandino en la región Centro de Colombia.
Suramérica en el bosque seco tropical de la cuenca media del río Revista de la Academia Colombiana de Ciencias Exactas, Físicas y
Cauca. Revista de la Academia Colombiana de Ciencias Exactas, Naturales, 43(169), 737-745. doi: 10.18257/raccefyn.886
Físicas y Naturales, 45(175), 415-420. doi: 10.18257/raccefyn.1322 Zedda L, Rambold G (2015) The diversity of lichenised fungi: ecosystem
Spribille T, Pérez-Ortega S, Tønsberg T, Schirokauer D (2010) Lichens and functions and ecosystem services. In: Upreti DK, Divakar PK, Shukla
lichenicolous fungi of the Klondike Gold Rush National Historic Park, V, Bajpai R (eds). Recent Advances in Lichenology Modern Methods
Alaska, in a global biodiversity context. The Bryologist, 113, 439- and Approaches in Lichen Systematics and Culture Techniques, Volume
515. doi: 10.1639/0007-2745-113.3.439 2. New Delhi: Springer. Pp. 121-145.
Spribille T, Tuovinen V, Resl P, Vanderpool D, Wolinski H, Aime MC,
Schneider K, Stabentheiner E, Toome-Heller M, Thor G, Mayrhofer
H (2016) Basidiomycete yeasts in the cortex of ascomycete
macrolichens. Science, 353, 488-492. doi: 10.1126/science.
aaf8287
Spribille T, Fryday AM, Pérez-Ortega S, Svensson M, Tønsberg T, Ekman
S, Holien H, Resl P, Schneider K, Stabentheiner E, Thüs H (2020)
Lichens and associated fungi from Glacier Bay National Park, Alaska.
The Lichenologist, 52, 61-181. doi: 10.1017/s0024282920000079
St. Martin P, Mallik AU (2017) The status of non-vascular plants in trait-
based ecosystem function studies. Perspectives in Plant Ecology,
Evolution and Systematics, 27, 1-8. doi: 10.1016/j.ppees.2017.04.002
Stenroos S, Velmala S, Pykälä J, Ahti T (2016) Lichens of Finland. Helsinki:
Botanical Museum, Finnish Museum of Natural History. 896p.
Tuovinen V, Millanes AM, Freire-Rallo S, Rosling A, Wedin M (2021)
Tremella macrobasidiata and Tremella variae have abundant and
widespread yeast stages in Lecanora lichens. Environmental
Microbiology, 23(5), 2484-2498. doi: 10.1111/1462-2920.15455
Umaña-Tenorio L, Sipman HJM, Lücking R (2002) Preliminary Checklist
of Lichens from Costa Rica, Version 1.2. Available at: http://archive.
fieldmuseum.org/ticolichen/checklist.html
Valencia-Islas NA, Arguello JJ, Rojas JL (2020) Antioxidant and Enchylium conglomeratum
photoprotective metabolites of Bunodophoron melanocarpum, a [Robert Lücking]
ABSTRACT
Colombia is recognised for its remarkable biodiversity as the country ranked first for bird and orchid richness, second
for amphibians, butterflies, and freshwater fishes, third for palms and reptiles, and fourth for mammals. By contrast, the
diversity of fungi in Colombia is not well known, with at least 7,241 species, a low number compared to that in other tropical
countries. Yeast diversity in paticular has been little explored, and little is known about the sampling efforts within this group
in Colombia. This study reviewed published studies from repositories, databases, and public collections reporting the isolation
and identification of environmental yeasts in Colombia in the years 2000 to 2020. All clinical isolations (human or related to
zoonotic diseases) were excluded. We found 112 species of environmental yeasts occurring in four regions and 11 departments
of Colombia, with 73.4% of yeasts belonging to Ascomycota and 26.6% to Basidiomycota. Thirty-two percent of publications
reported yeasts only to genus level, suggesting the potential to find and describe new species in Colombia. The most common
primary environments were associated with different parts of plants, soils, and animals. Secondary environments for the
reported yeasts were fermented fruits, fermented beverages, soils surrounding sugarcane crops, and soils contaminated
with hydrocarbons. The most representative species were Cryptococcus neoformans (544 strains), Cryptococcus vini (360),
Cryptococcus gattii (243), Saccharomyces cerevisiae (201), and Meyerozyma guilliermondii (107). There is an evident lack of
sampling efforts in several geographic regions of Colombia, such as the Amazon, Orinoquia, Caribbean, Islands, and Oceans.
We emphasise that future studies on the yeasts of Colombia should focus on natural ecosystems in these regions.
RESUMEN
Colombia es reconocida por su gran biodiversidad. Es el primer país con mayor biodiversidad en el mundo de aves y orquídeas,
segundo lugar en anfibios, mariposas y peces de agua dulce, tercer lugar en palmeras y reptiles y cuarto lugar en mamíferos.
La diversidad de hongos en Colombia no es bien conocida, donde hay al menos 7.241 especies, siendo un número bajo
cuando se compara con otros países. Las levaduras han sido poco exploradas y se sabe poco sobre su esfuerzo de muestreo.
Este trabajo tuvo como objetivo revisar todas las publicaciones, repositorios, bases de datos y colecciones públicas que
reportaron aislamiento e identificación de levaduras en Colombia, entre 2000 y 2020. Algunas de las combinaciones a buscar
en cada base de datos fueron: “yeast AND Colombia”, “yeasts AND diversity AND Colombia OR yeasts AND species AND
Colombia” en inglés y español. Se excluyeron todos los aislamientos clínicos (humanos o relacionados con enfermedades
zoonóticas). De 24.320 reportes iniciales, fueron seleccionados 97 reportes para este análisis. Fueron encontradas 112
especies de levaduras, ubicadas en cuatro regiones (11 departamentos). El 73.4% de las levaduras pertenecen a Ascomycota
y el 26.6% a Basidiomycota. Trienta-y-dos por ciento de las publicaciones reportaron solo géneros de levaduras, sugiriendo
el potencial para encontrar y describir nuevas especies de levaduras en Colombia. En total, fueron reportados 68 géneros
y 112 especies, donde las regiones geográficas con mayor información fueron el Pacífico y la región Andina. Los ambientes
primarios más comunes se asociaron con diferentes partes de plantas, suelo y animales. Los ambientes secundarios que
reportaron levaduras fueron frutas fermentadas, bebidas fermentadas, suelos que rodean el cultivo de caña de azúcar y
suelos contaminados con hidrocarburos. Las especies más representativas fueron Cryptococcus neoformans (544 cepas),
Cryptococcus vini (360), Cryptococcus gattii (243), Saccharomyces cerevisiae (201), y Meyerozyma guilliermondii (107). Fueron
identificados cinco clusters de géneros a partir del análisis de redes, donde es posible identificar grupos de levaduras que
podrían coexistir y ser aisladas simultáneamente. Por ejemplo, el cluster 1 incluye a los géneros Candida, Rhodotorula y
Geotrichum como los más frecuentes. El cluster 2 incluye a Pichia y Saccharomyces como nodos principales, mientras que
los cluster 3 y 4 fueron representados por Cryptococcus y Aureobasidium, respectivamente. Este análisis ofrece el potencial
para comparar, en una forma rápida y fácil, ecosistemas complejos y diferentes para entender como las asociaciones de
levaduras pueden influir las comunidades microbianas. Hay una evidente falta de esfuerzo de muestreo en varias regiones
geográficas, tales como Amazonas, Orinoquía, Caribe y océanos y se hace el énfasis en enfocar los muestreos en estas
regiones, principalmente en ecosistemas naturales.
FIGURE 2. PRISMA 2020 flow diagram for the systematic reviews performed in this analysis.
Inclusion criteria whereas 410 studies were excluded for one or several of the
a. Original papers and documents. following reasons: 1. reports without any yeast identification
(n=16); 2. literature reviews (n=31); 3. duplicated information
b. Papers and documents in English and Spanish.
between original articles and theses (n=13); 4. studies
c. Papers and documents that have isolated yeasts from evaluating other aspects unrelated to diversity, such as
Colombia and identified it, at least to the genus level. clinical isolation, genetic engineering, or bioprocessing
improvement, among others (n=127); 5. studies that were
Exclusion criteria not from Colombia (n=21); and 6. fungal reports that did not
a. Papers without any yeast identification or only metabar- include any yeasts or yeast-like organisms (n=86). We added
coding results. studies that were not found through the literature search but
that were available from other sources to the 66 selected
b. Review papers and documents.
studies, such as documents from university repositories
c. Duplicate papers and documents in different databases and reports from the Colombian collection database SIB
or repositories. (Sistema de Información Biológica). Those were processed
d. Theses already published as papers or books. as described above. Thirty-one reports were also included,
corresponding to three books or book chapters, 17 theses,
e. Papers with clinical isolates.
and nine SIB reports. Literature searches were classified as
f. Papers that do not involve yeast diversity assessments. research papers, book chapters, books, theses, conference
proceedings, book chapters, and database collections.
Data analyses Research papers and theses were the most common type
Selected papers were organised, standardised using of publications found in these years. We highlighted that
Mendeley, and exported to the RIS format (Research several reports corresponded to grey literature and were
Information System) in a plaintext file. This file was imported not submitted to peer-review. Therefore, 97 records of
into VOSviewer v1.6.17 (Van Eck & Waltman, 2010). Two environmental yeasts were selected for the period between
types of analyses were performed: 1. a co-occurrence 2000 and 2020 (Figure 2).
network, where the field “keywords” was updated with the Pioneer reports about yeasts in Colombia were identified,
genera found in each paper; and 2. a co-author network. We but not included in the analyses. The first reports of
showed all keywords or authors in the generated map. The environmental yeast isolates in Colombia were based on
nodes in the reconstructed network represent the genera, the serotyping of clinical and environmental isolates of
whereas the edges (i.e., connections) correspond to a strong Cryptococcus neoformans (Ordoñez & Castañeda, 1994).
and significant correlation between nodes. The clustering These isolates were recovered between 1972 and 1992.
coefficient was defined as how nodes are embedded in In total, 29 strains of C. neoformans were isolated from
their neighbourhood and, thus, the degree to which they environmental samples. Duarte et al. (1994) identified
tend to cluster together. Several characteristics, such as Cryptococcus yeasts associated with species of Eucalyptus
region, category, and specific substrate where yeasts were in a first evaluation of the ecology of C. neoformans var. gattii
isolated, were extracted from each report for analysis. We in Colombia. Caicedo et al. (1996) reported C. neoformans
also updated the taxonomy and nomenclature of all records in pigeon droppings from the urban perimeter of Santiago de
using Mycobank (Robert et al., 2013) or Index Fungorum Cali. In this study, 59 strains of this species were isolated.
(http://www.indexfungorum.org/) and computed the Another study characterised microorganisms with pectinolytic
frequency of inclusion for each genus or species (Table S1). activity from Mangifera indica (mango), such as Pichia
For example, Cryptococcus was updated and separated into kudriavzevii (formerly Candida krusei), Candida sorboxylosa,
the segregated Vanrija, Papiliotrema, Cutaneotrichosporon, Aureobasidium pullulans, and Candida insectorum (Feoli et
Solicoccozyma, among others. In addition, we organised al., 1997).
the richness and abundance of species in a spreadsheet
according to paper, and we classified reports according to the Diversity of yeasts of Colombia
substrate wherever available. Finally, we included additional We found 173 environmental yeasts belonging to 68 genera.
fields, such as region and year of publication (Table S1). From these, 112 yeasts were identified at the species
level. 73.4% were Ascomycota and 26.6% Basidiomycota.
RESULTS AND DISCUSSION 288 recorded yeasts, out of 3,557, were not identified by
Literature searches any specific method. However, this number is higher since
24,320 studies were identified from searches in the many reports were excluded from this study because they
abovementioned databases using different combinations of identified no yeast strains. The same species isolated from
English and Spanish keywords. After the removal of 14,977 a different substrate were assumed to be different samples.
duplicates, 8,814 further studies did not pass our initial According to Gaya et al. (2021), the recorded richness of
checks. The remaining 476 studies were assessed for fungi in Colombia differs in the five different geographical
eligibility by scanning their abstracts and results sections. regions recognised in the country. This fact is also true for
After this step, 66 studies were selected for full-text reviews, yeasts but with different geographic patterns evidenced.
The highest yeast species richness was recorded for the and 1,643 yeast strains found (Table 1). In general, different
Pacific region (103 species), followed by the Andean (77 parts of plants were assessed, including fruit pulp (Ramirez-
spp.), Caribbean (eight spp.), and Orinoquia (two spp.) Castrillon et al., 2019), fruit skin (Vanegas Cordoba et
regions. There are no reports of culturable yeasts for the al., 2004), phyllosphere (Medina et al., 2009; Zapata
Amazon region to date (Figure 3). However, Vasco-Palacios Narvaez et al., 2016), seed (Parsa et al., 2016), and cortex
et al. (2018) reported 21 operational taxonomic units (OTUs) (Contreras Martinez et al., 2011). A second group included
belonging to the genus Cryptococcus from metabarcoding non-fermented or fermented beverages and foods with
analyses in three different Amazon forests. The seawater 26 samples. For this group, we found 670 yeast strains.
and insular territory from the Caribbean and Pacific oceans The non-fermented beverages studied included sugarcane
also lack information on yeasts. There is no report from juice and fruit juices (i.e., pineapple, blackberry, grape,
islands such as San Andrés, Providencia, Santa Catalina, guanabana, and orange). On the other hand, fermented
or Gorgona. In this sense, mainly the Andean and Pacific beverages included “champús”, “chicha”, kumis, kefir (Alzate
regions, where the main research groups working with yeasts et al., 2016), and fermented juices. The base from each
(see section on Yeast Research Groups of Colombia) are fermented beverage is variable. For example, “Champús”
located, are represented in the sampling efforts for yeasts is generally made with pineapple fruit pulp, “lulo” fruit
in Colombia. We suggest increasing efforts in sampling and pulp and maize seed, while “kumis” uses fermented milk
isolating yeasts from other regions of Colombia, such as the (Chaves-Lopes et al., 2014). Some foods include milk and
Amazon, Orinoquia, Caribbean, Islands, and Oceans. derivatives (Lopez et al., 2010), beebread, and beehoney
Regarding the substrate of isolation, 3,559 strains were (Portillo Carrascal, 2016). We found nine samples from soil
categorised into 12 substrates (Table 1). Plant substrates substrates associated with parks, crops, composting, or
exhibited higher sampling efforts, with 36 samples reported natural ecosystems, such as forests (Miles et al., 2012).
FIGURE 3. Yeast strains, species, genera, and number of reports for different geographic regions of Colombia.
In this category, 162 yeast strains were reported. Yeasts importance of each species. For example, C. neoformans
isolated from animals (nine samples) were reported mainly and C. gattii are clinically significant, with several reports
from faeces (birds, cows, or pigs) or associated with pigs or studying the spreading of strains in different environments
fish (Londoño-Franco et al., 2014), where 239 strains were associated with possible hotspots of contamination for
found. From the air, reports assessed fungal communities humans, such as plants (Granados et al., 2005; Quintero
associated with gravimetric deposition of cells, including et al., 2005), soil (Firacative et al., 2011), animal faeces
environments such as libraries or buildings (Toloza Moreno (Quintero et al., 2005; Virviescas et al., 2018), food (Lopez
et al., 2012, Hernandez et al., 2012; Toloza & Lizarazo 2013; et al., 2010; Daza-Merchan 2011; Lopez Molinello, 2011), or
Mendez-Puentes et al., 2015). Two-hundred-and-forty yeast wastewater (Osorio-Vanegas, 2019).
strains were reported from water, both fresh and wastewater. In our analyses, yeast species were updated to current
Regarding freshwater samples, rivers, lakes and drinking names according to Mycobank (Robert et al., 2013). Several
water were assessed, whereas wastewater included organic yeast names were merged after the initiative 1 Fungus =
sludges, wastewater treatment plants, and leachate waters, 1 Name (Taylor, 2011), and other yeasts had taxonomic
among others. The fermentation industry category included changes, showing different nomenclatures depending on the
all modified environments related to ethanol distilleries. report. Therefore, we combined these cases as synonyms. For
Finally, contaminated environments included samples example, Wickerhamomyces anomalus is the accepted name
containing petroleum derivatives (Delgadillo Ordoñez et al., for several isolates reported in Colombia (Lopez-Arboleda et
2017; Ordoñez-Burbano et al., 2018). al., 2012; Cuervo-Mulet et al., 2015), although this species
was also reported as Pichia anomala (Delgadillo-Ordoñez
TABLE 1. Yeast strains by substrate type. Some reports were et al., 2017) or Candida pelliculosa (Vanegas et al., 2004).
included in more than one category. A similar case occurred with Pichia kudriavzevii, which was
reported with the current name (Delgado-Ospina et al., 2020),
Number of Yeast but also as Candida krusei (Herrera-Lopez et al., 2017; Castillo
Category
reports strains et al., 2018; Osorio-Vanegas 2019) or Issatchenkia orientalis
Plant 36 1,643 (Osorio-Cadavid et al., 2008). Finally, 30 morphotypes of
yeasts were identified only at the generic level. This number
Fermented beverages 6 367 might reflect the inability to identify species with phenotypical
or biochemical tests, the difficulty in separating cryptic
Animals 9 239
species, the inability of chosen molecular marker to separate
Non-fermented beverages 11 162 species within the same genus, or even the possibility that
the yeast under discussion is a species that is to science,
Soil 9 162
such as the case for Candida (Osorio-Vanegas, 2019) or
Freshwater 6 127 Wickerhamomyces (Ramirez-Castrillon et al., 2019).
Food 9 121
Yeast genera network
Wastewater 7 113 The growth and metabolic activities of yeasts are moderated
Fermentation industries 5 79 by a network of strain and species interactions, including
interactions with bacteria and other fungi (Fleet, 2007).
Air 7 19 With the network, we can identify the most frequent and
Contaminated connected yeasts and the association between them. In this
2 7 sense, we analysed the co-occurrence of different genera for
environments (petroleum)
all reports. Our results identified five yeast genera clusters,
9 (SIB where cluster 1 included Candida spp., Rhodotorula spp., and
Without information 520
Collection) Geotrichum spp. as the most frequent yeasts. Other clusters
Total 106 3,557 were represented by the most frequent genera. For example,
cluster 2 included Pichia spp. and Saccharomyces spp., cluster
3 included Cryptococcus spp., and cluster 4 was represented
by Aureobasidium spp. (Figure 4a). The most frequent and
The yeast species with the highest number of strains connected genus was Candida, with 42 reports and 61 genera
reported in the literature were Cryptococcus neoformans (544 connections. The most frequent and connected Basidiomycota
strains), followed by Cryptococcus vini (formerly Kloeckera yeast was Rhodotorula (36 reports; 58 connections). Other
apiculata), C. gattii (243 strains), and Saccharomyces yeast genera were also represented in the networks, such as
cerevisiae (201 strains). Some yeasts were mentioned in a Pichia (26 reports; 53 connections), Meyerozyma (16 reports;
larger number of studies, such as S. cerevisiae (35 studies), 45 connections), Saccharomyces (23 reports; 40 connections),
followed by Geotrichum sp. (23), Candida sp. (21), and C. and Hanseniaspora (11 reports; connections). We highlighted
neoformans (21). By contrast, 59 yeast species were only that Candida was overrepresented in this analysies with 28
isolated once or twice. These results correlate with the species found. On the other hand, we found only five species
of Cryptococcus from 1,297 strains. This network suggests ecosystems in order to understand how the yeast associations
that some coexisting yeasts should be found in the same may influence the microbial communities. Several reports
substrate. The probability of finding yeasts belonging to already support these findings (Barberán et al., 2012; Ma et
Candida or Rhodotorula is high due to their association with al., 2021).
other genera (up to 89.7%) and ubiquitous character (Urbina
& Aime, 2018). In the same sense, Pichia is highly associated Useful yeasts
with Saccharomyces strains, and this result can be related to Yeasts represent a significant potential for modern and
the artificial substrate of fermentation industries or fruit pulps traditional biotechnology. Since ancient times they have been
(Abranches et al., 2001). This analyses offers the potential recognised as protagonists in the production of food, such as
to compare, quickly and most easily, different and complex in the production of bread, or as beneficial participants in the
FIGURE 4. Yeast networks for Colombia. A Genera network. B Author network. Each colour represents different clusters of yeast genera.
The size of the node represents the occurrence of the genus or author by reports.
production of different cheeses (Lopez et al., 2010; Souza del Valle, Cali), who works with Dr Raul Cuervo-Mulet
et al. 2021), probably contributing to the maturation, texture, (Universidad de San Buenaventura, Cali) and Dr Mauricio
and flavour of the cheese. They are also protagonists in the Ramirez-Castrillon (Universidad Icesi, Cali) (Osorio-Cadavid
production of fermented beverages such as wine (Hernandez et al., 2008; Lopez-Arboleda et al., 2010; Gaviria & Osorio
et al., 2011), beer (Walteros-Pinzón et al., 2020; Amaya- 2012; Mambuscay et al., 2013; Silva-Bedoya et al., 2014;
Jimenez & Diaz-Pascal, 2019), and “guarapo” (Rodríguez Argote et al., 2015; Cuervo-Mulet et al., 2015; Ramirez-
Gonzales, 2015), contributing characteristics such as Castrillon et al., 2019). In addition, Dr Raul Cuervo-Mulet
flavour. Saccharomyces cerevisiae is the most studied yeast works in collaboration with Dr Clemencia Chavez-Lopez
and is very important to the fermentation industry, mainly (University of Teramo). This large group focuses on isolating
in bioethanol production (Tesfaw et al., 2014). Several yeasts from environmental samples for biotechnological
reports isolated wild strains with better performance than purposes, focusing on fermentative yeasts for bioethanol,
commercial strains used in bioethanol, brewing, or bread novelty enzymes, organoleptic characteristics, or diversity.
production. Different strategies were employed to isolate In AGROSAVIA, the group studies yeast for biological pest
S. cerevisiae: first, spontaneous juice fermentation (Daza- control and is led by Dr Alba Marina Cotes, who works with Dr
Merchan 2011; Argote et al., 2015; Cuervo-Mulet et al., Jimmy Zapata and Dr Fernando Rodriguez. They also have a
2015) to produce, fermented beverages, such as champús yeast collection in this research Institute (Cotes et al., 2004,
(Osorio-Cadavid et al., 2008) and chicha (Lopez-Arboleda et 2011; Zapata-Narvaez et al., 2016). In Bogotá, a research
al., 2010) (both made with corn) and kumis (whose base line of pathogenic yeasts, including Cryptococcus, is led by
is milk) (Chaves-Lopez et al., 2012); and second, use of Dr Patricia Escandón (Instituto Nacional de Salud, Bogotá),
resources directly associated to bioethanol distillery, such who works alongside Dr Elizabeth Castañeda, Dr Carolina
as sugarcane juice, molasses and tanks, among others Firacative, and others looking for yeasts in environmental
(Vasquez-Castillo et al., 2016b). samples with serotypes that could affect humans (Escandon
Yeasts are used as sources for obtaining B-complex et al., 2005, 2010; Firacative et al., 2011; Velez & Escandon
vitamins, pigments, proteins from unicellular organisms, 2017; Anacona et al., 2018; Virviescas et al., 2018). Several
extracts and biomass, among other products (Vasquez- studies are not connected with the authors mentioned above,
Castillo et al., 2016a). Some yeasts present biological and suggesting that these authors possibly did not work directly in
antifungal control alternatives that facilitate their use as a research line related towards yeasts or fungi but occasionally
controllers of spoilage of food (Buitrago-Estrada & Escobar- published information about yeast isolations. For example, Dr
Romero, 2009), flowers (Zapata-Narvaez et al., 2016), plants, Luz Marina Lizarazo (Universidad Pedagógica y Tecnológica
and crops (Cotes et al., 2004). Certain reported yeasts can de Colombia, Tunja), who works with environmental biology,
produce products that have potential industrial applications, focusing on aerobiology, has have frequently isolated yeasts.
such as lipases obtained from Candida rugosa (Domínguez de Other authors, such as Trujillo and Echeverry-Hernandez
María et al., 2006). These lipases have been widely used in (2015), Oviedo Zumaque et al. (2009) or Lara Mantilla &
fermentation technologies, biocatalytic assays, detergents, Burgos Portacio (2012), also were found in this analysis but
and solvents. Yeasts have been reported as the feedstock of without connections with research groups focusing on yeasts.
lipids for biodiesel production (Arcos-Velasco 2020). Likewise,
there have been some reports of high biomass production of CONCLUSIONS
yeasts with few nutrients, and their ease of cultivation reported From the 97 selected reports for our analyses, we found
by Olaya & Ossa (2020) represents an enzymatic capacity 173 yeasts belonging to 68 genera, with 73.4% of the
that will allow the development of bioremediation methods. yeasts belonging to Ascomycota. The geographic regions
To achieve these possibilities, several reports isolated with more samples were the Andean and Pacific regions,
yeasts from natural ecosystems or modified substrates. supported by research groups working with yeasts. We
For example, Romero et al. (2017) explored non-fermented suggest isolating and characterising the yeast diversity in
juices, but fruit pulps were also sampled (Argote et al., 2015). other regions of Colombia, such as the Amazon, Orinoquia,
Natural ecosystems included solid-state fermentation of Caribbean, Islands, and Oceans. Cryptococcus, Candida, and
cocoa (Delgado-Ospina et al., 2020) or coffee (Quintero et al., Saccharomyces were the most reported genera for Colombia,
2012), but also sampling from soils (Argote et al., 2015) or air having the most connections with other genera. Specifically,
(Toloza-Moreno & Lizarazo-Forero, 2011; Mendez-Puentes et the most frequent species were Cryptococcus neoformans,
al., 2015). Finally, aquatic systems were sampled to assess C. vini, C. gattii, Saccharomyces cerevisiae, Meyerozyma
lipid-accumulation or pigment production abilities, such as guilliermondii, and Pichia kudriavzevii. We identified five
wastewater, lakes or drinking water (Silva-Bedoya et al., 2014; clusters of co-occurrences of yeasts, where the probability
Osorio-Vanegas, 2019; Villota et al. 2020). to co-isolate yeasts such as Saccharomyces spp. and Pichia
spp. is high, and should be related to the purpose of the
Research groups working with yeasts in Colombia isolation. Finally, the yeast species reported in this chapter
Eight Colombian research groups that work with yeasts were showed many possibilities for biotechnological exploration,
identified from the published reports (Figure 4b). In Cali, ranging from fermentation, to food, and biofuel industries.
the group is led by Dr Esteban Osorio-Cadavid (Universidad
Acknowledgements
100 CATALOGUE OF FUNGI OF COLOMBIA
CHAPTER 7
We thank the Research Group in Mycology, led by M.Sc. Luz de Cryptococcus gattii aislado de Terminalia catappa en Montería,
Dary Caicedo, and the “Asociación Colombiana de Micología” Córdoba, Colombia. Revista Cubana de Medicina Tropical, 63(2),
(ASCOLMIC) for the information provided on some reports of yeasts 117-122.
Cotes AM, Villamizar LF, García Murillo PG, Gomez Alvarez MI, Fuentes
of Colombia; and the reviewers Aida Vasco-Palacios and Robert
O, Diaz García A, Florez J, Grijalba EP, Lopez J (2004) Las Levaduras
Lücking for their insightful suggestions. Universidad Santiago de como Alternativa de Control Biológico de Fitopatógenos. Bogotá DC:
Cali funded this study (934-621120-2388). Corporación Colombiana de investigación agropecuaria. 21p.
Cotes AM, Zapata Narvaez YA, Diaz García A, García M, Medina C,
Cristancho D, Rodriguez S, Rodriguez F, Uribe D (2011) Selection
References of yeasts with potential to biological control of Botrytis cinerea.
Fitopatología Colombiana, 35(2), 51-56.
Abranches J, Starmer WT, Hagler AN (2001) Yeast-yeast interactions Cuervo-Mulet R, Argote F, Fernandez-Daza F, Osorio-Cadavid E (2015)
in guava and tomato fruits. Microbial Ecology, 42(2), 186-192. doi: Hongos levaduriformes aislados de frutas y suelo de la región del Valle
10.1007/s002480000111 del Cauca, Colombia. Cali: Editorial Bonaventuriana. 100p.
Alzate BCS, Rodríguez MC, Campuzano OM (2016) Identification of some Daza Merchán ZT (2011) Evaluación a escala laboratorio de la producción
kefir microorganisms and optimization of their production in sugarcane de etanol en presencia de vinaza a partir de levaduras nativas.
juice. Revista Facultad Nacional de Agronomía Medellín, 69(2), 7935- Universidad del Valle. Available at: https://bibliotecadigital.univalle.
7943. doi: 10.15446/rfna.v69n2.59138 edu.co/handle/ 10893/7636
Amaya-Jiménez NA, Diaz-Pascagaza LM (2019) Evaluación de perfiles Deak T (2007) Handbook of food spoilage yeasts. Boca Ratón: CRC
fermentativos para la elaboración de cerveza artesanal por levaduras Press. 352p.
nativas. Fundación Universidad De América. Available at: http:// Delgadillo-Ordoñez NC, Posada-Suárez LR, Marcelo E, Cepeda-Hernández
repository.uamerica.edu.co/handle/20.500.11839/7724 ML, Sánchez-Nieves J (2017) Aislamiento e identificación de levaduras
Anacona C, González-Cuellar FE, Vásquez-A LR, Escandón P (2018) First degradadoras de hidrocarburos aromáticos, presentes en tanques de
isolation and molecular characterization of Cryptococcus neoformans gasolina de vehículos urbanos. Revista Colombiana de Biotecnología,
var. grubii in excreta of birds in the urban perimeter of the municipality 19(2), 141-151. doi: 10.15446/rev.colomb.biote.v19n2.70278
of Popayán, Colombia. Revista Iberoamericana de Micología, 35(3), Delgado-Ospina J, Triboletti S, Alessandria V, Serio A, Sergi M, Paparella
123-129. doi: 10.1016/j.riam.2018.01.005 A, Rantsiou K, Chaves-López C (2020) Functional biodiversity of
Arcos-Velasco LV (2020) Selección y caracterización de levaduras yeasts isolated from Colombian fermented and dry cocoa beans.
oleaginosas aisladas a partir de sistemas acuáticos de la ciudad de Microorganisms, 8(7), 1-17. doi: 10.3390/microorganisms8071086
Cali. Universidad Santiago de Cali. Available at: https://repository. Domínguez de María P, Sánchez-Montero JM, Sinisterra JV,
usc.edu.co/handle/20.500.12421/3538 Alcántara AR (2006) Understanding Candida rugosa lipases: an
Argote F, Cuervo-Mulet R, Osorio-Cadavid E, Delgado-Ospina J, Villada overview. Biotechnology Advances, 24, 180-196. doi: 10.1016/j.
HS (2015) Evaluación de la producción de etanol a partir de melaza biotechadv.2005. 09.003
con cepas nativas Saccharomyces cerevisiae. Biotecnología en el Duarte A, Ordoñez N, Castañeda E (1994) Asociación de levaduras del
Sector Agropecuario y Agroindustrial, 13(2), 40-48. doi: 10.18684/ género Cryptococcus con especies de Eucalyptus en Santa fé de Bogotá.
BSAA(13)40-48 Revista del Instituto de Medicina Tropical de Sao Paulo, 36(2): 125-130.
Barberán A, Bates ST, Casamayor EO, Fierer N (2012) Using network Escandón P, Quintero E, Granados D, Huérfano S, Ruiz A, Castañeda E
analysis to explore co-occurrence patterns in soil microbial (2005) Aislamiento de Cryptococcus gattii serotipo B a partir de
communities. The ISME Journal, 6(2), 343-351. doi: 10.1038/ detritos de Eucalyptus spp. en Colombia. Biomédica: revista del
ismej.2011.119 Instituto Nacional de Salud, 25(3), 390-397.
Becher PG, Hagman A, Verschut V, Chakraborty A, Rozpędowska E, Escandón P, Sánchez A, Firacative C, Castañeda E (2010) Isolation of
Lebreton S, Bengtsson M, Flick G, Witzgall P, Piškur J (2018) Chemical Cryptococcus gattii molecular type VGIII, from Corymbia ficifolia detritus
signaling and insect attraction is a conserved trait in yeasts. Ecology in Colombia. Medical Mycology, 48(4), 675-678.
and Evolution, 8(5), 2962-2974. doi: 10.1002/ece3.3905 Feoli M, Gómez Z, Muñoz A (1997) Aislamiento y caracterización de
Buitrago-Estrada J, Escobar-Romero AM (2009) Aplicación de levadura microorganismos con actividad pectinolítica a partir de Mangifera
Candida spp. como una alternativa viable para la retardación en indica. Revista Colombiana de Ciencias Químico-Farmacéuticas, 26(1),
la pudrición del banano (Musa acuminata). Pontificia Universidad 33-37.
Javeriana. Available at: https://repository.javeriana.edu.co/ Firacative C, Torres G, Rodríguez MC, Escandón P (2011) First environmental
handle/10554/8215 isolation of Cryptococcus gattii serotype B, from Calcuta, Colombia,
Caicedo LD, Álvarez MI, Llanos CA, Molina D (1996) Cryptococcus Biomedica, 31(1), 118-123.
neoformans en excretas de palomas del perímetro urbano de Cali. Fleet GH (2007) Yeasts in foods and beverages: impact on product quality
Revista Colombiana Médica, 27(3-4), 1-4. and safety. Current Opinion in Biotechnology, 18(2), 170-175.
Carlile MJ, Watkinson SC, Gooday GW (2001) The Fungi, 2nd edition. San Gaviria J, Osorio E (2012) Diversidad de levaduras asociadas a
Diego: Gulf Professional Publishing. 70p. inflorescencias de mango y flores de “lulo arbóreo”. Biotecnología en el
Castillo Arroyo PL, Betancur Hurtado CA, Pardo Pérez E (2018) Sector Agropecuario y Agroindustrial, 10(2), 160-169.
Caracterización de microorganismos con potencial probiótico aislados Gaya E, Vasco-Palacios AM, Vargas-Estupiñán N, Lücking R, Carretero
de estiércol de terneros Brahman en Sucre, Colombia. Revista de J, Sanjuan T, Moncada B, Allkin B, Bolaños-Rojas AC, Castellanos-
Investigaciones Veterinarias del Perú, 29(2), 438-448. doi: 10.15381/ Castro C, Coca LF, Corrales A, Cossu T, Davis L, dSouza J, Dufat A,
rivep.v29i2.14482 Franco-Molano AE, García F, Gómez-Montoya N, González-Cuellar FE,
Chaves-López C, Tofalo R, Serio A, Paparella A, Sacchetti G, Suzzi Hammond D, Herrera A, Jaramillo-Ciro MM, Lasso Benavides C, Mira
G (2012) Yeasts from Colombian Kumis as a source of peptides MP, Morley J, Motato-Vásquez V, Niño-Fernández Y, Ortiz-Moreno ML,
with Angiotensin I converting enzyme (ACE) inhibitory activity in Peña-Cañón ER, Ramírez-Castrillón M, Rojas T, Ruff J, Simijaca D,
milk. International Journal of Food Microbiology, 159(1), 39-46. doi: Sipman HJM, Soto-Medina E, Torres G, Torres-Andrade PA, Ulian T,
10.1016/j.ijfoodmicro.2012.07.028 White K, Diazgranados M (2021) ColFungi: Colombian resources for
Chaves‐López C, Serio A, Grande‐Tovar CD, Cuervo‐Mulet R, Delgado‐ Fungi Made Accessible. Richmond: Royal Botanic Gardens, Kew, 36p.
Ospina J, Paparella A (2014) Traditional fermented foods and Gonzales A, Valenzuela L (2003) Saccharomyces cerevisiae. In: Romero
beverages from a microbiological and nutritional perspective: the EM, Romero JCM (eds). Microbios en línea. Centro de Investigación
Colombian heritage. Comprehensive Reviews in Food Science and Food sobre fijación de nitrógeno. Cuernavaca: UNAM. Available at: http://
Safety, 13(5), 1031-1048. doi: 10.1111/1541-4337.12098 www.biblioweb.tic.unam.mx/libros/microbios/Cap16/
Contreras Martínez OI, Aycardi Morinelli MP, Alarcón Furnieles JL, Jaraba Granados DP, Castañeda E (2005) Isolation and characterisation of
Ramos AM, Martínez Contreras OI (2011) Identificación presuntiva Cryptococcus neoformans varieties recovered from natural sources
in Bogotá, Colombia, and study of ecological conditions in the area. Colombia Susceptibilidad a los antimicóticos de levaduras aisladas
Microbial Ecology, 49(2), 282-290. de sistemas de aguas de la ciudad de Santiago de Cali, Colombia.
Hernández CJD, Trujillo NYY, Durán ODS (2011) Contenido fenólico e Universidad Santiago de Cali. Available at: https://repository.usc.edu.
identificación de levaduras de importancia vínica de la uva isabella co/handle/20.500.12421/4481
(Vitis labrusca) procedente de villa del rosario (Norte de Santander). Oviedo Zumaqué L, Lara Mantilla C, Mizger Pantoja M (2009) Levaduras
Vitae, 18(1), 17-25. autóctonas con capacidad fermentativa en la producción de etanol
Hernández DR, Lopez-Valiente E, Lizarazo-Forero LM (2012) Caracterización a partir de pulpa de excedentes de plátano musa (aab simmonds)
Fúngica en el Archivo Histórico de la Universidad Pedagógica y en el departamento de Córdoba, Colombia. Revista Colombiana de
Tecnológica de Colombia. Ciencia en Desarrollo, 4(1), 115-128. Biotecnología, 11(1), 40-47.
Herrera-López MA, Lora-Suarez F, Loango-Chamorro N (2017) Identificación Page MJ, McKenzie JE, Bossuyt PM, Boutron I, Hoffmann TC, Mulrow
de microorganismos aislados a partir de lodos residuales de una CD (2021) The PRISMA 2020 statement: an updated guideline for
planta de tratamiento de un sector curtidor del Quindío. Revista de la reporting systematic reviews. BMJ, 71, 372. doi: 10.1136/bmj.n71
Asociación Colombiana de Ciencias Biológicas, 29(1), 103-118. Parsa S, García-Lemos AM, Castillo K, Ortiz V, López-Lavalle LAB, Braun
Kurtzman C, Fell JW, Boekhout T (2011) The Yeasts: a Taxonomic Study. 5th J, Vega FE (2016) Fungal endophytes in germinated seeds of the
edition. Amsterdam: Elsevier Science. 2354p. common bean, Phaseolus vulgaris. Fungal Biology, 120(5), 783-790.
Lara Mantilla C, Burgos Portacio Á (2012) Potencial probiótico de cepas doi: 10.1016/j.funbio.2016.01.017
nativas para uso como aditivos en la alimentación avícola, Revista Portillo Carrascal CY (2016) Identificación de levaduras presentes en
Colombiana de Biotecnología, (1), 31-40. el proceso de transformación de polen corbicular a pan de abejas
Londoño-Franco LF, Villamarín-Muñoz JA, David-Ruales CA (2014) por métodos tradicionales y moleculares. Universidad Nacional de
Identificación de hongos en cachama blanca (Piaractus brachypomus) Colombia. Available at: https://repositorio.unal.edu.co/handle/
en cultivos semi-intensivos de Norte de Santander, Colombia. unal/56727
Orinoquia, 18, 247-255. Quintero E, Castañeda E, Ruiz A (2005) Environmental distribution
López D, Jiménez M, López-Molinello A (2010) Identificación de hongos of Cryptococcus neoformans in the department of Cundinamarca-
benéficos que participan en el proceso de obtención del queso Paipa Colombia | Distribución ambiental de Cryptococcus neoformans en el
en lacteos Ibel, municipio de Belén (Boyacá). Revista Alimentos Hoy, departamento de Cundinamarca-Colombia. Revista Iberoamericana de
19(21), 85-108. Micologia, 22(2), 93-98.
López-Arboleda WA, Ramírez-Castrillón M, Mambuscay-Mena LA, Osorio- Quintero GIP, Mejía JM, Betancur GAO (2012) Microbiología de la
Cadavid E (2010) Diversidad de levaduras asociadas a chichas fermentación del mucílago de café según su madurez y selección.
tradicionales de Colombia. Revista Colombiana de Biotecnología, 12(2), Cenicafé, 63(2), 58-78.
176-186. Ramírez-Castrillón M, Usman LM, Silva-Bedoya LM, Osorio-Cadavid E
López-Arboleda WA, Mambuscay-Mena LA, Cuervo-Mulet RA (2012) Banco (2019) Dominant yeasts associated to mango (Mangifera indica) and
Levaduriformes. Cali: Universidad de San Buenaventura Cali. 54p. rose apple (Syzygium malaccense) fruit pulps investigated by culture-
López-Molinello A (2011) Diversidad de la microbiota fúngica del queso based methods. Anais da Academia Brasileira de Ciencias, 91(4),
paipa fabricado en Pacho, Cundinamarca. Revista ION, 24(1), 77-84. e20190052. doi: 10.1590/0001-3765201920190052
Ma Y, Pan Y, Liu Q, Huang L, Zhang W (2021) Co-occurrence patterns Riesute R, Salomskiene J, Moreno DS, Gustiene S (2020) Effect of
and assembly processes of microeukaryotic communities in a semi- yeasts on food quality and safety and possibilities of their inhibition.
enclosed aquaculture bay. Continental Shelf Research, 228, 104550. Trends in Food Science & Technology, 108, 1-10. doi: 10.1016/j.
doi: 10.1016/j.csr.2021.104550 tifs.2020.11.022
Mambuscay-Mena LA, López-Arboleda WA, Cuervo-Mulet R, Argote F, Robert V, Vu D, Amor ABH, van de Wiele N, Brouwer C, Jabas B, Szoke S,
Osorio-Cadavid E (2013) Identificación de las levaduras nativas Dridi A, Triki M, Daoud SB, Chouchen O, Vaas L, de Cock A, Stalpers
presentes en zumos de piña, mora y uva. Biotecnología en el Sector JA, Stalpers D, Verkley GJM, Groenewald M, dos Santos FB, Stegehuis
Agropecuario y Agroindustrial, 11(2), 136-144. G, Li W, Wu L, Zhang R, Ma J, Zhou M, Pérez S, Eurwilaichitr L,
Medina C, Cristancho DE, Uribe D (2009) Physiological responses and Ingsriswang S, Hansen K, Schoch C, Robbertse B, Irinyi L, Meyer W,
antagonistic capacity of yeast phyllospheric isolates obtained in Cardinali G, Hawksworth DL, Taylor JW, Crous PW (2013) MycoBank
blackberry crops (Rubus glaucus). Acta Biológica Colombiana, 14(3), gearing up for new horizons. IMA Fungus, 4(2), 371-379. doi: 10.5598/
181-198. imafungus.2013.04.02.16
Méndez-Puentes CA, Camacho-Suárez JG, Echeverry-Hernández S (2015) Rodríguez-Gonzales SA (2015) Evaluación Del Potencial Fermentativo De
Identificación de bacterias y hongos en el aire de Neiva, Colombia. Levaduras Nativas Para La Producción De Etanol a Partir De Mieles
Revista de Salud Pública, 17(5), 728-737. De Caña De Azúcar. Universidad Nacional de Colombia. Available at:
Miles LA, Lopera CA, González S, Cepero de García MC, Franco-Molano https://repositorio.unal.edu.co/handle/unal/55626
AE, Restrepo Restrepo S (2012) Exploring the biocontrol potential of Romero VC, Ortiz D, Serna-Jiménez JA (2017) Identificación de factores
fungal endophytes from an Andean Colombian Paramo ecosystem. microbiológicos asociados al deterioro de jugo (naranja-mandarina)
BioControl, 57(5), 697-710. mínimamente procesado para su bioconservación. Alimentos Hoy,
Monapathi ME, Bezuidenhout CC, James Rhode OH (2020) Aquatic yeasts: 24(39), 156-167.
diversity, characteristics and potential health implications. Journal of Sanjuan TI, Franco-Molano AE, Kepler RM, Spatafora JW, Tabima J, Vasco-
water and health, 18(2), 91-105. Palacios AM, Restrepo S (2015) Five new species of entomopathogenic
Olaya Hurtado DS, Ossa Cabra LT (2020) Actividad enzimática de fungi from the Amazon and evolution of neotropical Ophiocordyceps.
levaduras en diferentes ambientes del Valle del Cauca. Programa de Fungal Biology, 119(10), 901-916. doi: 10.1016/j.funbio.2015.06.010
Microbiología, Universidad Santiago de Cali. Cali, Colombia. Schoch CL, Seifert KA, Huhndorf S, Robert V, Spouge JL, Levesque CA,
Ordóñez N, Castañeda E (1994) Serotipificación de aislamientos clínicos Chen W, Fungal Barcoding Consortium (2012) Nuclear ribosomal
y del medio ambiente de Cryptococcus neoformans en Colombia. internal transcribed spacer (ITS) region as a universal DNA barcode
Biomédica, 14(3), 131-139. marker for Fungi. Proceedings of the National Academy of Sciences
Ordoñez Burbano DE, Abella Medina CA, Echeverry Tamayo A, Paz Lasprilla U S A , 109(16), 6241-6246. doi: 10.1073/pnas.1117018109
LM, Benítez-Campo N (2018) Biodegradación de hidrocarburos SiB Colombia (2021) Portal de Datos, Sistema de Información sobre
alifáticos saturados por microorganismos aislados de suelo Biodiversidad de Colombia. Available at: https://datos.biodiversidad.
contaminado con derivados del petróleo. Revista de Ciencias, 22(2), co/
33-44. Silva-Bedoya LM, Ramírez-Castrillón M, Osorio-Cadavid E (2014) Yeast
Osorio-Cadavid E, Chaves-López C, Tofalo R, Paparella A, Suzzi G (2008) diversity associated to sediments and water from two Colombian
Detection and identification of wild yeasts in Champús, a fermented artificial lakes. Brazilian Journal of Microbiology, 45(1), 135-142. doi:
Colombian maize beverage. Food Microbiology, 25(6), 771-777. 10.1590/S1517-83822014005000035
Osorio-Vanegas LS (2019) Susceptibilidad a los antimicóticos de levaduras Souza PV, Grecellé CBZ, Barreto F, Ramírez-Castrillon M, Valente P, Costa
aisladas de sistemas de aguas de la ciudad de Santiago de Cali, M (2021) Bacteria and yeasts associated to colonial cheese production
chain and assessment of their hydrolytic potential. Brazilian Journal of Vu D, Groenewald M, De Vries M, Gehrmann T, Stielow B, Eberhardt U,
Food Technology, 24, e2020286. doi: 10.1590/1981-6723.28620 Al-Hatmi A, Groenewald JZ, Cardinali G, Houbraken J, Boekhout T,
Taylor JW (2011) One fungus = one name: DNA and fungal nomenclature Crous PW, Robert V, Verkley GJM (2019) Large-scale generation and
twenty years after PCR. IMA Fungus, 2(2), 113-120. doi: 10.5598/ analysis of filamentous fungal DNA barcodes boosts coverage for
imafungus.2011.02.02.01 kingdom fungi and reveals thresholds for fungal species and higher
Tesfaw A, Assefa F (2014) Current trends in bioethanol production by taxon delimitation. Studies in Mycology, 92, 135-154. doi: 10.1016/j.
Saccharomyces cerevisiae: substrate, inhibitor reduction, growth simyco.2018.05.001
variables, coculture, and immobilisation. International Scholarly Walteros-Pinzón CD, Fernández MA, Reyes LH (2020) Caracterización
Research Notices, 2014, 532852. doi: 10.1155/2014/532852. de cepas de levadura colombiana Saccharomyces cerevisiae para
Toloza-Moreno DL, Lizarazo-Forero LM (2011) Aeromicrobiología del archivo su potencial uso en la producción de cerveza “Colombian Ale”.
central de la Universidad Pedagógica Y Tecnológica De Colombia Universidad de los Andes. Available at: https://repositorio.uniandes.
(Tunja-Boyacá). Acta Biológica Colombiana, 16(1), 185-194. edu.co/handle/1992/44911
Toloza-Moreno DL, Lizarazo-Forero LM, Blanco-Valbuena JO (2012) Zapata Narvaez YA, Diaz García A, Grijalba EP, Rodriguez F, Elad Y, Cotes
Concentración y composición microbiana en el ambiente de la AM (2016) Phyllosphere yeasts with potential for biological control
biblioteca central jorge palacios preciado de la Universidad Pedagógica of Botrytis cinerea in rose. Acta Horticulturae, 1144, 77-84. doi:
y Tecnológica De Colombia, Tunja. Actualidades Biológicas, 34(97), 10.17660/ActaHortic.2016.1144.11
241-252.
Toloza-Moreno DL, Lizarazo-Forero LM (2013) Calidad microbiológica
del ambiente de la Biblioteca Alfonso Patiño Rosselli, Tunja-Boyacá
(Colombia). Revista U.D.C.A Actualidad and Divulgación Científica, 16(1),
43-52.
Tournas VH, Katsoudas E (2005) Mould and yeast flora in fresh berries,
grapes and citrus fruits. International Journal of Food Microbiology,
105(1), 11-17.
Trujillo LC, Echeverry-Hernández S (2015) Aislamiento y caracterización
de levaduras presentes en el fruto del Syzygium malaccense (l.). Merr.
and L. M Perry (pomorroso) en la comuna 1 de la ciudad de Neiva.
Ingeniería y Región, 13(1), 37-37. doi: 10.25054/22161325.707
Urbina H, Aime MC (2018) A closer look at Sporidiobolales: ubiquitous
microbial community members of plant and food biospheres.
Mycologia, 110(1), 79-92. doi: 10.1080/00275514.2018.1438020
Van Eck, NJ, Waltman L (2010) Software survey: VOSviewer, a computer
program for bibliometric mapping. Scientometrics, 84(2), 523-538.
doi: 10.1007/s11192-009-0146-3
Vanegas Córdoba I, Yepes Pérez MS, Ruiz Villadiego OS (2004) Producción
de xilitol a partir de levaduras nativas colombianas. Revista Colombiana
de Biotecnología, 6(2), 31-36.
Vargas N, Restrepo S (2020) A checklist of ectomycorrhizal mushrooms
associated with Quercus humboldtii in Colombia. In: Pérez-Moreno J,
Guerin-Laguette A, Arzú RF, Yu FQ (eds). Mushrooms, Humans and
Nature in a Changing World. Berlin: Springer. Pp. 425-450.
Vasco-Palacios AM, Hernandez J, Peñuela-Mora MC, Franco-Molano AE,
Boekhout T (2018) Ectomycorrhizal fungi diversity in a white sand
forest in western Amazonia. Fungal Ecology, 31, 9-18. doi: 10.1016/j.
funeco.2017.10.003
Vásquez JA, Ramírez Castrillón M, Monsalve ZI (2016a) Update on
molecular characterization for yeast of industrial interest. Revista
Colombiana de Biotecnología, 18(2), 129-139. doi: 10.15446/rev.
colomb.biote.v18n2.61530
Vasquez-Castillo J, Laguado JA, Lopez Genera J, Gil NJ (2016b) New
sources and methods to isolate vinasse-tolerant wild yeasts efficient
in ethanol production. Annals of Microbiology, 66(1), 187-195. doi:
10.1007/s13213-015-1095-0
Vélez N, Escandón P (2017) Report on novel environmental niches
for Cryptococcus neoformans and Cryptococcus gattii in Colombia:
Tabebuia guayacan and Roystonea regia. Medical Mycology, 55(7), 794-
797. doi: 10.1093/mmy/myw138
Villota SV, Vargas AF, Martínez CA, Caicedo LD, Osorio E, Ramírez-Castrillón
M (2020) Producción de carotenoides en levaduras nativas aisladas
de sistemas acuáticos en Cali, Colombia. Revista De La Asociación
Colombiana De Ciencias Biológicas, 1(32), 103-114. doi: 10.47499/
revistaaccb.v1i32.215
Virviescas BC, Aragón FM, Vásquez-A LR, González-Cuellar FE, Escandón
P, Castro SH (2018) Caracterización molecular de Cryptococcus
neoformans recuperado de guano de palomas en Rivera y Neiva,
Colombia. Revista MVZ Córdoba, 23, 6991-6997. doi: 10.21897/
rmvz.1422
Vu D, Groenewald M, Szöke S, Cardinali G, Eberhardt U, Stielow B, de Vries
M, Verkleij GJ, Crous PW, Boekhout T, Robert V (2016) DNA barcoding
analysis of more than 9,000 yeast isolates contributes to quantitative
thresholds for yeast species and genera delimitation. Studies in
Mycology, 85, 91-105. doi: 10.1016/j.simyco.2016.11.007
Epicoccum sp.
[Marcela Guevara-Suarez]
104 CATALOGUE OF FUNGI OF COLOMBIA
Chapter 8
Micromycetes of Colombia: Focusing on the Hidden Diversity
Martha Cárdenas1, Marcela Guevara-Suárez2, Angelis Marbello1, Natalia Bolaños1, Pedro Jiménez3 & Silvia Restrepo1*
1
Laboratorio de Micología y Fitopatología (LAMFU), Departamento de ingeniería Química y de alimentos, Universidad de los Andes, Colombia.
2
Applied Genomics Research Group, Vicerrectoría de Investigación y Creación, Universidad de los Andes, Colombia.
3
Facultad de Ciencias Básicas y Aplicadas, Universidad Militar Nueva Granada, Bogotá, Colombia.
ABSTRACT
Colombia is a Neotropical country with a highly diverse fauna, flora, and microbial biota. However, mycology is still a developing
field, and currently, Colombia has reported only 5% of the fungal species known worldwide. The goal of this chapter is to
revisit the current state of the knowledge on Colombian microfungi, according to species registered in the ColFungi database,
excluding lichens. For this purpose, we categorised each taxon as macro- or micromycete according to the presence or
absence of macroscopically visible structures. In our definition, a total of 2,498 (57%) out of the 4,406 non-lichenized species
included in the database correspond to microfungi, most of them being members of the subkingdom Dikarya (92%). We found
that early diverging fungal lineages are underrepresented and need to be prioritised in future diversity surveys. Although next-
generation sequencing tools are being used worldwide to explore biodiversity, few studies to date have employed this approach
to assess fungal diversity in Colombia. In general, the relevance of performing an assertive polyphasic identification to species
level and the necessity to enrich our national culture collections are discussed.
RESUMEN
Colombia es considerado un país megadiverso en fauna, flora y microorganismos. En cuanto a los hongos, nuestro país
ha reportado el 5% de las especies conocidas en el mundo. El objetivo de este capítulo fue estudiar el estado actual de la
diversidad de microhongos en Colombia y resaltar los grupos taxonómicos que deben ser priorizados en futuros estudios de
diversidad de hongos en el país. Para este fin, se utilizaron las especies registradas en la base de datos de ColFungi y se realizó
una categorización adicional de las especies, asignándole a cada una la clasificación de macro- o micromicete, de acuerdo con
la presencia o ausencia de estructuras macroscópicamente visibles. Los líquenes fueron excluidos. Del total de 4,406 especies
pertenecientes al reino Fungi registradas en la base de datos de ColFungi, excluyendo los que son liquenizados, el 57% (2,498)
corresponden a registros de microhongos, siendo la mayoría miembros del subreino Dikarya (92%). También encontramos que
linajes basales de hongos, como por ejemplo los zigomicetes (filos Mucoromycota y Zoopagomycota), están subrepresentados
y deben ser priorizados en futuros estudios de diversidad. En los últimos años, las herramientas de secuenciación de nueva
generación han sido usadas para explorar la biodiversidad a nivel mundial. Aunque Colombia ha realizado esfuerzos para
evaluar la diversidad fúngica en el país usando estas aproximaciones, aún siguen siendo pocos los estudios que emplean
estas estrategias para estimar la diversidad fúngica. En este capítulo resaltamos la importancia de realizar estudios polifásicos
dirigidos hasta el nivel de especie, con el fin de tener identificaciones rigurosas y una estimación más precisa de los grupos
más representativos de comunidades fúngicas. Además, recalcamos la necesidad de enriquecer las colecciones nacionales de
microhongos, para incentivar así las investigaciones en torno al aprovechamiento de la diversidad y bioeconomía.
special attention has been paid to the study of microfungi. more effectively and to assess even complex habitats
Several efforts have been made to collect specimens with a rich microbiota (Lücking et al., 2020). However,
from different natural regions, elucidate their taxonomy, these technically advanced approaches are currently only
understand their ecology, and exploit their biotechnological available for a limited number of laboratories in Colombia.
value (Landínez-Torres et al., 2020). However, there is a need Therefore, hitherto published studies on micromycetes
to continue studying microbial life, for example to better focusing on traditional identification methods often have
understand the functioning of the soil and its ecosystem taxa identified only to the genus or morphospecies level,
services (Álvarez-Yela et al., 2017). Thus, the main goal of leading to an underestimation of the diversity of this fungal
this chapter was to revisit the current state of knowledge group in Colombia.
of the microfungi diversity of Colombia, according to the Fungal identifications remain challenging due to
species registered in the ColFungi database, and to highlight constant taxonomic changes. Where possible, species
the taxonomic groups that need to be prioritised in future identifications should be based on a polyphasic
fungal diversity studies because they have been overlooked, approach, including morphological, anatomical, chemical,
and also due to their potential biotechnological applications. physiological, and molecular data (Lücking et al., 2020).
To this end, we categorised each species, excluding In 2003, DNA barcoding was proposed to standardise
lichenised fungi, as either macro- or micromycete, depending the identification of all organisms to the species level
on the presence or absence of macroscopically visible (Hebert et al., 2003). The Internal Transcribed Spacer (ITS)
structures reported for each species. In addition, we cross- region was proposed as a DNA barcode for the kingdom
checked Mycobank (https://www.mycobank.org/) and Index Fungi (Schoch et al., 2012). Since then, ITS barcoding
Fungorum (http://www.indexfungorum.org/Names/Names. has become an essential tool for species identification in
asp) for the current nomenclature and taxonomy placement fungi. Nonetheless, in some genera, the ITS region is not
of each species. We determined the proportion (number variable enough for species-level identification, making it
of species / taxonomic group) of micromycetes within the necessary to use additional markers, such as β-tubulin
current accepted phyla and subphyla of Fungi (Li et al., 2021; (TUB), the DNA-directed RNA polymerase II largest (RPB1)
James et al., 2020). We also performed a search of Google and second-largest (RPB2) subunits, and the elongation
Scholar, without time restriction, to look for studies related to factor 1α (EF1) for accurate identification (Lücking et al.,
Colombian fungal diversity with the keywords "Micromycetes 2020). The method to achieve precise identification of
diversity" AND "Colombia"; "Microfungal diversity" AND
"Colombia"; "Microfungi diversity" AND "Colombia", excluding
lichens, and the same terms for Macromycetes. We also
searched for metagenomics and metabarcoding using the
modifiers "Metagenomics" AND "Microfungi" AND "Colombia".
Within the Mucoromycota, the Glomeromycotina, subphylum is reported, being represented by seven species.
comprising arbuscular mycorrhizal fungi, is represented by Basidiobolus ranarum and Conidiobolus coronatus are
106 species, 50 of which belong to three main genera: animal pathogens and can cause a human disease known
Glomus (20), Acaulospora (19), and Scutellospora (11). In as entomophthoromycosis (Morales-López et al., 2018).
a literature review conducted in Google Scholar and Web The remaining five species are insects pathogens (Barta &
of Knowledge by Landínez-Torres et al. (2020), members Cagáň, 2006).
of these subphyla were reported as the most abundant Although not yet known from Colombia, members of the
microfungi of Colombia from the soils in the Amazonian, Zoopagomycotina have bioprospective potential as biological
Caribbean, and Orinoquia regions (Landínez-Torres et al., controllers, as they can be found as nematode predators
2020). Owing to their agronomic relevance, arbuscular and mycoparasites (Spatafora et al., 2017). In the same
mycorrhizal fungi are being studied mainly in cultivated way, species in the Kickxellomycotina are found in the gut
plants, such as Physalis peruviana (Solanaceae), Manihot of arthropods that have an aquatic stage (Spatafora et
esculenta (Euphorbiaceae), Bactris gasipaes (Arecaceae), al., 2017). Owing to the high diversity of insects and other
and Euterpe oleracea (Arecaceae), among others (Molineros arthropods in Colombia, we expect that surveys of their
Hurtado et al., 2013; Gómez et al., 2020; Peña-Venegas et associated fungal diversity will reveal species of these
al., 2019; Ramírez-Gómez et al., 2019). fungal lineages in the country.
In a biogeographic database of arbuscular mycorrhizal The early-diverging lineages of the fungal kingdom,
fungi, 77 species were recorded for Colombia over the past the zoosporic fungi, are represented by nine species of
50 years, making Colombia one of the Neotropical countries Chytridiomycota and three of Blastocladiomycota. Among
with the highest number of reported species, except for the Chytridiomycota found in Colombia, six species
Brazil, with 182, and México, with 87 species. Countries are catalogued in the database as plant pathogens.
neighboring Colombia, such as Ecuador (28), Peru (52), and Batrachochytrium is the only vertebrate pathogen in this
Venezuela (37), had fewer species listed, indicating that phylum (Cepero et al., 2012), and in Colombia, only B.
in the Andean region, this group of micromycetes is better dendrobatidis is reported. The latter is one of the most
studied in Colombia. However, additional studies are still relevant fungal pathogens because it is responsible for
needed (Stürmer & Kemmelmeier, 2020). chytridiomycosis, a disease that threatens the amphibian
Also, six genera (Dissophora, Actinomortierella, diversity in Colombia and globally (Flechas et al., 2013).
Linnemannia, Podila, Entomortierella, and Mortierella) and In the Blastocladiomycota, Coelomomyces reticulatus
26 species of the Mortierellomycotina are reported in is an animal pathogen that attacks mosquitoes in
the ColFungi database. These species were found in soil particular (Gleason et al., 2010). No anaerobic fungi
samples (Veerkamp & Gams, 1983; Gualdrón-Arenas et (Neocallimastigomycota) have so far been registered in the
al., 1997; Landínez-Torres et al., 2019). Species in the ColFungi database. Members of this lineage are recognised
Morteriellaceae have been isolated worldwide from many for their outstanding lignocellulolytic activity (Solomon et
sources, including roots or decomposing plant material al., 2016).
(Vandepol et al., 2020). New species are expected to be Colombia is likely to be a promising source of zoosporic
described from countries like Colombia, where this fungal fungi due to its abundance of marine and freshwater
group is still underrepresented in listings (Vandepol et al.,
ecosystems. These underrepresented phyla deserve
2020). Members of this family, as well as members of the
more attention as they are expected to include emerging
orders Mucorales and Umbelopsidales (Mucoromycotina),
pathogens (e.g., Batrachochytrium spp., Synchytrium
are considered oleaginous (oil-producing) fungi, valued
spp.). The number of species waiting to be described in
in the food and bioenergy industries (Zhao et al., 2020),
the zoosporic fungi at a global level is high (Voigt et al.,
providing an additional incentive for the study of diversity
2013). The trophic modes associated with the microfungi
and distribution of these fungi in Colombia.
registered in ColFungi are essentially pathogens, either
Mucoromycotina is represented by 16 genera and 47
of plants (1,205 species) or animals (236 species), or
species, with Mucor being the most speciose genus, with 14
saprotrophs (450 species). In addition, 252 species are
recorded species in Colombia. Members of this subphylum,
reported as endophytes. This group of fungi is gaining
such as Absidia, Actinomucor, Circinella, Cuninghamella,
attention among the Colombian research community,
Rhizopus, and Syncephalastrum, are commonly isolated
possibly because of their bioprospecting potential (Strobel
from soil (Landínez-Torres et al., 2020). Apophysomyces
& Daisy, 2003; see Chapter 15).
variabilis (Rodríguez et al., 2018), A. elegans (Ruiz et al.,
2004), Sacksenaea erythrospora (Rodríguez et al., 2016),
Lichtheimia corymbifera, and several species of Rhizopus, CONCLUSIONS
are responsible for mucormycosis disease in humans, The diversity of Colombian micromycetes in the ColFungi
being associated with necrotizing fasciitis (Morales-López database is mainly represented by Ascomycota and
et al., 2018). Basidiomycota. The remaining phyla in the kingdom Fungi are
The Zoopagomycota comprises three subphyla: underrepresented or absent. We encourage the mycology
Zoopagomycotina, Kickxellomycotina, and Entomophthoro community to contribute to diversity assessments,
mycotina. In the ColFungi database, only the latter emphasising the early-diverging fungal lineages, such as
zygomycetes and zoosporic fungi, which need further studies of Bactris gasipaes Kunth in Buenaventura, Colombia. Acta Agronómica,
to better reflect the true diversity of microfungi in Colombia. 62(4), 344-351.
Morales-López S, Ceballos-Garzón A, Parra-Giraldo CM (2018) Zygomycete
In addition, we need to enrich our culture collections to fungi infection in Colombia: literature review. Current Fungal Infection
preserve microfungal diversity and to make it accessible for Reports, 12(4), 149-154. doi: 10.1007/s12281-018-0326-9
applied research. Furthermore, we encourage the use of a Peña-Venegas CP, Kuyper TW, Davison J, Jairus T, Vasar M, Stomph
polyphasic approach, choosing the best molecular markers TJ, Struik PC, Öpik M (2019) Distinct arbuscular mycorrhizal fungal
communities associate with different manioc landraces and Amazonian
for each genus, to assess the microfungal diversity and to soils. Mycorrhiza, 29(3), 263-275. doi: 10.1007/s00572-019-00891-5
achieve assertive identifications to species level. Ramírez-Gómez M, Pérez-Moncada U, Serralde-Ordoñez D, Peñaranda-
Rolón A, Roveda-Hoyos G, Rodriguez A (2019) Diversity of arbuscular
mycorrhizal fungi communities associated with cape gooseberry
References (Physalis peruviana L.) crops. Agronomía Colombiana, 37(3), 239-254.
doi: 10.15446/agron.colomb.v37n3.74008
Ainsworth GC (2008) Ainsworth & Bisby's Dictionary of the Fungi. London: Rodríguez JY, Morales-López SE, Rodríguez GJ, Álvarez-Moreno CA,
CABI. 771p. Ocampo W, Cepeda ML, Mora-Valderrama MA (2018) Necrotizing
Álvarez-Yela AC, Alvarez-Silva MC, Restrepo S, Husserl J, Zambrano MM, fasciitis caused by Apophysomyces variabilis in an immunocompetent
Danies G, Gómez JM, Barrios AFG (2017) Influence of agricultural patient. Medical Mycology Case Reports, 20, 4-6. doi: 10.1016/j.
activities in the structure and metabolic functionality of paramo mmcr.2017.12.002
soil samples in Colombia studied using a metagenomics analysis Rodríguez JY, Rodríguez GJ, Morales-López SE, Cantillo CE, Le Pape
in dynamic state. Ecological Modelling, 351, 63-76. doi: 10.1016/j. P, Álvarez-Moreno CA (2016) Saksenaea erythrospora infection
ecolmodel.2017.02.010 after medical tourism for aesthetic breast augmentation surgery.
Barta M, Cagáň LU (2006) Aphid-pathogenic Entomophthorales (their International Journal of Infectious Diseases, 49, 107-110. doi:
taxonomy, biology and ecology). Biologia, 61(21), S543-S616. doi: 10.1016/j.ijid.2016.05.032
10.2478/s11756-007-0100-x Ruiz CE, Arango M, Correa AC, Saider L, Restrepo A (2004) Fascitis
Cepero C, Restrepo S, Franco-Molano A, Cárdenas M, Vargas N (2012) necrosante por Apophysomyces elegans, moho de la familia
Biología de hongos. Bogotá DC: Ediciones Uniandes. 520p. Mucoraceae, en paciente inmunocompetente. Biomédica, 24(3), 239-
Flechas SV, Medina E, Crawford A, Sarmiento C, Cárdenas M, Amézquita 251.
A, Restrepo S (2013) Characterisation of the first Batrachochytrium Schoch CL, Seifert KA, Huhndorf S, Robert V, Spouge JL, Levesque CA,
dendrobatidis isolate from the Colombian Andes, an amphibian Chen W, Fungal Barcoding Consortium (2012) Nuclear ribosomal
biodiversity hotspot. EcoHealth, 10(1), 72-76. doi: 10.1007/s10393- internal transcribed spacer (ITS) region as a universal DNA barcode
013-0823-9 marker for Fungi. Proceedings of the National Academy of Sciences
Gleason FH, Marano AV, Johnson P, Martin WW (2010) Blastocladian U S A , 109(16), 6241-6246. doi: 10.1073/pnas.1117018109
parasites of invertebrates. Fungal Biology Reviews, 24(1-2), 56-67. doi: Schulz B, Boyle C (2005) The endophytic continuum. Mycological Research,
10.1016/j.fbr.2010.03.004 109(6), 661-686.
Gualdrón-Arenas C, Suárez-Navarro AL, Valencia-Zapata H (1997) Hongos Solomon KV, Haitjema CH, Henske JK, Gilmore SP, Borges-Rivera D, Lipzen
del suelo aislados de zonas de vegetación natural del páramo de A, Brewer HM, Purvine SO, Wright AT, Theodorou MK (2016) Early-
Chisacá, Colombia. Caldasia, 19, 235-245. branching gut fungi possess a large, comprehensive array of biomass-
Gómez SPM, Berdugo SEB, Mena RAM (2020) Occurrence of indigenous degrading enzymes. Science, 351(6278), 1192-1195. doi: 10.1126/
arbuscular mycorrhizal fungi associated with the rhizosphere of the science.aad1431
naidí palm in Colombia. Ciencia & Tecnología Agropecuaria, 21(3), 1-14. Spatafora JW, Aime MC, Grigoriev IV, Martin F, Stajich JE, Blackwell M
doi: 10.21930/rcta.vol21_num3_art:1275 (2017) The fungal tree of life: from molecular systematics to genome-
Hebert PD, Cywinska A, Ball SL, DeWaard JR (2003) Biological scale phylogenies. Microbiology Spectrum, 5(5), 5503. doi: 10.1128/
identifications through DNA barcodes. Proceedings of the Royal Society microbiolspec.FUNK-0053-2016
of London Series B: Biological Sciences, 270(1512), 313-321. doi: Strobel G, Daisy B (2003) Bioprospecting for microbial endophytes and
10.1098/rspb.2002.2218 their natural products. Microbiology and Molecular Biology Reviews,
James TY, Stajich JE, Hittinger CT, Rokas A (2020) Toward a fully resolved 67(4), 491-502. doi: 10.1128/MMBR.67.4.491-502.2003
fungal tree of life. Annual Review of Microbiology, 74, 291-313. doi: Stürmer SL, Kemmelmeier K (2020) The Glomeromycota in the Neotropics.
10.1146/annurev-micro-022020-051835 Frontiers in Microbiology, 11, 3200. doi: 10.3389/fmicb.2020.553679
Landínez-Torres A, Panelli S, Picco AM, Comandatore F, Tosi S, Capelli Vandepol N, Liber J, Desirò A, Na H, Kennedy M, Barry K, Grigoriev IV,
E (2019) A meta-barcoding analysis of soil mycobiota of the upper Miller AN, O'Donnell K, Stajich JE (2020) Resolving the Mortierellaceae
Andean Colombian agro-environment. Scientific Reports, 9(1), 1-12. phylogeny through synthesis of multi-gene phylogenetics and
doi: 10.1038/s41598-019-46485-1 phylogenomics. Fungal Diversity, 104(1), 267-289. doi: 10.1007/
Landínez-Torres AY, Becerra Abril JL, Tosi S, Nicola L (2020) Soil s13225-020-00455-5
microfungi of the Colombian Natural Regions. International Journal Veerkamp J, Gams W (1983) Los Hongos de Colombia - VIII some new
of Environmental Research and Public Health, 17(22), 8311. doi: species of soil fungi from Colombia. Caldasia, 13(65), 709-717.
10.3390/ijerph17228311 Voigt K, Marano AV, Gleason FH (2013) Ecological and Economical
Li Y, Steenwyk JL, Chang Y, Wang Y, James TY, Stajich JE, Spatafora Importance of Parasitic Zoosporic True Fungi. Agricultural Applications.
JW, Groenewald M, Dunn CW, Hittinger CT (2021) A genome-scale Berlin: Springer. Pp. 243-270.
phylogeny of the kingdom Fungi. Current Biology, 31(8), 1653-1665. Vélez N, Escandón P (2020) Multilocus sequence typing (MLST) of
doi: 10.1016/j.cub.2021.01.074 clinical and environmental isolates of Cryptococcus neoformans
Liu XZ, Wang QM, Göker M, Groenewald M, Kachalkin A, Lumbsch HT, and Cryptococcus gattii in six departments of Colombia reveals high
Millanes A, Wedin M, Yurkov A, Boekhout T (2015) Towards an genetic diversity. Revista da Sociedade Brasileira de Medicina Tropical,
integrated phylogenetic classification of the Tremellomycetes. Studies 53, e20190422. doi: 10.1590/0037-8682-0422-2019
in Mycology, 81, 85-147. doi: 10.1016/j.simyco.2015.12.001 Wang QM, Begerow D, Groenewald M, Liu XZ, Theelen B, Bai FY, Boekhout
Lücking R, Aime MC, Robbertse B, Miller AN, Ariyawansa HA, Aoki T, T (2015a) Multigene phylogeny and taxonomic revision of yeasts and
Cardinali G, Crous PW, Druzhinina IS, Geiser DM (2020) Unambiguous related fungi in the Ustilaginomycotina. Studies in Mycology, 81, 55-
identification of fungi: where do we stand and how accurate and precise 83. doi: 10.1016/j.simyco.2015.10.004
is fungal DNA barcoding? IMA Fungus, 11(1), 1-32. doi: 10.1186/ Wang QM, Yurkov A, Göker M, Lumbsch H, Leavitt S, Groenewald M, Theelen
s43008-020-00033-z B, Liu XZ, Boekhout T, Bai FY (2015b) Phylogenetic classification of
Molineros Hurtado FH, Mosquera-Espinosa AT, Gómez-Carabalí A, Otero JT yeasts and related taxa within Pucciniomycotina. Studies in Mycology,
(2013) Temporal variation in arbuscular mycorrhizal fungi colonization 81, 149-189. doi: 10.1016/j.simyco.2015.12.002
Neoscytalidium dimidiatum
[Álvaro Rúa]
Curvularia sp.
[Álvaro Rúa]
Chardoniella gynoxidis
[Mauricio Salazar-Yepes]
112 CATALOGUE OF FUNGI OF COLOMBIA
Chapter 9
Diversity of Rust and Smut Fungi of Colombia
Mauricio Salazar-Yepes1* & Meike Piepenbring2
1
Universidad Nacional de Colombia, sede Medellín, Facultad de Ciencias, Museo Micológico-MMUNM, Calle 59A, No. 63-20, Medellín, Colombia.
2
Mycology Working Group, Institute of Ecology, Evolution and Diversity, Faculty of Biosciences, Goethe University Frankfurt am Main, Biologicum, Max-von-
Laue-Str. 13, 60438 Frankfurt am Main, Germany.
ABSTRACT
Rust and smut fungi are biotrophic and obligate pathogens of vascular plants with a high species richness on plants growing
in the wild. This group comprises species that cause economically relevant plant diseases affecting cultivated plants. Rust
fungi are represented by Pucciniales (Pucciniomycotina, Basidiomycota), and smut fungi mostly comprise several orders
from Ustilaginomycotina (Basidiomycota) and some additional orders from other Basidiomycota subdivisions. The updated
checklist of rust fungi of Colombia includes 493 species, and the checklist for smut fungi 71 species, with an additional
ten species of Ustilaginomycotina found as yeasts or belonging to Exobasidiales. While the number of rust fungi known
for Colombia has increased during recent years, the knowledge on smut fungi has stagnated since 2002, with relatively
few new records of yeasts being added. Rust and smut records for different departments of Colombia show substantial
information in regions with large urban centres, universities, and active mycologists. By contrast, no records or systematic
surveys are available for many other regions of the country. Estimations of species diversity for the rust and smut fungi of
Colombia suggest that even though the knowledge regarding these fungi in Colombia has increased throughout the past
three decades, we are far from completely documented. Field surveys and analyses of specimens by traditional taxonomic
methods and the generation of molecular sequence data are urgently needed for rust and smut fungi of Colombia, including
closely related groups such as the Exobasidiales.
RESUMEN
Colombia es considerado un país biodiverso, lo cual se debe principalmente a su privilegiada ubicación geográfica,
localizado en la esquina noroccidental de América del Sur en la zona intertropical. Junto a una alta diversidad de plantas,
existe igualmente una elevada diversidad de los microorganismos asociados, entre ellos las royas y carbones, los cuales se
caracterizan por ser patógenos obligados de plantas vasculares con una alta diversidad de especies. Se encuentran sobre
plantas nativas y varias de sus especies pueden causar pérdidas económicamente relevantes en plantas cultivadas de
importancia económica como: arroz, avena, café, caña de azúcar, cebada, frijol, maíz, pinos, soja y trigo entre otras. Ambos,
hacen parte de la división Basidiomycota (Fungi), las royas pertenecientes al orden Pucciniales (Pucciniomycotina) y los
carbones a diversos órdenes de Ustilaginomycotina, así como algunos órdenes presentes en otros grupos relacionados. El
inventario de royas de Colombia incluye 493 especies, y el de carbones 71, además de 10 especies de levaduras del orden
Exobasidiales (Ustilaginomycotina). Si bien el número de especies de royas conocidas para Colombia aumentó durante los
últimos años, el conocimiento de los carbones se mantiene igual desde 2002. Solo se adicionaron algunos nuevos reportes
de especies de levaduras. Los registros de royas y carbones para diferentes departamentos de Colombia muestran que existe
un conocimiento relativamente bueno en las regiones con grandes ciudades, universidades y micólogos activos, mientras
que en otras regiones del país no se cuenta con estudios micológicos de campo relacionados con este importante grupo
de patógenos de plantas. Las estimaciones realizadas sobre la diversidad de especies de royas y carbones existentes en
el país confirman que estamos lejos de un registro completo de la Biota de royas y carbones de Colombia. Son necesarias
cuanto antes expediciones de campo que permitan recolectar en diferentes departamentos plantas nativas o introducidas
de diferentes familias botánicas parasitadas con estos hongos, para luego, realizar los métodos tradicionales de taxonomía
morfológica, así como la determinación de secuencias moleculares para las royas y carbones, y grupos relacionados como
los Exobasidiales. En Colombia, se hace necesario un mayor esfuerzo por estudiar las royas y carbones presentes, y de
esta manera aportar la mayor cantidad de información sobre la biodiversidad presente en las cinco regiones biogeográficas
del país, en las cuales tenemos ecosistemas frágiles, amenazados, y con una deforestación descontrolada en algunos de
ellos, lo que llevará a que muchas de las nuevas especies de royas y carbones no se conozcan. Este hecho, una vez más,
agrega un argumento adicional para la urgente necesidad de su conservación.
FIGURE 2. The life cycle of the rust fungus Puccinia graminis FIGURE 3. The life cycle of the smut fungus Ustilago maydis
(Pucciniales) on Berberis sp. (generations 0, I) and on Triticum (Ustilaginales) on corn (Zea mays). 1n: haploid, P!: plasmogamy,
aestivum (generations II and III). 1n: haploid, P!: plasmogamy, n+n: n+n: dikaryotic, K!: karyogamy, 2n: diploid, M!: meiosis. (Illustration
dikaryotic, K!: karyogamy, 2n: diploid, M!: meiosis. (Illustration by by M. Piepenbring).
M. Piepenbring).
recognized for species belonging to this order: spermogonia, to this developmental stage as a plant parasite, many smut
two anamorphs (aecia and uredinia), teleomorph (telia), fungi grow as yeasts during a saprotrophic developmental
and phragmobasidium (basidia). Different spores are stage. Smut fungi present a high diversity of ultrastructural
formed inside or by the abovementioned structures, such characteristics of septal pores and host plant interaction
as spermatia, aeciospores, urediniospores, teliospores, structures, reflecting complex systematic relationships
and basidiospores (Buriticá et al., 2014). Rust fungi can (Bauer et al., 2001). These micromorphological features,
comprise up to five or six morphological and functionally alongside macromorphological and molecular characters,
different types of spores within their life cycle, with several justify the current classification of smut fungi into different
species requiring up to two, usually unrelated species orders (Bauer et al., 2001, 2006; Begerow et al., 2018).
of host plants to complete their life cycle (Figure 2) Species of rust fungi present themselves as different
(Piepenbring, 2015). Additional ultrastructural characters stages of spores and smut fungi and as plant pathogens
that are typical for rust fungi are nuclei without spindle or yeasts. Until recently, it was permissible to apply more
pole bodies and simple septal pores with characteristic than one name to different developmental forms of the
caps formed by the endoplasmic reticulum (Bauer, 1987; same species. Nonetheless, since 2013, according to the
O’Donnell & McLaughlin, 1981). International Code of Nomenclature for Algae, Fungi and
Smut fungi are the second-largest parasitic group of Plants, only a single name is valid for all developmental
Basidiomycota, currently comprising more than 120 genera forms of a species. This rule has been applied for yeasts of
and ca. 1,700 species (Kruse et al., 2017). These fungi used smut fungi by Wang et al. (2015). In the case of rust fungi,
to be placed in one (Ustilaginales) or two (Ustilaginales, there are still many species with names that need to be
Tilletiales) orders (Vánky 2012). Owing to their polyphyletic revised according to this rule.
nature revealed by molecular phylogenetics, they are
currently placed into an increasing number of orders within MATERIALS AND METHODS
Ustilaginomycotina (e.g., Dossansiales, Entylomatales, We compiled the current knowledge on rust and smut
Georgefischeriales, Tilletiales, Urocystidales, Ustilaginales), fungi known for Colombia based on literature searches
alongside other plant-parasitic fungi (e.g., Exobasidiales) and on academic databases (i.e., Google Scholar, MMUNM),
some yeast-forming lineages, such as Malasseziales (Vánky, including keywords such as “Pucciniales”, “roya”, “smut
2012). Other lineages traditionally classified as smut fungi fungi”, “Tilletiales”, “Uredinales”, and “Ustilaginales”. All
belong to Microbotryales (Pucciniomycotina) or Entorrhizales retrieved data were compiled into an excel sheet, including
(Entorrhizomycetes) (Begerow et al., 2018). Host plants of the names of families, genera, and species of rust and
smut fungi are primarily herbaceous, with Poaceae and smut fungi reported for Colombia, besides the plant species
Cyperaceae harbouring most of the known smut fungi species they were found parasitizing. In addition, we compared our
(Piepenbring, 2015). The life cycle of smut fungi comprises results from the literature review with the database available
a single generation of spores (teliospores) developing on a at the ColFungi website (https://colfungi.org/) and updated
single host plant (Figure 3) (Piepenbring, 2015). In addition it whenever needed.
RESULTS AND DISCUSSION Frank D. Kern, who published their findings from 1927 to
History of the study and cataloguing of rust and smut fungi 1954 (Buriticá & Pardo-Cardona, 1996). In the second half
in Colombia of the 20th century and after the turn of the millennium
(~1968-2021), research on rust and smut fungi was mainly
The first observations of rust fungi in Colombia date back performed by Colombian researchers, such as Pablo Buriticá,
to those made in 1803 by Francisco José de Caldas, who Luis Alfredo Molina-Valero, Víctor Manuel Pardo-Cardona,
associated the «polvillo” (powder) of cereals with a parasitic and Mauricio Salazar-Yepes (Piepenbring, 2002; Buriticá et
fungus that consumes its nutrients, today known as the al., 2014). The second author of this chapter (MP) visited
wheat stem rust (Puccinia graminis) (Caldas, 1803). The first Colombia in 1998 and included her discoveries of smut fungi
record of a rust fungus in Colombia was based on specimens in an updated annotated checklist of smut fungi known for
collected by Triana and Planchon (Comisión Corográfica- this country (Piepenbring, 2002).
Colombia) identified as Trichobasis oxalidis (=Puccinia oxalidis)
parasitizing Oxalis pubescens (Oxalidaceae) collected in the Diversity of rust fungi in Colombia
department of Cundinamarca (Léveillé, 1867). There are three studies presenting catalogues of Pucciniales
After these initial observations, sporadic records of known from Colombia. The first study reported 204 species
rust and smut fungi were reported by foreign mycologists (Kern & Whetzel, 1930), the second 316 species (Buriticá &
throughout the 19th and early 20th centuries, until the first Pardo-Cardona, 1996), and the third 456 species (Buriticá
mycological expedition in Colombia was carried out by Mayor et al., 2014). The current diversity of Pucciniales of Colombia
(1913). As a result of this expedition, 158 species of rust fungi comprises 493 species (see Chapter 16). These species
(Pucciniales) were collected and reported for Colombia, 84 represent 67 genera (58 teleomorphs and nine anamorphs),
of which were new to science. Based on Mayor's collections, with Puccinia (226), Uromyces (60), and Phakopsora (18)
Sydow & Sydow (1914) also published new records of fungi being the most species-rich genera in Colombia. Four
for Colombia, including the first three species of smut fungi. teleomorph genera of Pucciniales have been described on
In 1926, Carlos Chardón and his colleague and compatriot the basis of Colombian specimens, namely Chardoniella,
Rafael Toro made numerous collections of fungi, including Chrysocelis, Hennenia (endemic to Colombia), and Uncol. In
several Pucciniales and smut fungi (Chardón & Toro, 1930). addition, 86 species of rust fungi have been described as
Samples collected were sent to Herbert H. Whetzel and new to science based on type specimens from Colombia.
A B
FIGURE 4. Departments of Colombia with numbers of known species. A Rust fungi. B Smut fungi.
Host plants parasitised by rust fungi in Colombia TABLE 1. Diversity of rusts in Colombia and other tropical countries.
comprise 88 vascular plant families (angiosperms and Numbers for smut fungi are very low and do not justify such an
ferns), with Asteraceae (host species for 173 species of rust analysis. a Hennen et al. (2005); Forzza et al. (2010); b Berndt
fungi), Poaceae (130 spp.), and Leguminosae (90 spp.) being (2008); 1 Bernal et al. (2019); 2 Flora do Brasil (2020).
the families with the highest number of reported species of
Pucciniales. According to Cárdenas-López & Salinas (2007), Costa South
Teleomorph Colombia Brazil
some of these host plants are threatened in Colombia, Rica Africa
genera n(%) n(%)a
including Quercus humboldtii (vulnerable). Their host-specific n(%)b n(%)b
parasite, Cronartium quercuum, is equally threatened (López Puccinia 226(46) 264(35) 136(45) 217(40)
& Salinas, 2007).
Uromyces 60(12) 107(14) 39(13) 103(19)
The first molecular phylogenetic study of rust fungi in
Colombia was carried out by Zuluaga et al. (2011), presenting Phakopsora 18(3) 28(3) 11(4) 8(5)
the first molecular sequences for specimens of rust fungi
from Colombia deposited in GenBank. There are sequences Ravenelia 9(2) 41(5) 13(4) 27(5)
for only 37 species of rust fungi, representing only 7% of Prospodium 7(1) 31(4) 10(3) –
the diversity currently occurring in Colombia. This figure is
a meagre percentage, considering the biological diversity of Total number of
493 745 300 546
this large group of plant pathogens. species
Number of vascular
Distribution of known species of rust fungi in Colombia 24,5301 49,9872 12,119 21,000
plant species
A distribution map highlighting the 32 departments of Rust plant ratio 1:49 1:67 1:40 1:39
Colombia is presented (Figure 4), with different colours
reflecting the number of species known for each department. Rust estimates
1,226 2,499 605 1,050
The Antioquia (324 species of rust fungi), Cundinamarca 1:20
(143 spp.), and Caldas (121 spp.) departments are 1:4 6,132 12,496 3,029 5,250
highlighted in green as they show the highest number of rust
species in Colombia. However, 35 out of 125 municipalities 1:9 2,725 5,554 1,346 2,333
in Antioquia (28%) have not yet been explored, lacking any
recorded specimen of rust fungi. No rust fungi have been Diversity of smut fungi of Colombia
reported yet for the five departments highlighted in red
According to the Checklist of Fungi of Colombia (Chapter 16),
(Guainía, Guaviare, La Guajira, Vaupés, and Vichada) (Figure
71 species of smut fungi are currently known on the basis
4). Lack of research and inadequate rust species surveys
of collections of infected plants identified by host species
in the remaining 24 departments suggest that the diversity
and morphology. This number has not increased since 2002,
of Pucciniales in Colombia might be higher than current
the year of the publication of the most recent checklist
estimations. Therefore, intensive research is required to
(Piepenbring, 2002). It is a relatively similar magnitude as
increase our understanding of these fascinating fungi.
the corresponding numbers published for other countries in
the Neotropics, such as Brazil (77), Costa Rica (52), Ecuador
Estimation of the diversity of species of rust fungi of Colombia
(37), and Panama (29) (Piepenbring, 1996, Piepenbring et
Based on studies carried out in Brazil, Central America, and al., 2011).
the United States, Hennen & McCain (1993) proposed that Among the 71 species of smut fungi reported for
the number of species of rust fungi in a region should be Colombia, 66 species belong to Ustilaginomycotina
between 5% (1:20) and 25% (1:4) of the number of plant (Ustilaginales: 42 species, Entylomatales: 8 spp., Tilletiales:
species in the same region. The current rust/plant ratio in 7 spp., Urocystidales: 5 spp., and Georgefischeriales: 4 spp.)
Colombia is 1:49 (Table 1). Using the 1:20 (5%) rust fungi/ and five to Microbotryales (Pucciniomycotina). No species
host estimate, the number of species for Colombia would be of Entorrhizales (Entorrhizomycetes) and Doassansiales
around 1,226, meaning that we currently know only 40% of (Ustilaginomycotina) are known from Colombia. The 71
the species present in this country. When applying the 1:4 species of smut fungi are placed into 25 genera, with
ratio, we obtain an even higher estimate of 6,132 species, Ustilago (11 species), Sporisorium (11 spp.), and Entyloma
of which we currently only know 8%, in tropical countries, (8 spp.) being the most diverse genera in Colombia. In
such as Brazil, only 6% of the existing species would be comparison to the species richness reported from temperate
known to science, which is similar for Costa Rica (9%) and regions (e.g., Europe; Vánky, 1994), the number of species
South Africa (10%) (Table 1). An intermediate ratio of 1:9 belonging to Sporisorium, infecting the plant family Poaceae
(11%) was proposed by Salazar-Yepes & Carvalho (2013). and Cintractia and related genera from the plant family
Based on this ratio, Colombia would have 2,725 species of Cyperaceae is relatively high for Colombia, while in regions
rust fungi (Table 1), of which only 18% are currently known. with a temperate climate, species of Anthracoidea dominate
Therefore, numerous species of tropical rust fungi might still infections on Cyperaceae and species of Ustilago on Poaceae
be unknown to Colombia. (Piepenbring, 2015).
Five species of smut fungi (Aurantiosporium colombianum, our present knowledge (71 species) will correspond to only
Entyloma galinsogae, Kuntzeomyces ruizianae, Thecaphora 12.5% of the existing diversity of smut fungi in Colombia.
polymniae, and Tilletia colombiana) were described as new
species based on specimens from Colombia and are still CONCLUSIONS
recognized as valid taxa. In addition to the aforementioned Regions that have never been visited for mycological surveys
species of smut fungi, five species of Ustilaginales and and estimates of species richness show that our knowledge
three species of Malasseziales are cited for Colombia as of the diversity of rust and smut fungi in Colombia is still
saprotrophic yeasts (Chapter 16). The order Exobasidiales is quite incomplete. Future additional studies are necessary
represented only by two species in the checklist. As evident to sample these fungi in all biogeographic regions of
by records published for neighbouring countries, such as Colombia to order to better understand the composition,
Panama (Piepenbring et al., 2010), many more species of species diversity, biology, life cycles, and geographic and
this group are undoubtedly present in Colombia. Host plants altitudinal distribution of rust and smut fungi, besides other
parasitized by smut fungi in Colombia belong to seven related fungi and their host plants. In addition, it is vital to
families of vascular plants, with Poaceae (34 species) and investigate morphological characters and to generate new
Cyperaceae (22 spp.) being the families with the highest DNA sequences for the collected specimens. Considering
numbers of reported species. All host plants are herbaceous, the elevated rates of deforestation in Colombia and the
except for Cissus spp. (Vitaceae), which are woody lianas lack of human and infrastructure resources available for
and hosts of the smut fungus Mycosyrinx cissi. research, several species of rust and smut fungi might
Several recent studies provided identifications of smut be already extinct and never be known to science. This
yeasts and non-smut fungi based on the sequences of single highlights the urgent need to preserve Colombia’s native
gene regions obtained from isolated yeasts (e.g., Gaviria & and endemic host plants alongside their rust and smut
Osorio, 2012) or in the context of metabarcoding studies (e.g., fungi species.
Větrovský et al., 2020). These records should be treated with
care as their identifications are not as reliable as those from
traditional taxonomic studies. They suffer from incomplete Acknowledgements
reference databases and improper identification algorithms,
We want to thank the curators of rust and smut herbaria in Colombia
such as percentage identities using predefined thresholds. for access to their collections, specimen loans, and literature
As far as we know, no study to date has ever focused on information; Laura Carolina Alvarez-Morales for elaborating the
generating DNA sequences of smut fungi collected in distribution maps; Bibiana Moncada and Robert Lücking for their
Colombia. In addition, thresholds for species delimitation revisions; and the Universidad Nacional de Colombia and Goethe
are not established (and not recommended, see Lücking University Frankfurt for their institutional support.
et al., 2020), and the ecological significance of a fungal
species detected only by the presence of its DNA cannot
be assessed. Nevertheless, environmental sequence data References
are helpful to learn about fungal diversity, distribution, and Aime MC, McTaggart AR (2021) A higher-rank classification for rust fungi,
ecological niches used by saprotrophic smut yeasts. with notes on genera. Fungal Systematics and Evolution, 7(1), 21-47.
doi: 10.3114/fuse.2021.07.02
Álvarez Morales L, Salazar Yepes M (2014) Caracterización morfológica de
Distribution of known species of smut fungi in Colombia las royas (Pucciniales) que afectan el limoncillo (Cymbopogon citratus
The distribution map indicating the numbers of species of smut (DC.) Stapf) en Colombia. Bioagro, 26(3), 171-176.
Andrade MG (2011) Estado del conocimiento de la biodiversidad en
fungi known for different departments of Colombia (Figure 4B)
Colombia y sus amenazas. Consideraciones para fortalecer la
was based on records published by Chardon & Toro (1930), interacción ciencia-política. Revista de la Academia Colombiana de
Molina-Valero (1980), and Piepenbring (2002). It shows Ciencias Exactas, Físicas y Naturales, 35(137), 491-508.
patterns similar to those documented for rust fungi because Barrera-Enríquez VP, Salazar-Yepes M (2019) New Pucciniales records
from Colombia, including Uredendo anthurii sp. nov. and Uromyces
of reasons similar to those cited for rust fungi. Maximum
colombiana sp. nov. Boletín Científico Centro de Museos Museo de
numbers of species of smut fungi are reported for Antioquia Historia Natural, 23(2), 95-105. doi: 10.17151/bccm.2019.23.2.4.
(24 species) and Cundinamarca (27 spp.), followed by Valle Bauer R (1987) Uredinales-germination of basidiospores and pycnospores.
del Cauca (14 spp.). There are no records for the Amazonas, Studies in Mycology, 30, 111-125.
Bauer R, Begerow D, Oberwinkler F, Piepenbring M, Berbee ML (2001)
Arauca, Bolivar, Guainía, and Guaviare departments.
Ustilaginomycetes. In: McLaughlin DJ, McLaughlin E, Lemke PA (eds).
Mycota VII. Systematics and Evolution. Berlin: Springer. Pp. 57-83.
Estimation of the diversity of smut fungi of Colombia Bauer R, Begerow D, Sampaio JP, Weiβ M, Oberwinkler F (2006) The
simple-septate basidiomycetes: a synopsis. Mycological Progress,
Using the numbers of species known worldwide (8,000 5(1), 41-66. doi: 10.1007/s11557-006-0502-0
species of rust fungi:1,650 species of smut fungi), we may Begerow D, McTaggart A, Agerer R, Frey W (2018) Engler’s Syllabus der
assume a ratio of 4.8:1 for these two groups. By applying Pflanzenfamilien, Part 1/3, Basidiomycota and Entorrhizomycota.
the same ratio to estimate the diversity of smut fungi for Stuttgart: Gebrueder Borntraeger. 471p.
Bernal R, Gradstein SR, Celis M (2019) Catálogo de plantas y líquenes
Colombia, based on the assumption of 2,725 species of rust de Colombia. Instituto de Ciencias Naturales, Universidad Nacional
fungi existing for Colombia (see above), 568 species of smut de Colombia, Bogotá. Available at: http://catalogoplantasdecolombia
fungi should exist in Colombia. If this calculation is correct, .unal.edu.co
Berndt R (2008) The rust mycobiota of southern Africa: species richness, Molina-Valero LA (1980) Los hongos de Colombia- VI. Reconocimiento e
composition, and affinities. Mycological Research, 112(4), 463-471. doi: identificación de Ustilaginales en Colombia. Caldasia, 13, 50-96.
10.1016/j.mycres.2007.05.005 Morin L, Evans KJ, Sheppard AW (2006) Selection of pathogen agents in
Buriticá P, Pardo-Cardona VM (1996) Flora uredineana colombiana. Revista weed biological control: critical issues and peculiarities in relation to
de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, arthropod agents. Australian Journal of Entomology, 45(4), 349-365. doi:
20(77), 183-236. 10.1111/j.1440-6055.2006.00562.x
Buriticá P, Salazar-Yepes M (2007) Nuevos registros de royas (Uredinales) O’Donnell KL, McLaughlin DJ (1981) Ultrastructure of meiosis in the hollyhock
potencialmente importantes en Colombia. Revista Facultad Nacional de rust fungus, Puccinia malvacearum I. Prophase I-Prometaphase I.
Agronomía-Medellín, 60(1), 3645-3655. Protoplasma, 108(3), 225-244.
Buriticá P, Salazar-Yepes M, Pardo-Cardona VM (2014) Pucciniales (Fungi), Otálora MA, Berndt R (2020) Overview of Puccinia (Uredinales) species
Royas de Colombia. Revista Facultad Nacional de Agronomía-Medellín, on Solanaceae. Nova Hedwigia, 3-4, 307-346. doi: 10.1127/nova_
67(2), 1-93. hedwigia/2020/0572
Caldas FJ (1803) Memoria sobre la nivelación de las plantas que se cultivan Pardo-Cardona VM (2006) Uredinales de plantas cultivadas de interés floral
en la vecindad del Ecuador. Colección general. In: Posada E (1912) en Colombia. Revista Facultad Nacional de Agronomía-Medellín, 59(1),
Obras de Caldas. Biblioteca Nacional de Historia Natural, 9, 85-95. 3335-3353.
Cárdenas-López D, Salinas NR (2007) Libro rojo de plantas de Colombia. Piepenbring M (1996) Smut fungi (Ustilaginales and Tilletiales) in Costa
Volumen 4. Especies maderables amenazadas: Primera parte. Leticia: Rica. Nova Hedwigia, 113, 1-155.
Instituto Amazónico de Investigaciones Científicas - SINCHI. Ministerio Piepenbring M (2002) Annotated checklist and key for smut fungi in
de Ambiente, Vivienda y Desarrollo Territorial. 232p. Colombia. Caldasia, 24, 103-119.
Chardón C, Toro RA (1930) Mycological explorations of Colombia. Journal of Piepenbring M (2015) Introducción a la Micología en los Trópicos. St. Paul:
the Department of Agriculture Porto Rico, 14, 195-369. APS Press. 376 p.
Cummins GB, Hiratsuka Y (2003) Illustrated genera of rust fungi, 3 ed. St. Piepenbring M, Espinoza J, Saldaña L, Cáceres O (2010) New records,
Paul: American Phytopathological Society Press. 225p. host plants, morphological and molecular data of Exobasidiales
Flora do Brasil 2020. Jardim Botânico do Rio de Janeiro. Available at: http:// (Basidiomycota) from Panama. Nova Hedwigia, 91, 231-242. doi:
floradobrasil.jbrj.gov.br/ 10.1127/0029-5035/2010/0091-0231
Forzza RC, Baumgratz JFA, Bicudo CEM, Canhos DAL, Carvalho-Jr AA, Costa Piepenbring M, Hofmann TA, Kirschner R, Mangelsdorff R, Perdomo O,
A, Costa DP, Hopkins M, Leitman PM, Lohmann LG, Lughadha EN, Maia Rodríguez Justavino D, Trampe T (2011) Diversity patterns of Neotropical
LC, Martinelli G, Menezes M, Morim MP, Coelho MAN, Peixoto AL, Pirani plant parasitic microfungi. Ecotropica, 17, 27-40.
JR, Prado J, Queiroz LP, Souza S, Souza VC, Stehmann JR, Sylvestre Salazar-Yepes M, Carvalho Jr AA (2013) Uredinales (rust fungi) biota of the
LS, Walter BMT, Zappi D (2010) Síntese da diversidade Brasileira. In: Parque Nacional do Itatiaia, Brazil: an analysis of composition, species
Catálogo de plantas e fungos do Brasil. Vol. 1. Available at: https://static. diversity and altitudinal distribution. Caldasia, 35(1), 165-176.
scielo.org/scielobooks/z3529/pdf/forzza-9788560035083.pdf Salazar-Yepes M, Pardo-Cardona VM (2020) Puccinia fumosa (Pucciniales)
Gaviria J, Osorio E (2012) Yeast diversity associated to inflorescences and the host Loeselia glandulosa (Polemoniaceae), new records for
of mango and flowers of ‚Lulo arboreo‘. Biotecnología en el Sector Colombia and South America. Caldasia, 42(1), 166-169. doi: 10.15446/
Agropecuario y Agroindustrial, 10(2), 160-169. caldasia.v42n1.73420
González D, Salazar-Yepes M (2019) Nuevos registros de Pucciniales sobre Sydow H, Sydow P (1914) Contribution à l’étude des champignons parasites
la familia Solanaceae en Colombia. Revista de la Facultad de Ciencias, de Colombie. Mémoires de la Société Neuchâteloise des Sciences
8(1), 20-30. doi: 10.15446/rev.fac.cienc.v8n1.70330 Naturelles, 5, 432-441.
Hennen JF, Figueiredo MB, de Carvalho Jr AA, Hennen PG (2005) Catalogue Vanegas-Berrouet KM, Salazar-Yepes M (2015) Adiciones a la biota
of the species of plant rust fungi (Uredinales) of Brazil. Rio de Janeiro: de Pucciniales sobre Fabaceae en Colombia. Revista Mexicana de
Jardim Botânico do Rio de Janeiro. Available at: http://www.jbrj.gov.br Biodiversidad, 86(4), 849-855. doi: 10.1016/j.rmb.2015.08.004
Hennen JF, McCain JW (1993) New species and records of Uredinales from Vanegas-Berrouet KM, Pardo-Cardona VM, Salaza-Yepes M (2018) Estado
the Neotropics. Mycologia, 85(6), 970-986. actual de los Pucciniales en páramos de Antioquia (Colombia), sobre
Hershenhorn J, Casella F, Vurro M (2016) Weed biocontrol with fungi: past, la familia Cyperaceae. Acta Biológica Colombiana, 23(1), 88-94. doi:
present and future. Biocontrol Science and Technology, 26(10), 1313- 10.15446/abc.v23n1.60640
1328. doi: 10.1080/09583157.2016.1209161 Vanegas-Berrouet KM, Salazar-Yepes M (2018) Novedades del orden
Kern FD, Whetzel Jr, HE (1930) Mycological exploration of Colombia. Pucciniales sobre la familia Asteraceae en paramos de Antioquia
Uredinales. Journal of Department of Agriculture, Puerto Rico, 14, 301- (Colombia). Caldasia, 40(2), 284-295. doi: 10.15446/caldasia.
348. v40n2.72053
Kruse J, Mishra B, Choi YJ, Sharma R, Thines M (2017) New smut-specific Vanegas-Berrouet KM, Salazar-Yepes M (2020) Nuevos registros
primers for multilocus genotyping and phylogenetics of Ustilaginaceae. de Pucciniales sobre Araceae, Bromeliaceae, Dryopteridaceae,
Mycological Progress, 16(9), 917-925. doi: 10.1007/s11557-017-1328-7 Grossulariaceae, Juncaceae, Lamiaceae y Poaceae en páramos de
Léveillé JH (1867) Fungi. In: Triana J, Planchon JE (eds). Prodromus florae Antioquia, Colombia. Revista de la Academia Colombiana de Ciencias
novo-granatensis: Cryptogamie vol 2. Paris: V. Masson et fils. 382p. Exactas, Físicas y Naturales, 44(170), 227-240. doi: 10.18257/
López-Alzate LM, Salazar-Yepes M (2017) Pucciniales sobre la familia raccefyn.986
Bignoniaceae en Colombia. Revista Mexicana de Biodiversidad, 88(3), Vánky K (1994) European smut fungi. Stuttgart: Gustav Fischer. 570p.
487-495. doi: 10.1016/j.rmb.2017.06.001 Vánky K (2012) Smut fungi of the world. St. Paul: APS press. 1458p.
Lücking R, Aime MC, Robbertse B, Miller AN, Ariyawansa HA, Aoki T, Větrovský T, Morais D, Kohout P, Lepinay C, Algora C, Hollá SA, Baldrian P
Cardinali G, Crous PW, Druzhinina IS, Geiser DM, Hawksworth DL (2020) GlobalFungi, a global database of fungal occurrences from high-
(2020) Unambiguous identification of fungi: where do we stand and how throughput-sequencing metabarcoding studies. Scientific Data, 7(1),
accurate and precise is fungal DNA barcoding? IMA Fungus, 11(1), 1-32. 1-14. doi: 10.1038/s41597-020-0567-7
doi: 10.1186/s43008-020-00033-z Wang QM, Begerow D, Groenewald M, Liu XZ, Theelen B, Bai FY, Boekhout
Mayor E (1913) Contribution a l’étude des Uredinées de Colombie. Mémoires T (2015) Multigene phylogeny and taxonomic revision of yeasts and
de la Société des Sciences Naturelles, 5, 442-509. related fungi in the Ustilaginomycotina. Studies in Mycology, 81, 55-83.
McTaggart AR, Shivas RG, van der Nest MA, Roux J, Wingfield BD, Wingfield doi: 10.1016/j.simyco.2015.10.004
MJ (2016) Host jumps shaped the diversity of extant rust fungi Zea-Fernández C, Gutiérrez-Castaño MC, Salazar-Yepes M (2021) Nuevos
(Pucciniales). New Phytologist, 209(3), 1149-1158. doi: 10.1111/ registros de Pucciniaceae (Pucciniales, Fungi) para Colombia. Revista
nph.13686 de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales,
McTaggart AR, Prychid CJ, Bruhl JJ, Shivas RG (2020) The PhyloCode 45(175), 500-507. doi: 10.18257/raccefyn.1348
applied to Cintractiellales, a new order of smut fungi with unresolved Zuluaga C, Buriticá P, Marín M (2011) Filogenia de hongos roya (Uredinales)
phylogenetic relationships in the Ustilaginomycotina. Fungal Systematics en la zona andina colombiana mediante el uso de secuencias del ADN
and Evolution, 6, 55-64. doi: 10.3114/fuse.2020.06.04 ribosomal 28S. Revista de Biología Tropical, 59(2), 517-540.
Cora elephas
[Robert Lücking]
120 CATALOGUE OF FUNGI OF COLOMBIA
Chapter 10
A Critical Assessment of Biogeographic Distribution Patterns of Colombian Fungi
Aída M. Vasco-Palacios 1,2,3*, Robert Lücking4, Bibiana Moncada3,5, Melissa Palacio6,7 & Viviana Motato-Vásquez3,8,9
1
Grupo de Microbiología Ambiental y Grupo BioMicro, Escuela de Microbiología, Universidad de Antioquia UdeA, Calle 70 No. 52–21, Medellín 050010, Colombia.
2
Fundación Biodiversa Colombia, Carrera 22 # 41 - 80 Apto. 004, 111311 Bogotá D.C., Colombia.
3
Asociación Colombiana de Micología, ASCOLMIC.
4
Botanischer Garten und Botanisches Museum, Freie Universität Berlin, Königin-Luise-StraßeStraße 6–8, 14195 Berlin, Germany.
5
Licenciatura en Biología, Universidad Distrital Francisco José de Caldas, Cra. 4 No. 26D-54, Torre de Laboratorios, Herbario, Bogotá D.C., Colombia.
6
Laboratorio de Taxonomía y Ecología de Hongos, TEHO, Instituto de Biología, Facultad de Ciencias Exactas y Naturales, Universidad de Antioquia UdeA,
A.A.1226, Medellín, Colombia.
7
Laboratório de Micologia, Departamento de Botânica, Instituto de Biociências, Universidade Federal do Rio Grande do Sul, Av. Bento Gonçalves, 9500,
Prédio 43.433, Campus do Vale, Agronomia, 91501-970, Porto Alegre, Rio Grande do Sul, Brasil.
8
Grupo de Investigación en Biología de Plantas y Microorganismos (BPM), Departamento de Biología, Facultad de Ciencias Naturales y Exactas, Universidad
del Valle, Calle 13 # 100-00 Cali, Valle del Cauca, Colombia.
9
Calima, Fundación para la Investigación de la Biodiversidad y Conservación en el Trópico, Cali, Colombia.
ABSTRACT
This chapter attempts to assess the distribution patterns of three selected groups of fungi, namely polypores, ectomycorrhizal
fungi (EcM), and lichenised fungi in Colombia. Assessing the biogeography of fungi is difficult because of two key issues:
1. information gaps on their distribution and biology in biodiversity-rich countries, such as Colombia, and 2. the traditional,
phenotype-based species concepts, which make it difficult to recognise cryptic species or species complexes. This latter
aspect is very frequent in fungi, as currently revealed by advanced molecular biology and phylogenetic analysis techniques. For
instance, Polyporus is a widespread genus, commonly found in Colombia with numerous species recorded as cosmopolitan.
However, detailed studies in some Polyporus sensu lato in Brazil and Argentina showed a hidden diversity now accommodated in
different genera such as Atroporus, Neodictyopus, and Bresadolia. On the other hand, the various fungal lineages show different
distribution patterns depending on their biology. This fact evidences how the biogeographic distribution of ectomycorrhizal
symbiont fungi (EcM) is closely linked with the distribution patterns of their host plants, but due to information gaps, their
real distribution ranges are unknown. It has been observed that the EcM fungi associated with Fagaceae in the Andean
region belong to Holarctic lineages, while the species associated with the Fabaceae or Dipterocarpaceae in the Colombian
Amazonia Region come from tropical lineages of Gondwanan origin. Finally, for lichenised fungi, we looked at eight genera
in four families and two classes and phyla: Bunodophoron (Sphaerophoraceae), Neoprotoparmelia (Parmeliaceae), Crocodia,
Lobariella, Podostictina, Pseudocyphellaria, Sticta (Peltigeraceae, all Ascomycota), and Cora (Hygrophoraceae, Basidiomycota).
Comparing traditional taxonomy with modern taxon concepts derived from integrative approaches using molecular and
phenotype data revealed that the proportion of distribution types assessed from the data changed substantially. When using
a traditional taxon concept, 45% of the species were inferred to have a broad, intercontinental distribution, 39% neotropical,
and 12% endemic. On the other hand, using the modern taxon concept indicated that only 4% of them were widespread, 20%
neotropical, and 76% potentially endemic. These findings underline the importance of accurate taxon concepts and proper
knowledge of evolutionary relationships when performing biogeographical analyses of Colombian fungi. As mycologists,
we must then continue to generating information that allows us to understand the historical processes responsible for the
geographical distributions of the different lineages of fungi present in the national territory.
RESUMEN
La biogeografía se encarga de los patrones de distribución global de los organismos, dilucidando los factores y procesos
que conducen a estos patrones. Los estudios en esta área se han centrado en organismos macroscópicos, como plantas
vasculares y vertebrados, dejando de lado microorganismos como bacterias, protistas, algas, plantas no vasculares y hongos.
Por mucho tiempo se consideró que los microorganismos suponían rangos de distribución amplios e intercontinentales para
muchas especies. El «todo está en todas partes» se aplicó durante décadas al pensar en la distribución de la mayoría de
los hongos. Sin embargo, los enfoques filogenéticos moleculares, combinados con el estudio de distribuciones y de las
comunidades de hongos, han dejado ver que la biogeografía de los hongos es compleja. Estos procesos dependen tanto de
la biología de las especies como de modelos de dispersión (expansión de las distribuciones) y vicarianza (fragmentación de
las distribuciones), así como de la historia geológica de los continentes, i.e., migración de los hongos a través de puentes
continentales, y la disyunción relictual continental. Muchas especies consideradas de distribución cosmopolita representan
complejos de especies difíciles de diferenciar con caracteres morfológicos, pero visibles en análisis filogenéticos. Este es el
caso del basidiolíquen Cora y de las especies del grupo neotropical Polyporus. Como se ha explicado en capítulos anteriores, el
conocimiento que tenemos hoy en día acerca de la diversidad, ecología, distribución, asociaciones y estado de conservación de
los hongos en Colombia es aún incipiente, y esto dificulta entender y definir patrones de distribución de las diferentes especies
presentes en el país. En este capítulo hacemos un primer acercamiento para evaluar los patrones de distribución de los hongos
poliporoides comparando la riqueza y composición de las especies. Por ejemplo, en el genero Polyporus son evidentes los
sesgos existentes por la falta de estudios en el género Polyporus muestra los sesgos existentes por la falta de estudios que
integren datos moleculares y análisis filogenéticos, este es el caso de especies que se pensaban ampliamente distribuidas
como Polyporus udus y Polyporus dictyopus, las cuales se consideran complejos de especies e incluso representan géneros
nuevos neotropicales, como es el caso de Neodictyopus. Por otro lado, diversos linajes de hongos muestran diferentes patrones
de distribución. Así es como la distribución biogeográfica de los hongos simbiontes ectomicorrízicos (EcM) está íntimamente
ligada con los patrones de distribución de sus plantas hospederas, sin embargo, debido a los vacíos de información no se
conocen los rangos reales de distribución. Se han reportado un total de 202 especies de hongos EcM en Colombia, de ellas,
56 son especies nuevas descritas a partir de especímenes colombianos y 36 son endémicas, lo que representa una tasa de
endemismo de cerca del 20%. Sin embargo, hay que confirmar la distribución de muchas especies con registros antiguos que
se conocen de pocos o de un único espécimen. En general se ha observado que los hongos EcM asociados a Fagaceae en la
región Andina pertenecen a linajes holárticos, mientras que las especies asociadas a las familias Fabaceae o Dipterocarpaceae
en la Amazonía Colombiana provienen de linajes tropicales de origen Gondwanico. Por último, se realizó un análisis con ocho
géneros de líquenes y comparando conceptos de la taxonomía tradicional con conceptos de taxón modernos; i.e. derivados de
enfoques integradores con datos moleculares y fenotípicos. Encontramos que la proporción de tipos de distribución evaluados
a partir de los datos cambió sustancialmente. Al utilizar un concepto tradicional, se infirió un concepto tradicional, se infirió
que el 45% de las especies tienen una amplia distribución intercontinental, 39% neotropical y 12% endémica. Mientras
que, siguiendo un concepto moderno, solo el 4% tiene distribución amplia, el 20% neotropical y 76 % son potencialmente
endémicas. Estos hallazgos subrayan la importancia de conceptos taxonómicos precisos y un conocimiento adecuado de las
relaciones evolutivas al realizar análisis biogeográficos de hongos colombianos. Como micólogos, debemos seguir generando
información que nos permita comprender los procesos históricos responsables de las distribuciones geográficas del pasado
al presente de los diferentes linajes de hongos en el territorio nacional.
FIGURE 1. A–B Schizophyllum commune (split gill fungus) photographed in Colombia (A) and Germany (B). C–D Sticta fuliginoides,
photographed in Colombia (C) and New Zealand (D). (Photographs by Robert Lücking.)
(Moncada et al., 2014, 2020; Magain & Sérusiaux, 2015). BIOGEOGRAPHY OF THE POLYPORES OF COLOMBIA
Yet, the true S. fuliginosa remains a subcosmopolitan Polypores are a highly diverse group of fungi characterised
taxon, and a newly recognised taxon within this complex, by their growth form and ecology (Figure 2a–f), representing
Sticta fuliginoides, is also subcosmopolitan (Figure 1c–d). In 2,300 species from the 21,000 species of Agaricomycetes
microfungi, the situation is challenging: human-pathogenic within Basidiomycota (Kirk et al., 2008). Species of
fungi and those attacking widely utilised crops tend to polypores are found in various orders but are concentrated
become widespread following human-induced expansions of mainly in Polyporales and Hymenochaetales, which together
the host range and usually evolve into new regional lineages contain around 80–90% of all polypores. These fungi are
(Summerell et al., 2010). most diverse in forest ecosystems on all continents and
These issues make it extremely difficult to assess the are morphologically characterised by a typically poroid
overall biogeographical patterns of fungi in a biodiversity- hymenophore and bracket-shaped to resupinate basidiomata.
rich country, such as Colombia, when relying primarily on They almost exclusively occur as saprotrophs or parasites on
phenotype-based or other taxonomic concepts, besides the dead or living trees, not rarely first attacking living trees and
fact that we do not know the total diversity of the Colombian then continuing as saprotrophs on the dead trees. Only a few
funga and the distribution of these species within and species grow on humus or mineral soil, and even fewer are
outside the country. This chapter refrains from deriving mycorrhizal (Väisänen et al., 1992; Tedersoo et al., 2007).
biogeographical patterns from the entire list of known fungal Polypores are the most critical wood decomposers, playing
species. Instead, we focus on selected groups and specific a pivotal role in forest ecosystems and their food webs by
examples to assess the realistic distribution patterns of recycling wood, the most critical reservoir of organic carbon
species present in Colombia. in the living world (Watkinson et al., 2006, Krah et al., 2018).
A B
D E
FIGURE 2. Fresh basidiomes of polypores. A–B Polyporaceae family. A Polyporus tricholoma. B Polyporus
dictyopus species complex. C Hydnopolyporus fimbriatus (Irpicaceae). D Ganoderma resinaceum
(Ganodermataceae). E Flaviporus liebmanii (Steccherinaceae). F Hymenochaete iodina (Hymenochaetaceae).
(Photographs A, D by Melissa Palacio; B–C, E–F by Viviana Motato-Vásquez).
The species richness and composition of polypores all polypores recorded from Colombia, published through
are influenced by climate and tree species composition in the ColFungi project (Gaya et al., 2021; https://colfungi.org),
forest ecosystems. Higher tree species diversity may result assembling all (reliable) fungi species records for Colombia
in higher polypore species richness (Hattori, 2017), and available in the taxonomic and biodiversity literature. With
tropical forests are recognised for their high diversity of tree this dataset, we tried to answer accurately questions such
species – commonly hundreds of species within 50 ha areas as “where does this polypores species occur in Colombia?”
(Condit et al., 2000). However, assessing host specificity Or even “how many polypore species are known to occur in
can become a challenge in such settings because it may this specific part (department, region, etc.) of Colombia?”
be difficult to identify host species reliably, mainly when the Although we are still far from completing this goal, we hope
basidiomata are produced on dead trees, fallen trunks, or to show the potential of our initiative in this case study.
high up in mature trees. Most polypore species are rare in A total of 84 genera and 223 polypore species have
species-rich tropical forests, whereas there is little evidence been recorded from different Colombian biomes (Figure 3a).
for host specificity in more common species (Lindblad, 2000; However, the representation of data in this kind of map is
Gilbert et al., 2002). Preferences for distinctive habitat types limited as a tool to represent the distribution of species
may be more important than host specificity in determining records (instead of representing species distribution per se).
the distribution of polypores (Lindblad, 2001). Each dot on this map can represent either a single record
Despite significant efforts to characterise the richness of a species or many records of several species collected at
of Neotropical funga, resulting in essential checklists and the same locality. Our data showed a high number of species
inventories of Neotropical polypores (e.g. Carranza & Ruiz- with a single record (35.7% of the records, 80 species),
Boyer, 2005; Silveira & Wright, 2005; Robledo & Rajchenberg, which could be understood more as an index of sampling
2007; Baltazar & Gibertoni, 2009; Vasco-Palacios & Franco- bias rather than as an indicator of the level of endemism.
Molano, 2013), the distribution patterns of polypores
remain poorly known. Biogeographic studies on Neotropical A
polypores are scarce. For instance, De Lima et al. (2018) used
polypores growing in the Brazilian rainforests to reconstruct
biogeographical relationships between the Amazon Forest,
Atlantic Forest, and Caatinga domains. Their results showed
that polypores have distribution patterns similar to those of
woody plants present in Brazilian rainforests. Another study
was done with Phellinotus piptadeniae in the Neotropics
(Salvador-Montoya et al., 2015). This species is a parasitic
polypore, causing heart-rot in some Fabaceae, with a disjunct
distribution in moist and dry forests, including Argentina,
Brazil, and Peru (Elias et al., 2020). The study emphasised
the importance of using host distributions in biogeographical
analyses of parasitic polypores to assess distribution
patterns. This fact illustrates another knowledge gap in
Neotropical polypores, namely species biology, as it is often
not known whether a species is parasitic or saprotrophic (or
both), whether it causes white or brown rot, and whether it is
a specialist or a generalist. B
Based on accurate taxon concepts, the geographic
distribution of species provides the data for biogeography,
macroecology, and conservation science assessments.
Species distributions are typically assessed through point
records, either through human observation or ideally
accompanied by voucher specimens that are permanently
stored in scientific collections. Unfortunately, data about
the geographic distribution of most fungi species known
from Colombia are still insufficient, generating the so-called
Wallacean knowledge shortfall, defined by the fact that
geographical distribution for the majority of taxa is poorly
FIGURE 3. A Distribution and occurrence of records of polypore
understood and contains many gaps (Hortal et al., 2015). This
fungi in Colombia. Each dot represents at least one record of a
situation is particularly difficult for species less well-known single species. B Main areas of the geographical distribution of the
than more charismatic species. Consequently, any effort 223 species of polyporoid fungi registered in Colombia. Blue tones
to compile and analyse distributional data for Colombian indicate species distributed at the American continent,whereas
polypores must be considered preliminary at this point. species distributed in two continents are represented by red tones,
We used a large, recently compiled database containing and those distributed in more than two continental areas in green.
In addition, we provide a first quantitative approach the Andes biome, compared with that in other Colombian
to determining the distribution patterns of Colombian biomes, may also result from the evolutionary history and
polypores based on the assessment of 223 species (Figure geo-climatic characteristics of this biome. Currently, in
3b). Our results showed that most species are currently Colombia, nine species of polyporoid fungi are recognised
reported with a Neotropical distribution (34.1%), followed by as endemic: three of them are described from the Andean
Pantropical (27.1%) and Cosmopolitan (25.1%) distributions, region, three from the Pacific, two from the Amazon, and one
whereas endemic species represent only 4%. However, this from the Caribbean. It is very premature to assess which
information may not be accurate because the taxonomic region supports the greatest richness of endemic species
and phylogenetic concepts of many species in the group with our data. For this reason, although logic indicates
are still debated. In many cases, it has been shown that that we should focus on sampling poorly known places, it
species believed to be widely distributed represent species is important to consider that better-sampled and species-
complexes, which is a factor that contributes to substantial rich areas, such as the Andes biome, may still hold many
gaps and bias in the assessment of fungal biogeography. undescribed and narrowly distributed species.
For instance, in recent decades, phylogenetic studies
have shown that traditional genera, such as Polyporus, are BIOGEOGRAPHY OF ECTOMYCORRHIZAL FUNGI IN COLOMBIA
polyphyletic, and several new genera have been segregated
Ectomycorrhizal (EcM) fungi present different patterns of
or reinstated since then. For example, Polyporus udus,
symbiotic associations with diverse families of Angiosperms
originally described from Indonesia, has been widely
and Gymnosperms (Brundett & Tedersoo, 2018; Corrales et
recorded in the Neotropics. Motato-Vásquez et al. (2018)
al. 2018). Owing to the obligate nature of these relationships,
investigated the phylogenetic relationship of P. udus and
the occurrence of EcM fungi generally coincides with
its purported taxonomic synonyms in South America. This
study showed that specimens from Argentina, Brazil, and the distribution of their associated plant families, which
Paraguay are not conspecific with Paleotropical specimens renders plant distributions critical to understanding the
of P. udus. The Neotropical records are now recognised as the biogeography and abundance of EcM fungi. However,
Bresadolia paradoxa. There are several records of P. udus processes of dispersal and migration in EcM fungi are
in Colombia, and future studies should attempt to untangle not always known. Recent studies have provided data to
these specimens’ true identity, especially because it is help to elucidate apparent disjunct distribution patterns,
a species of importance as a food source for Amazonian with possible scenarios including long-distance dispersal,
indigenous tribes (Sanuma et al., 2016). community migration across land bridges (with possible
In the same way, Polyporus dictyopus is a species that symbiont exchange), relictual continental disjunction and
has been recorded as Cosmopolitan and presents a large multi-hosts (Moser & Horak, 1975; Halling, 1996; Halling
number of heterotypic synonyms (with at least 16 known et al., 2008; Hosaka et al., 2008; Lumbsch et al., 2008;
from tropical and subtropical America). Palacio et al. (2017) Matheny et al., 2009; Hackel et al., in press).
showed that at least five distinct species were hidden under In Colombia, the best - studied ectomycorrhizal systems
P. dictyopus. The authors accommodated these species are those associated with the dominant tree in Andean
in two different genera, Atroporus and Neodictyopus. The Mountain ranges, Quercus humboldtii (Fagaceae) (Vargas &
study only included samples from Brazil, but P. dictyopus Restrepo, 2020; Peña-Vanegas & Vasco-Palacios, 2019).
has been widely recorded from the Amazonian, Andean, and There is another Fagaceae that forms monodominant EcM
Pacific regions in Colombia. However, the identity of these systems, Trigonobalanus excelsa, but this species is poorly
specimens remains unknown. studied (Peña-Vanegas & Vasco-Palacios, 2019). In the
Our compiled data show that the highest concentration of Amazonian region in Colombia, studies have been carried
recorded polypores is in the Andes region within Colombia. out on the diversity of EcM fungi associated with the host
This fact is not surprising since taxonomists tend to collect tree Pseudomonotes tropenbosii (Dipterocarpaceae) and
more intensively in the vicinity of their workplace. So, areas the Fabaceae hosts Aldina sp. and Dicymbe uaiparuensis
near important research institutions tend to show a higher (Vasco-Palacios et al., 2018; Peña-Vanegas & Vasco-
concentration of species distribution records (Sobral & Palacios, 2019). Most of the research on EcM fungi in
Stehmann, 2009). Most of the undercollected regions Colombia has focused on inventories in few localities in
contain diverse habitats that are suitable for polypores, the Andean and Amazon regions, so we do not know the
so the absence of records indicates strong sampling bias total distribution, biology, or ecology of EcM species in the
(Figure 3a). Nonetheless, sampling bias may not be the country (Chapter 4).
only explanation for the high diversity of polypores in the A total of 207 species of EcM fungi are known for Colombia.
Andean region. This biome is known for its high diversity Of those, 56 are new species described from Colombian
and endemism for many animals and plant taxa (Mutke et specimens, including 36 putative endemics, representing
al., 2014, and references therein). The high biodiversity a rate of endemism of nearly 18% (Table 1, Figure 4 a–f).
of this biome is possibly related to its topographic and Most of these species are associated with Fagaceae in
hydrologic heterogeneity, which resulted in species with the Andean region of the country (43 species), and about
narrow distribution ranges and the differentiation of small 55% of these new species are endemic (24 species) to the
areas of endemism. Thus, the high diversity of polypores in country. Most of these endemic species are known only from
TABLE 1. Data about new species of EcM fungi described from Colombian specimens based on their hosts, including the total of endemic
species. Biogeographic regions are based on biogeographic realms (https://ecoregions2017.appspot.com/)
the type specimen, or, in some cases, are known just from humboldtii being the only species present in South America,
few collections from the type locality (e.g. Lactarius caucae restricted to the Serranía del Darien, a small mountain
Singer, Russula idroboi Singer, Boletus orquidianus Halling). range on the border with Panama, and the Andes Mountains
Even though mycological expeditions have been conducted in Colombia (Hooghiemstra, 2006; Rangel & Avella, 2011).
in Antioquia, Boyacá, Santander, and Tolima in the past 20 Floristic studies in Neotropical Central and South America
years, those did not yield any additional specimens and show patterns similar to that of the fauna concerning the
localities for most of these endemic species (Vargas & Great American Biotic Interchange (GABI), which started with
Restrepo, 2020; Peña-Vanegas & Vasco-Palacios, 2019). It the closure of the Panama Isthmus approximately 3.0 Mya
is important to consider that a large number of unidentified (Leigh et al., 2014; O’Dea et al., 2016). Plant communities in
fungal specimens are deposited in fungal collections around montane regions of Central America and the northern portion
of the country. For example, in the fungal collection of the of South America are primarily composed of lineages of
Herbarium of the Universidad de Antioquia (HUA), which is the either Holarctic or Austral-Antarctic origin that subsequently
largest in the country, houses more than 12,500 specimens, dispersed south- or northwards. Phytogeographically, Costa
of which only 49% are identified at the species level (David Rica and Panama are more similar to Colombia than either
et al. 2019; Chapter 14). In addition, most of the endemic are to Mexico (Kappelle, 2006). Quercus and its associated
species have no associated barcode sequences that would EcM fungi migrated southwards across the Panama Isthmus
allow corroboration of their inferred restricted distribution. during the last glaciation. The remaining populations are now
Based on the information on the distribution of the geographically isolated because of the absence of cooler
species available on GBIF (http://www.gbif.org), the 207 environments, like those from high mountains, connecting
EcM species have a primarily Neotropical distribution (44%), Panama and Colombia (Halling 1996; Hooghiemstra, 2006;
of those 21% occur in lowland forests in South America, and Halling et al., 2008). Regardless of the available research
16% in mountain areas. Near one third of the species have on Neotropical oak communities, much of the taxonomy,
nordic distribution (14% Nearctic-Neotropical and 12.6% diversity, and biogeographic history of their associated
holarctic-Neotropical). Regarding the EcM species hosted EcM communities in Colombia remains to be explored in-
by Fagaceae, 18% are endemic, 37% have a Holarctic- depth. This lack of knowledge is even more pronounced
Neotropical (17.6%) or Nearctic-Neotropical (19.6%) for Trigonobalanus excelsa. The other two known species
distribution, and only 22% are Neotropical from mountain of Trigonobalanus, are from SE Asia and this Neotropical
areas. In the case of EcM fungi from tropical lowland forests, species is an example of tropical Amphi-Pacific disjunctions
11% are endemic, and 76.8% occur in Neotropical-South (van der Hammen & Cleef, 1983). Therefore, it is very
America (Figure 5A). interesting to know the EcM fungi lineages associated with T.
Quercus (Fagaceae) is an important EcM plant host with excelsa, which could also have a Pantropical origin, contrary
Holarctic distribution. In Costa Rica, Panama, and Colombia, to those lineages associated with Quercus humboldtii, which
oak forests are primarily found in tropical montane has Holactic origins. Studies of EcM agarics and boletes
ecosystems. The presence of oak forests in Colombia from Quercus forests in Costa Rica and Colombia indicate
represents the southernmost limit of the geographic genus-level affinities with the Northern Hemisphere (59% of
distribution of Quercus in the Americas, with Quercus all EcM species) rather than those from tropical lowlands
FIGURE 4. Endemic species of EcM fungi. A Russula floriformis subsp. floriformis, symbiont with Quercus humboldtii.
B Phylloporus fibulatus symbiont with Q. humboldtii. C Sarcodon rufobrunneus associated with Dicymbe uaiparuensis.
D Gloeocantharellus uitotanus putatively associated with Pseudomonotes tropenbosii. E Austroboletus amazonicus
associated with P. tropenbosii. G Ramaria chocoënsis without information about the possible plant host.
(Photographs A by Adriana Corrales; B–E by Aída Vasco-Palacios; and G by Danny Newman.)
and temperate areas in South America (22%) (Halling et during the Pleistocene (Vera et al., 2021). While there are
al., 2008) (Figure 5b). These groups also tend towards high some specific examples, in general there is a big gap in the
endemism at the species level (Halling et al., 2008). In knowledge of EcM fungi, particularly those associated with
Russulaceae, a highly diverse family of EcM fungi, Andean Fagaceae. For example, the EcM diversity associated with
species have different biogeographical patterns compared T. excelsa is almost unknown, and therefore we do not know
to those occurring in lowlands in tropical South America. whether this plant host share species of EcM fungi with Q.
The association of Andean Russulaceae with northern humboldtii or not, considering that they present different
temperate plant lineages suggested recent co-immigration origins (Tropical Amphi-pacific vs Holartic). Other little-
with Quercus when the latter colonised the rising Andes known patterns that affect the distribution of the species is
during the Pleistocene (Hackel et al., in press). For example, the arrival of invasive species, such as Amanita muscaria,
Russula floriformis subsp. floriformis and R. floriformis subsp. which has been detected on roots of Q. humboldtii (Vargas
symphoniae were recently described as new from montane et al., 2019).
forest dominated by Quercus and/or Oreomunnea (Fagales) In wet tropical lowland forests, Dipterocarpaceae and
from Colombia and Panama, respectively (Vera et al., 2021) Fabaceae represent two distantly related plant lineages
(Figure 4a). These two subspecies’ morphological and within the angiosperms that have separately evolved the
phylogenetic proximities supported their diversification due ability to form EcM symbioses (Wang & Qiu, 2006). As
to a co-immigration, adaptation and geographic isolation of mentioned before, these hosts occur in tropical rainforests in
Quercus and their symbionts along the Isthmus of Panama the Amazon region, supporting a surprising diversity of EcM
fungi, with a total of 114 morphospecies. However, based
A on the available information (descriptions or keys), only 61
of those morphospecies are correlated to species names,
with 20 corresponding to speciesthat are to science, only
some of which are in process of description (Vasco-Palacios
2016; Vasco-Palacios et al., 2014, 2018). Of the 61 species,
45 were associated with Pseudomonotes tropenbosii
(Dipterocarpaceae) and 42 species with Fabaceae (Table 1).
The discovery of the endemic tree P. tropenbosii emphasises
a phytogeographical link between the Colombian Amazon and
continental Africa and Madagascar (Morton et al., 1999). In
addition, the EcM status of P. tropenbosii indicates that the
EcM habit evolved before the continental separation, and
that radiations of EcM fungi associated with dipterocarps
across continents are explained by co-migrations of the
B fungus and host partners (Halling et al., 2008; Brearley
EcM fungi associated with
Trigonabalanus excelsa et al., 2012; Moyersoen, 2012). In addition, several EcM
(Fagaceae) fungal species that were previously restricted to the Guiana
Holartic lineages
EcM fungi associated with in three years of sampling, representing 72% of the total
Dipterocarpaceae species reported from Guyana (Vasco-Palacios & Boekhout,
2022). Similarities in the EcM fungi community structure
EcM fungi associated with between P. tropenbosii, Fabaceae forests, and Pakaraimaea
Fabaceaes (Cistaceae) suggested a low level of host specificity. Nearly
43% of the species associated with P. tropenbosii have also
been reported from Fabaceae forests in the Neotropics
FIGURE 5. A Proportions of distribution of EcM fungi species (Vasco-Palacios, 2016). This scenario is the case for the
with occurrence in Colombia, Neotropical. Blue tones indicate rare ascomycete, Pseudotulostoma volvatum, whose type
species distributed on the American continent, whereas red
specimen was collected associated with ectomycorrhizal
species distributed in two continents, and green are species
in more than two continental areas. B EcM fungi in neotropical
Dicymbe corymbosa (Fabaceae) trees and was also found
oak communities belong to Holarctic lineages. By contrast, associated with P. tropenbosii in Colombia. Species such as
Gondwanan lineages are represented in the EcM fungi of tropical Amanita xerocybe, Craterellus atratus, Clavulina amazonensis,
lowland forests in the Colombian Amazon, African Fabaceae and and Coltriciella oblectabilis are widely distributed in the
Asian Dipterocarpaceae. Amazonian region and the Guyana plateau being associated
FIGURE 6. Lichenised fungi analysed in this chapter. A Bunodophoron melanocarpum. B Neoprotoparmelia multifera. C Crocodia
aurata. D Lobariella sipmanii. E Sticta hirsuta. F Cora elephas. (Photographs by Robert Lücking.)
with multiple hosts (Singer et al., 1983; Henkel et al., 2002, widely distributed, including Gondwanan (American-African)
2012; Smith et al., 2013; Roy et al., 2016). These findings and circumpacific (American-Asian) disjunctions, species
show that, although EcM symbioses seem rare in lowland found across the Northern or the Western Hemisphere,
tropical forests and host plants are often distributed in and pantropical and (sub-)cosmopolitan taxa. The largest
isolated patches, lowland forest EcM fungi may be abundant proportion of a particular distribution type is provided
and may present broad distribution ranges. by Neotropical species (35%), whereas endemic species
Long-distance dispersal combined with low host presumably correspond only to 8.5% (Chapter 6).
specificity may increase the possibility of gene flow To assess this pattern, we looked at eight genera for
between geographically distant populations of these EcM which a large amount of molecular data are available:
fungi (Roy et al., 2016; Tedersoo et al., 2010; Moyersoen, Bunodophoron (Sphaerophoraceae), Neoprotoparmelia
2012; Vasco-Palacios et al., 2018). Other important hosts (Parmeliaceae), Crocodia, Lobariella, Podostictina,
of EcM fungi in tropical lowland forests are Nyctaginaceae Pseudocyphellaria, Sticta (Peltigeraceae, all Ascomycota), and
(e.g., Neea, Guapira) and Polygonaceae (e.g., Coccoloba), Cora (Hygrophoraceae, Basidiomycota). These lichens include
but little is known about these ecosystems and the role crustose (Neoprotoparmelia), fruticose (Bunodophoron), and
of those plant hosts in species distribution throughout foliose lichens (all other genera; Figure 6a–f). We used the
the vast territory of the Amazon region (Vasco-Palacios checklist published by Sipman et al. (2008) to assess the
et al., 2020). Phylogenetic studies in tropical lowland number of species corresponding to these eight genera
EcM lineages are consistent with a Gondwanan origin, in Colombia and their presumed distribution patterns
i.e., phylogeographic links between South America and before molecular studies. We then analysed published and
Africa (Figure 5b) (Moyersoen, 2012; Hosaka et al., 2008; unpublished molecular data to estimate the actual number
Matheny, 2009; Koch et al., 2019). Further studies will help of species in the corresponding trees and their distribution
us to identify distribution patterns of EcM fungi (dispersal ranges (Moncada et al., 2013, 2014; Lücking et al., 2017a,
vs vicariance), as well as host specificity and preferences. b; Soto-Medina et al., 2018; Santos et al., 2019).
Based on Sipman et al. (2008), the eight genera
BIOGEOGRAPHY OF COLOMBIAN LICHENS: TRADITIONAL correspond to 49 mostly phenotypically defined species
TAXONOMY VERSUS MOLECULAR DATA (Table 2), with primarily broad distribution ranges: about
Chapter 6 presents an analysis of the distribution patterns 20% (sub-)Cosmopolitan or in the Southern Hemisphere,
of Colombian lichens, primarily relying on a phenotype- 10% pantropical, 12% circumpacific or in the Neotropics
based species concept but including published results from and Hawaii, and 2% Gondwanan (Figure 7a). By contrast,
molecular data available for some groups. According to that 39% were inferred as Neotropical and 12% as endemic,
analysis, more than half of the species (53%) are presumably somewhat higher than the overall proportions for all
TABLE 2. Species corresponding to the lichenised fungal genera Bunodophoron, Cora, Crocodia, Lobariella, Neoprotoparmelia, Podostictina,
Pseudocyphellaria, and Sticta in the checklist by Sipman et al. (2008).
TABLE 2. (continued)
TABLE 2. (continued)
lichen fungi (see Chapter 6). However, when analysing the Neotropical species will likely shift in favour of the latter.
same genera based on molecular data, the total number However, the global data for these groups suggest that
of species was estimated at 283, almost six times the these species are not intercontinentally widespread.
previous number, demonstrating the high degree of The observed difference between traditional phenotypic
hidden diversity in these taxa. The bulk of these additional and combined molecular-phenotypic species concepts in
species is found in the genera Cora and Sticta, where many these groups is substantial, showing an entirely different
of the phylogenetically defined clades still await formal picture of biogeographic patterns of Colombian lichens
description (Lücking et al., 2014, 2017a). When inferring the (Figure 7a, b). Nevertheless, it is not possible to extrapolate
distribution of these lineages, the overwhelming majority these results to all lichenised lineages in Colombia. The
(95%) was reconstructed as either endemic or Neotropical hyperdiversity of Cora in particular is distorting the picture
(Figure 7b). The exact proportion of truly endemic versus due to the high degree of previously unrecognised, hidden
more widespread Neotropical species is difficult to assess diversity in this genus, from just one to currently 78 species
due to the lack of dedicated inventories in many other recognised in Colombia. Even so, this exercise suggests
Neotropical areas. However, it is to be expected that that biogeographical distribution patterns derived from
many of these species also occur in other South American traditional, phenotype-based species concepts are not
countries, and some may extend to Central America and/ reliable.
or the Caribbean, so the current 75% endemic versus 20%
CONCLUSIONS
A B
FIGURE 7. Proportions of distribution types among species of eight selected lichen-forming fungal genera in Colombia.
Our assessment of three selected groups of fungi in Carranza J, Ruiz-Boyer A (2005) Checklist of polypores of Costa Rica.
Colombia shows that various factors contribute to Revista Mexicana de Micología, 20, 45-52.
Corrales A, Henkel TW, Smith ME (2018) Ectomycorrhizal associations in
substantial gaps and bias in assessing fungal biogeography the tropics—biogeography, diversity patterns and ecosystem roles.
in species-rich tropical countries. One of these factors is New Phytologist 220:1076–1091.
the often poor knowledge on the ecology and distribution Condit R, Ashton PS, Baker P, Bunyavejchewin S, Gunatilleke S,
of species and their accurate delimitation. If these factors Gunatilleke N, Hubbell SP, Foster RB, Itoh A, LaFrankie JV, Lee HS,
Losos E, Manokaran N, Sukumar R, Yamakura T (2000) Spatial
are not considered, biogeographical assessments can be patterns in the distribution of tropical tree species. Science, 288,
significantly misleading, both for individual species and 1414-1418. doi: 10.1126/science.288.5470.1414
for fungi as a whole. This has implications for evaluating David H, Cardona-Naranjo F, Calderón Árias AM, Martínez Figueroa Y,
their status as potential endemics and their conservation Murillo J (2019) HUA Herbario Universidad de Antioquia: 50 años
documentando algas, hongos y plantas de Colombia. Medellin:
assessment. Precise knowledge of species is one Editorial Universidad de Antioquia. 96p.
requirement to address this problem, and cataloguing all de Lima VX, Nogueira-Melo GS, Baltazar JM, de Medeiros PS, Soares AM,
species on the planet should be the main goal of biodiversity de Lira CRS, Gibertoni TB (2018) Fungal perspective on neotropical
research. However, discovering and naming species is just biogeography: poroid fungi (Agaricomycetes: Hymenochaetales and
Polyporales) and the Brazilian moist forests. Fungal Ecology, 36, 1-7.
part of the challenge. Knowing their ecology and geographic doi: 10.1016/j.funeco.2018.06.004
distribution is equally important if we intend to fully De Wit R, Bouvier T (2006) Everything is everywhere, but the environment
understand and preserve Earth’s biota. Using polypores, selects; what did Baas Becking and Beijerinck really say?
ectomycorrhizal fungi, and lichenised fungi as examples, Environmental Microbiology, 8(4), 755-758. doi: 10.1111/j.1462-
2920.2006.01017.x
we have demonstrated that species in these groups are Elias SG, Salvador-Montoya CA, Costa-Rezende DH, Guterres DC,
generally poorly known, and their biogeographical patterns Fernandes M, Olkoski D, Drechsler-Santos ER (2020) Studies on
are hard to assess. the biogeography of Phellinotus piptadeniae (Hymenochaetales,
Therefore, fungal distribution ranges are often Basidiomycota): expanding the knowledge on its distribution and
clarifying hosts relationships. Fungal Ecology, 45, 100912. doi:
misrepresented by available records within and outside 10.1016/j.funeco.2020.100912
Colombia. Targeted field explorations, particularly in Feuerer T, Hawksworth DL (2007) Biodiversity of lichens, including a
remote habitats, and phylogeographic studies are needed world-wide analysis of checklist data based on Takhtajan’s floristic
to provide reliable data to correct these issues. This kind regions. Biodiversity and Conservation, 16(1), 85-98. doi: 10.1007/
s10531-006-9142-6
of study should complement important but undervalued, Galloway DJ (2008) Lichen biogeography. In: Nash III TH (ed). Lichen
continuously updated species lists and distribution reports. Biology. Cambridge: Cambridge University Press. Pp.315-335. doi:
The maintenance and improvement of collection databases, 10.1017/CBO9780511790478
such as ColFungi (https://colfungi.org), GBIF (http://www. Gaya E, Vasco-Palacios AM, Vargas-Estupiñan N, Lücking R, Carretero
J, Sanjuan T, Moncada B, Allkin B, Bolaños-Rojas AC, Castellanos-
gbif.org), and SIB (https://sibcolombia.net) will contribute Castro C, Coca LF, Corrales A, Cossu T, Davis L, dSouza J, Dufat A,
to the broad distribution of knowledge about the Colombian Franco-Molano AE, Garcia F, Gómez-Montoya N, González-Cuellar FE,
funga. All of these efforts require the continued work of Hammond D, Herrera A, Jaramillo-Ciro MM, Lasso-Benavides C, Mira
trained taxonomists, who continue to be essential assets MP, Morley J, Motato-Vásquez V, Niño-Fernandez Y, Ortiz-Moreno ML,
Peña-Cañón ER, Ramirez-Castrillón M, Rojas T, Ruff J, Simijaca D,
in the mission to document and understand Colombia’s Sipman HJM, Soto-Medina E, Torres G, Torres-Andrade PA, Ulian T,
and the world’s biodiversity. White K, Diazgranados M (2021) ColFungi: Colombian Resources for
Fungi Made Accessible. Richmond: Royal Botanic Gardens, Kew. 36p.
Gilbert GS, Ferrer A, Carranza J (2002) Polypore fungal diversity and host
Acknowledgements density in a moist tropical forest. Biodiversity and Conservation, 11,
947-957. doi: 10.1023/A:1015896204113
We would like to thank the reviewer Manuela dal Forno for her Hackel J, Henkel T, Moreau PA, De Crop E, Verbeken A, Buyck B, Neves
insightful suggestions. MA, Vasco-Palacios AM (submitted) Distinct biogeographic origins of
neotropical ectomycorrhizal fungi. New Phytologist.
Halling RE (1996) Boletaceae (Agaricales): Latitudinal biodiversity and
biological interactions in Costa Rica and Colombia. Boletaceae
References (Agaricales). Revista de Biología Tropical, 44(4), 111-114.
Aguilar M, Fiore-Donno AM, Lado C, Cavalier-Smith T (2014) Using Halling RE, Osmundson TW, Neves MA (2008) Pacific boletes: implications
environmental niche models to test the ‘everything is everywhere’ for biogeographic relationships. Mycological Research, 112(4), 437-
hypothesis for Badhamia. The ISME Journal, 8(4), 737-745. doi: 447. doi: 10.1016/j.mycres.2007.11.021
10.1038/ismej.2013.183 Hattori T (2017) Biogeography of polypores in Malesia, Southeast Asia.
Allen JL, Lendemer JC (2015) Fungal conservation in the USA. Endangered Mycoscience, 58(11), 1-13. doi: 10.1016/j.myc.2016.09.004
Species Research, 28(1), 33-42. doi: 10.3354/esr00678 Henkel TW, Aime MC, Chin MML, Miller SL, Vilgalys R, Smith ME
Baas Becking LGM (1934) Geobiologie of inleiding tot de milieukunde. The (2012) Ectomycorrhizal fungal sporocarp diversity and discovery of
Netherlands: WP Van Stockum & Zoon NV. 263p. new taxa in Dicymbe monodominant forests of the Guiana Shield.
Baltazar JM, Gibertoni TB (2009) A checklist of the aphyllophoroid Biodiversity and Conservation, 21, 2195-2220. doi: 10.1007/
fungi (Basidiomycota) recorded from the Brazilian Atlantic Forest. s10531-011-0166-1
Mycotaxon, 109, 439-442. doi: 10.5248/109.439 Henkel TW, Terborgh J, Vilgalys RJ (2002) Ectomycorrhizal fungi and their
Brearley FQ (2012) Ectomycorrhizal associations of the leguminous hosts in the Pakaraima Mountains of Guyana. Mycological
Dipterocarpaceae. Biotropica, 44, 637-648. doi: 10.1111/j.1744- Research, 106, 515-531. doi: 10.1017/S0953756202005919
7429.2012.00862.x Hooghiemstra H (2006) Immigration of oak into Northern South America:
Brundrett MC, Tedersoo L (2018) Evolutionary history of mycorrhizal a paleo-ecological document. In: Kappelle M (ed). Ecology and
symbioses and global host plant diversity. New Phytologist, 220(4), conservation of neotropical montane oak forests. Berlin: Springer.
1108-1115. doi: 10.1111/nph.14976 Pp. 17-28.
Hortal J, de Bello F, Diniz-Filho JAF, Lewinsohn TM, Lobo JM, Ladle Moncada B, Lücking R, Betancourt-Macuase L (2013) Phylogeny of
RJ (2015) Seven shortfalls that beset large-scale knowledge of the Lobariaceae (lichenised Ascomycota: Peltigerales), with a
biodiversity. Annual Review of Ecology, Evolution and Systematics, reappraisal of the genus Lobariella. The Lichenologist, 45(2), 203-
46(1), 523-549. doi: 10.1146/annurev-ecolsys-112414-054400 263.
Hosaka K, Castellano MA, Spatafora JW (2008) Biogeography of Moncada B, Lücking R, Suárez A (2014) Molecular phylogeny of the
Hysterangiales (Phallomycetidae, Basidiomycota). Mycological genus Sticta (lichenised Ascomycota: Lobariaceae) in Colombia.
Research, 112(4), 448-462. doi: 10.1016/j.mycres.2007.06.004 Fungal Diversity, 64(1), 205-231. doi: 10.1007/s13225-013-0230-0
James TY, Moncalvo JM, Li S, Vilgalys R (2001) Polymorphism at the Moncada B, Lücking R, Lumbsch HT (2020) Rewriting the evolutionary
ribosomal DNA spacers and its relation to breeding structure of the history of the lichen genus Sticta (Ascomycota: Peltigeraceae
widespread mushroom Schizophyllum commune. Genetics, 157(1), subfam. Lobarioideae) in the Hawaiian Islands. Plant and Fungal
149-161. Systematics, 65(1), 95-119. doi: 10.35535/pfsyst-2020-0005
Kappelle M (2006) Structure and composition of Costa Rican montane Morton CM, Dayanandan S, Dissanayake D (1999). Phylogeny and
oak forests. In: Kappelle M (ed). Ecology and conservation of biosystematics of Pseudomonotes (Dipterocarpaceae) based on
neotropical montane oak forests. Berlin: Springer. Pp. 127-139. molecular and morphological data. Plant Systematics and Evolution,
Kirk PM, Cannon PF, David JC, Stalper JA (2008) Dictionary of the Fungi, 216, 197-205. doi: 10.1007/BF01084398
11th ed. Wallingford: CABI Publishing. 784p. Moser M, Horak E (1975) Cortinarius und nahe verwandte Gattungen in
Koch R, Dentinger B, Henkel T, Aime C (2019) Phylogenomics of Sudamerica. Beihefte zur Nova Hedwigia, 52, 1-628.
ectomycorrhizal genera suggest that some may have a common, Motato-Vásquez V, Grassi E, Gugliotta AM, Robledo G (2018) Evolutionary
ancient origin in northern Gondwana. Systematic and Biodiversity relationship of Bresadolia (Basidiomycota, Polyporales) based on
Session. Mycological Society of America annual meeting, molecular and morphological evidence. Mycological Progress, 17(9),
Minneapolis. 1031-1048. doi: 10.1007/s11557-018-1416-3
Krah FS, Bässler C, Heibl C, Soghigian J, Schaefer H, Hibbett DS (2018) Moyersoen B (2012) Dispersion, an important radiation mechanism for
Evolutionary dynamics of host specialisation in wood-decay fungi. ectomycorrhizal fungi in Neotropical lowland forests? In: Sudarshana
BMC Evolutionary Biology, doi: 10.1186/s12862-018-1229-7 P, Nageswara-Rao M, Soneji JR (eds). Tropical Forests. Rijeka: InTech.
Leigh EG, O’Dea A, Vermeij GJ (2014) Historical biogeography of the Pp. 93-116.
Isthmus of Panama. Biological Reviews, 89(1), 148-172. doi: Mueller GM, Halling RE, Carranza J, Mata M, Schmit JP (2006)
10.1111/brv.12048 Saprotrophic and ectomycorrhizal macrofungi of Costa Rica oak
Lindblad I (2000) Host specificity of some wood-inhabiting forest. In: Kapelle M (ed). Ecology and Conservation of Neotropical
fungi in a tropical forest. Mycologia, 92, 399-405. doi: Montane Oak Forest. Berlin: Springer. Pp. 55-68.
10.1080/00275514.2000.12061174 Mutke J, Jacobs R, Meyers K, Henning T, Weigend M (2014) Diversity
Lindblad I (2001) Diversity of poroid and some corticoid wood- patterns of selected Andean plant groups correspond to topography
inhabiting fungi along the rainfall gradient in tropical forests, and habitat dynamics, not orogeny. Frontiers in Genetics, 5, 1-15.
Costa Rica. Journal of Tropical Ecology, 17, 353-369. doi: 10.1017/ doi: 10.3389/fgene.2014.00351
S0266467401001249 O’Malley MA (2007) The nineteenth-century roots of ‘everything is
Lomolino MV, Riddle BR, Whittaker RJ (2017) Biogeography, 5th edition. everywhere’. Nature Reviews Microbiology, 5(8), 647-651. doi:
Sunderland: Sinnauer Associates, Inc. 850p. 10.1038/nrmicro1711
Lücking R (2003) Takhtajan’s floristic regions and foliicolous lichen O’Dea A, Lessios HA, Coates AG, Eytan RI, Restrepo-Moreno SA, Cione
biogeography: a compatibility analysis. The Lichenologist, 35(1), AL, Jackson JB (2016) Formation of the Isthmus of Panama. Science
33-53. doi: 10.1006/lich.2002.0430 Advances, 2(8), e1600883. doi: 10.1126/sciadv.1600883
Lücking R, Dal-Forno M, Sikaroodi M, Gillevet PM, Bungartz F, Moncada Olson DM, Dinerstein E, Wikramanayake ED, Burgess ND, Powell GV,
B, Yánez-Ayabaca A, Chaves JL, Coca LF, Lawrey JD (2014) A Underwood EC, D’amico JA, Itoua I, Strand HE, Morrison JC, Loucks
single macrolichen constitutes hundreds of unrecognised species. CJ (2001) Terrestrial ecoregions of the world: a new map of Life
Proceedings of the National Academy of Sciences U S A, 111(30), on Earth, a new global map of terrestrial ecoregions provides an
11091-11096. doi: 10.1073/pnas.1403517111 innovative tool for conserving biodiversity. BioScience, 51(11), 933-
Lücking R, Dal Forno M, Moncada B, Coca LF, Vargas-Mendoza LY, 938. doi: 10.1641/0006-3568(2001)051[0933:TEOTWA]2.0.CO;2
Aptroot A, Arias LJ, Besal B, Bungartz F, Cabrera-Amaya DM, Palacio M, Robledo GL, Reck MA, Grassi E, Góes-Neto A, Drechsler-
Cáceres ME (2017a) Turbo-taxonomy to assemble a megadiverse Santos ER (2017) Decrypting the Polyporus dictyopus complex:
lichen genus: seventy new species of Cora (Basidiomycota: recovery of Atroporus Ryvarden and segregation of Neodictyopus gen.
Agaricales: Hygrophoraceae), honouring David Leslie Hawksworth’s nov. (Polyporales, Basidiomycota). PLoS ONE, 12 (10), e0186183.
seventieth birthday. Fungal Diversity, 84(1), 139-207. doi: 10.1007/ doi: 10.1371/journal.pone.0186183
s13225-016-0374-9 Peay KG, Bidartondo MI, Arnold A (2010) Not every fungus is everywhere:
Lücking R, Moncada B, McCune B, Farkas E, Goffinet B, Parker D, scaling to the biogeography of fungal–plant interactions across
Chaves JL, Lőkös L, Nelson PR, Spribille T, Stenroos S, Wheeler roots, shoots and ecosystems. New Phytologist, 185(4), 878-882.
T, Yanez-Ayabaca A, Dillman K, Gockman OT, Goward T, Hollinger J, doi: 10.1111/j.1469-8137.2009.03158.x
Tripp EA, Villella J, Álvaro-Alba WR, Arango CJ, Cáceres MES, Coca Peña-Venegas CP, Vasco-Palacios AM (2019) Endo- and ectomycorrhizas
LF, Printzen C, Rodríguez C, Scharnagl K, Rozzi R, Soto-Medina E, in tropical ecosystems of Colombia. In: Pagano M, Lugo M (eds).
Yakovchenko LS (2017b) Pseudocyphellaria crocata (Ascomycota: Mycorrhizal Fungi in South America. Berlin: Springer. Pp. 111-146.
Lobariaceae) in the Americas is revealed to be thirteen species, Ramírez-Camejo LA, Zuluaga-Montero A, Lázaro-Escudero M, Hernández-
and none of them is P. crocata. The Bryologist, 120(4), 441-500. Kendall V, Bayman P (2012) Phylogeography of the cosmopolitan
doi: 10.1639/0007-2745-120.4.14 fungus Aspergillus flavus: is everything everywhere? Fungal Biology,
Lumbsch HT, Buchanan PK, May TW, Mueller GM (2008) Phylogeography 116(3), 452-463. doi: 10.1016/j.funbio.2012.01.006
and biogeography of fungi. Mycological Research, 112(4), 423-424. Rangel Ch JO, Avella A (2011) Oak forests of Quercus humboldtii
doi: 10.1016/j.mycres.2008.02.002 in the Caribbean region and distribution patterns related with
Magain N, Sérusiaux E (2015) Dismantling the treasured flagship lichen environmental factors in Colombia. Plant Biosystems-, 145(sup1),
Sticta fuliginosa (Peltigerales) into four species in Western Europe. 186-198. doi: 10.1080/11263504.2011.602727
Mycological Progress, 14(10), 1-33. doi: 10.1007/s11557-015- Robledo GL, Rajchenberg M (2007) Preliminary polypore mycota
1109-0 (Basidiomycetes) from northwestern Argentinean Yungas. Mycotaxon,
Matheny PB, Aime MC, Bougher NL, Buyck B, Desjardin DE, Horak E, 100, 5-9.
Hibbett DS (2009) Out of the Palaeotropics? Historical biogeography Roy M, Schimann H, Braga‐Neto R, Da Silva RA, Duque J, Frame D, Neves
and diversification of the cosmopolitan ectomycorrhizal mushroom MA (2016) Diversity and distribution of ectomycorrhizal fungi from
family Inocybaceae. Journal of Biogeography, 36(4), 577-592. doi: Amazonian lowland white‐sand forests in Brazil and French Guiana.
10.1111/j.1365-2699.2008.02055.x Biotropica, 48(1), 90-100. doi: 10.1111/btp.12297
Salvador-Montoya CA, Robledo GL, Cardoso D, Borba-Silva MA, van der Hammen T, Cleef AM (1983) Trigonobalanus and the tropical
Fernandes M, Dreschler-Santos ER (2015) Phellinus piptadeniae amphi-pacific element in the North Andean Forest. Journal of
(Hymenochaetales: Hymenochaetaceae): taxonomy and host range Biogeography, 437-440. doi: 10.2307/2844750
of a species with disjunct distribution in South American seasonally Vargas N, Restrepo S (2020) A checklist of ectomycorrhizal mushrooms
dry forest. Plant Systematics and Evolution, 301, 1887-1896. doi: associated with Quercus humboldtii in Colombia. In: Pérez-Moreno J,
10.1007/s00606-015-1201-6 Guerin-Laguette A, Flores R, Yu AQ (eds). Mushrooms, humans and
Santos LA, Aptroot A, Lücking R, da Silva Cáceres ME (2019) nature in a changing world. Berlin: Springer. Pp. 425-450.
High diversification in the Neoprotoparmelia multifera complex Vargas N, Gonçalves SC, Franco-Molano AE, Restrepo S, Pringle A (2019)
(Ascomycota, Parmeliaceae) in northeast Brazil revealed by DNA In Colombia the Eurasian fungus Amanita muscaria is expanding its
barcoding and phenotypical characters. The Bryologist, 122(4), 539- range into native, tropical Quercus humboldtii forests. Mycologia,
552. doi: 10.1639/0007-2745-122.4.539 111(5), 758-771. doi: 10.1080/00275514.2019.1636608
Sanuma OI, Tokimoto K, Sanuma C, Autuori J, Sanuma LR, Sanuma M, Vasco Palacios AM (2016) Ectomycorrhizal fungi in Amazonian tropical
Martins MS, Menolli Jr N, Ishikawa NK, Apiamö RM (2016) Sanöma forests in Colombia. Doctoral dissertation. Utrecht: Utrecht
samakönö sama tökö nii pewö oa wi ĩ tökö waheta: Ana amopö = University. 203p.
Enciclopédia dos alimentos yanomami Sanöma: Cogumelos. São Vasco-Palacios AM, Franco-Molano AE (2013) Diversity of Colombian
Paulo: Instituto Socioambiental. 108p. macrofungi (Ascomycota-Basiciomycota). Mycotaxon, 121, 48.
Schoch CL, Seifert KA, Huhndorf S, Robert V, Spouge JL, Levesque CA, Vasco-Palacios AM, López-Quintero C, Franco-Molano AE, Boekhout
Fungal Barcoding Consortium (2012) Nuclear ribosomal internal T (2014). Austroboletus amazonicus sp. nov. and Fistulinella
transcribed spacer (ITS) region as a universal DNA barcode marker campinaranae var. scrobiculata, two commonly occurring boletes from
for Fungi. Proceedings of the National Academy of Sciences U S A, a forest dominated by Pseudomonotes tropenbosii (Dipterocarpaceae)
109(16), 6241-6246. doi: 10.1073/pnas.1117018109 in Colombian Amazonia. Mycologia, 106(5), 1004-1014.
Silveira RMB, Wright JE (2005) The taxonomy of Echinochaete and Vasco-Palacios AM, Hernandez J, Peñuela MC, Boekhout T (2018)
Polyporus s. str. in South America. Mycotaxon, 93,1-59. Ectomycorrhizal fungi diversity in a white sand forest in
Singer R, Araujo I, Ivory MH (1983) The ectotrophically mycorrhizal fungi western Amazonia. Fungal Ecology, 31, 9-18. doi: 10.1016/j.
of the neotropical lowlands, especially central Amazonia. Vaduz: funeco.2017.10.003
Lubrecht & Cramer. 352p. Vasco-Palacios AM, Bahram M, Boekhout T, Tedersoo L (2020) Carbon
Sipman HJM, Hekking W, Aguirre-C J (2008) Checklist of Lichenized and content and pH as important drivers of fungal community structure in
Lichenicolous Fungi from Colombia. Bogotá DC: Instituto de Ciencias three Amazon forests. Plant and Soil, 450(1), 111-131. doi: 10.1007/
Naturales, Universidad Nacional de Colombia. 242p. s11104-019-04218-3
Smith ME, Henkel TW, Aime MC, Fremier AK, Vilgalys R (2011) Vasco-Palacios AM, Boekhout T (2022) Pseudomonotes tropenbosii, an
Ectomycorrhizal fungal diversity and community structure on three endemic dipterocarp tree from a neotropical terra-firme forest in
co‐occurring leguminous canopy tree species in a Neotropical Colombian Amazonia that hosts ectomycorrhizal fungi. In: Lugo A,
rainforest. New Phytologist, 192, 699-712. doi: 10.1111/j.1469- Pagano M, editors. Mycorrhizal fungi in South America, biodiversity,
8137.2011.03844.x conservation, and sustainable food production. Springer.
Smith ME, Henkel TW, Uehling JK, Fremier AK, Clarke HD, Vilgalys R Vera M, Slavomír A, Adamčíková K, Hampe F, Caboň M, Manz C,
(2013) The ectomycorrhizal fungal community in a Neotropical Forest Ovrebo C, Piepenbring M, Corrales A (2021) Morphological and
dominated by the endemic Dipterocarp Pakaraimaea dipterocarpacea. genetic diversification of Russula floriformis, sp. nov., along
PLoS One, 8, e55160. doi: 10.1371/journal.pone.0055160 the Isthmus of Panama. Mycologia, 113(4), 807-827. doi:
Sobral M, Stehmann JR (2009) An analysis of new angiosperm species 10.1080/00275514.2021.18973 77
discoveries in Brazil (1990–2006). Taxon, 58, 227-232. doi: Wang B, Qiu YL (2006) Phylogenetic distribution and evolution of
10.1002/tax.581021 mycorrhizas in land plants. Mycorrhiza, 16(5), 299-363. doi: 10.1007/
Song J, Cui BK (2017) Phylogeny, divergence time and historical s00572-005-0033-6
biogeography of Laetiporus (Basidiomycota, Polyporales). BMC Watkinson SC, Bebber D, Darrah PR, Fricker M, Tlalka M, Boddy L (2006)
Evolutionary Biology, 17(1), 1-12. doi: 10.1186/s12862-017-0948-5 The role of wood-decaying in the carbon and nitrogen dynamics of
Soto-Medina ES, Prieto M, Wedin M (2018) A new Bunodophoron the forest floor. In: Gadd GM (ed). Fungi in Biogeochemical Cycles.
species (Sphaerophoraceae, Lecanorales) from the Neotropics. The Cambridge: Cambridge University Press. Pp 151-181.
Lichenologist, 50(3), 255-266. doi: 10.1017/S0024282917000743 Werth S (2011) Biogeography and phylogeography of lichen fungi and
Summerell BA, Laurence MH, Liew EC, Leslie JF (2010) Biogeography and their photobionts. In: Fontaneto D (ed). Biogeography of microscopic
phylogeography of Fusarium: a review. Fungal Diversity, 44(1), 3-13. organisms: is everything small everywhere. Cambridge: Cambridge
doi: 10.1007/s13225-010-0060-2 University Press. Pp. 191-208. doi: 10.1017/CBO9780511974878
Takhtajan A, Crovello TJ, Cronquist A (1986) Floristic regions of the world. Wicklow DT (1981) Biogeography and conidial fungi. In: Biology of conidial
Los Angeles: University of California Press. 522p. fungi, Vol. 1. Cambridge: Academic Press. Pp. 417-447. doi: 10.1016/
Tedersoo L, Suvi T, Beaver K, Saar I (2007) Ectomycorrhizas of C2012-0-01472-X
Coltricia and Coltriciella (Hymenochaetales, Basidiomycota) on Yang H, Dai Y, Xu M, Zhang Q, Bian X, Tang J, Chen X (2016) Metadata-
Caesalpiniaceae, Dipterocarpaceae and Myrtaceae in Seychelles. mining of 18S rDNA sequences reveals that “everything is not
Mycological Progress, 6(2), 101-107. doi: 10.1007/s11557-007- everywhere” for glomeromycotan fungi. Annals of Microbiology, 66(1),
0530-4 361-371. doi: 10.1007/s13213-015-1116-z
Tedersoo L, Sadam A, Zambrano M, Valencia R, Bahram M (2010) Low
diversity and high host preference of ectomycorrhizal fungi in Western
Amazonia, a neotropical biodiversity hotspot. The ISME Journal, 4(4),
465-471. doi: 10.1038/ismej.2009.131
Tedersoo L, Bahram M, Põlme S, Kõljalg U, Yorou NS, Wijesundera R,
Vasco-Palacios A, Abarenkov K (2014) Global diversity and geography
of soil fungi. Science, 346, 6213. doi: 10.1126/science.1256688
Uehling JK, Henkel TW, Aime MC, Vilgalys R, Smith ME (2012) New species
and distribution records for Clavulina (Cantharellales, Basidiomycota)
from the Guiana Shield, with a key to the lowland Neotropical taxa.
Fungal Biology, 116, 1263-1274. doi: 10.1016/j.funbio.2012.09.004
Universidad de Antioquia (2020) Herbario Universidad de Antioquia
(HUA), Colección de Hongos en línea.
Väisänen R, Heliövaara K, Kotiranta H, Niemelä T (1992) Biogeographical Amanita xerocybe
analyses of Finnish polypore assemblages. Karstenia, 32, 17–28. [Aida Marcela Vasco-Palacios]
Scleroderma flavidum
[Robert Lücking]
138 CATALOGUE OF FUNGI OF COLOMBIA
Chapter 11
Diversity, Functional Groups, and Community Structure of Fungi of Colombia
Adriana Corrales1*, Cristina Benjumea2 & Nataly Gomez-Montoya2
1
Centro de Investigaciones en Microbiología y Biotecnología-UR (CIMBIUR), Facultad de Ciencias Naturales, Universidad del Rosario, Bogotá DC, Colombia.
2
Laboratorio de Taxonomía y Ecología de Hongos -TEHO, Instituto de Biología, Facultad de Ciencias Exactas y Naturales, Universidad de Antioquia - UdeA,
Calle 70 No. 52-21, Medellín, Antioquia, Colombia.
ABSTRACT
Owing to its tropical location and great diversity of ecosystems, Colombia hosts a high diversity of fungi, which play
fundamental roles in ecosystems as mycorrhizas, saprophytes, endophytes, and pathogens. In this chapter, we describe
a literature review of ecological studies focusing on the most - studied fungal functional groups of Colombia: arbuscular
mycorrhizas, ectomycorrhizas, saprotrophs, and endophytes. To complement the literature review, we used the University of
Antioquia Herbarium (HUA) database, which includes more than 11,000 macrofungi specimens, to run alpha diversity and
network analyses. The alpha diversity analyses were performed using species accumulation curves for the total number of
specimens and specimens divided by ecosystem types (oak forests, mixed montane forests, Amazonian lowland forests,
and extra-Amazonian lowland forests) and with diversity indexes based on Chao 1 and bootstrap estimators. Network
analysis was based on correlation matrices among main ecosystem types. These analyses made it possible to compare the
species richness and diversity patterns among the main ecosystem types. Our results show a high number of collections of
ectomycorrhizal fungi from oak forests, which are typical of this type of ecosystem, and a high alpha diversity for Colombia’s
Amazonian lowland forests. We highlight significant knowledge gaps regarding ecological research in the country as the
diversity patterns of many functional groups of fungi are still unknown. We recommend increasing the number of studies
based on environmental sequencing techniques because these will allow the inclusion species of microfungi.
RESUMEN
Colombia por su ubicación en la región tropical y su gran diversidad de ecosistemas, favorece una alta diversidad en
hongos, los cuales son fundamentales en los ecosistemas dado que juegan papeles importantes como hongos micorrízicos,
saprófitos, endófitos y patógenos. Para el presente trabajo se realizó una revisión de literatura de estudios ecológicos
enfocados en los grupos funcionales de hongos más estudiados en el país. Para complementar la información obtenida en
la revisión de literatura, se utilizó la base de datos del herbario de la Universidad de Antioquia (HUA), la cual incluye más de
11.000 colecciones de macrohongos, para realizar análisis de diversidad alfa y análisis de redes. Los análisis de diversidad
alfa se realizaron con curvas de acumulación de especies para el total de las colecciones y para las colecciones divididas
por tipos de ecosistemas (bosque de roble, bosque montano mixto, tierras bajas amazónicas y otras tierras bajas) y con
estimadores de diversidad basados en Chao 1 y análisis de bootstrap. Los análisis de redes se realizaron con base en
matrices de correlación entre los principales ecosistemas. Con estos análisis fue posible comparar la riqueza de especies
entre los principales tipos de ecosistemas, y a su vez se pudo identificar los principales patrones de diversidad beta para el
país. Los resultados muestran una alta diversidad alfa para los bosques amazónicos de tierras bajas del país, ya que con
menor número de colecciones se obtuvo un número total de 524 especies, con un número estimado de especies entre 638
y 975. Además, se observó un elevado número de especies (474) en los bosques de roble en su mayoría representados por
especies ectomicorrízicas, lo que lo posiciona en el segundo ecosistema más rico en especies de hongos del país con una
riqueza estimada de especies entre 566 y 847. Los análisis de redes mostraron un bajo número de especies compartidas
entre los bosques de roble y los demás ecosistemas; mientras que los bosques montanos mixtos y los bosques de tierras
bajas comparten un número alto de especies. Nuestros resultados muestran la existencia de grandes vacíos de información
en cuanto a investigación ecológica en el país, ya que aún se desconocen los patrones de diversidad de muchos grupos
funcionales de hongos. En especial, se recomienda realizar estudios basados en técnicas de secuenciación de muestras
ambientales, ya que estos permitirán incluir especies de microhongos los cuales no pueden ser estudiados con el uso de
colecciones de herbario.
spore number counts (e.g., Restrepo et al., 2019; Sandoval analysis. In addition, molecular data showed a higher
& Ordoñez, 2019), evaluation of mycorrhizal colonisation number of Archaeosporaceae and Gigasporaceae and
on roots (e.g., Restrepo et al., 2019), and identification of other rare species that are usually undetectable when
morphotypes based on spore morphology (e.g., Posada et using spore morphology due to their infrequent sporulation
al., 2018). However, during the past five years, there has (Peña-Venegas et al., 2021). In another study, Peña-
been an increase in ecological studies using metagenomic Venegas et al. (2020) also compared the diversity of AMF
approaches (Peña-Venegas et al., 2021). Questions regarding using spore morphology and molecular methods. They
the influence of biotic and abiotic factors on AMF spore found similar species richness with both methods (88 with
number, species richness, and species composition have spore morphology and 85 with molecular methods), with
been addressed in anthropogenic and natural ecosystems Glomus being the dominant genus. Several studies from
throughout the Colombian territory (e.g., López, 2009; other sites of Colombia at low and high altitudes also found
Tirado-Ardila, 2017). Glomus to be the most dominant genus (Restrepo et al.,
In general, it has been shown that soil variables have 2019; Peña-Venegas et al., 2021).
a substantial effect on all aspects of AMF ecology. For Regarding changes of AMF communities along
example, Restrepo et al. (2019), studying AMF from cattle altitudinal gradients, there have been two studies. Peña-
ranching systems in the department of Antioquia, found Venegas et al. (2020) studied AMF in an altitudinal gradient
that in lowland ecosystems of the Magdalena Medio, spore between 500m and 1,850 m altitude. They found that the
abundance was negatively correlated with Ca, Mg, K, P, and abundance of AMF spores had a positive correlation with
pH, while in the montane systems of northern Antioquia AMF higher altitudes. This positive correlation with altitude could
colonisation was positively correlated with K. Also, Sandoval be associated with increased phosphorus limitation with
& Ordoñez (2019) found a negative correlation between altitude (Peña-Venegas et al., 2020). Species composition
pH and AMF spore abundance in more acid soils, showing also changed with elevation, with each site showing a
higher spore counts in secondary montane and bamboo characteristic community with few species distributed
dominated forests. By contrast, in coffee plantations in among altitudes. Regardless of this increase in spore
the same sites, AMF spore abundance was negatively abundance and changes in composition, species richness
correlated with Na, K, Mg, Ca, and soil compaction. In a was constant along the altitudinal gradient Peña-Venegas et
similar montane ecosystem, Peña & Botía (2018), studying al., 2020). Studies performed in the Andes mountain range
forests in different successional stages, found that AMF at higher altitudes showed a negative relationship between
soil spore abundance was positively correlated with soil the number of AMF spores in the soil and P content at
organic matter content, temperature, and K, but negatively increasing altitudes (López, 2009; Tirado-Ardila, 2017).
correlated with Fe, P, Ca, Zn, and soil compaction. The degree of anthropic intervention or successional
Regarding AMF species richness and composition, states has been shown to alter the species richness of
Peña-Venegas et al. (2021), using molecular approaches AMF in Colombia. A study by Sandoval & Ordoñez (2019),
in lowland rubber plantations, found that AMF species looking at sites with different levels of anthropogenic
richness was negatively correlated with pH, Ca, Mg, and disturbance (coffee plantation, bamboo - dominated forest,
P. These authors also found that AMF species composition and secondary forest) in the Cundinamarca department,
changed along a Na - availability gradient, with soil texture found that the number of AMF spores in the soil was
being also an important factor structuring the communities. negatively correlated with the degree of disturbance. Soil
Using morphotypes extracted from soil and bait cultures from secondary and bamboo dominated forests showed the
from coffee plantations in the Risaralda department, highest number of AMF spores, while the coffee plantation
Posada et al. (2018) found that AMF species richness had the lowest spore count (Sandoval & Ordoñez, 2019).
was negatively correlated with available P and soil organic In another study, including five different successional
carbon content, and had a positive correlation with soil water stages (pioneer, early-intermediate, intermediate, late-
content and pH. Mahecha-Vásquez et al. (2017), in a study intermediate, and old mature forest), also in the department
done in banana agricultural systems in several Colombian of Cundinamarca, Peña & Botía (2018) did not find significant
departments with high soil P availability, reported low AMF differences in the number of AMF spores in the soil from
species diversity, with species composition being exclusive different successional stages. However, they reported an
to each site. They also found that species richness was inverse unimodal trend between the number of AMF spores
positively correlated with pH, finding more AMF species in in the soils and time after abandonment, with old grown
plantations with higher pH. forest and the “pioneer” stage showing greater abundances
Studies using metagenomic approaches have increased of AMF spores in the soil. This same trend was found by
in recent years. A study done by Peña-Venegas et al. Peña-Venegas et al. (2021) in rubber plantations from the
(2021) comparing traditional morphological and molecular Caquetá department, where plantations over nine years old
approaches in rubber plantations in Caquetá found, with showed a higher richness of AMF than younger plantations.
both approaches, that Glomeraceae was the dominant García et al. (2004) studied the richness of AMF morphotypes
taxon. However, molecular data showed a higher diversity and root mycorrhizal colonisation in plants in different
and more significant differences in species composition developmental stages (seedlings, saplings, and adults)
among sites when compared with spore morphotype in well - preserved montane forest and disturbed paramo
communities near Bogotá. They found that individuals in the number of collections, but our ecological knowledge of
early stages of development growing in sites with minor them is still incomplete. Oak forests are a characteristic
disturbance showed higher mycorrhizal colonisation than ecosystem of the Colombian Andes, present near main
adult individuals or areas with anthropogenic disturbance. cities and universities, and therefore there has been a lot of
In addition, when comparing mycorrhizal colonisation interest in their study. Vargas & Restrepo (2020) and Peña-
between dry and rainy seasons, there was a higher degree Venegas & Vasco-Palacios (2019) documented many species
of mycorrhizal colonisation during the rainy season in all associated with oak forests in Colombia. However, very few
species. In terms of AMF species richness, they report a studies have analysed the influence of abiotic factors on
decrease in richness in the sites with less intervention. EcM systems in Colombia to date. The only study comparing
EcM fungal communities associated with Quercus humboldtii
Ectomycorrhizal (EcM) fungi in Colombia is that from Sanchez-Tello (2021). This author
In Colombia, EcM-dominated systems are rare and compared EcM fungal communities in natural and urban
represented mainly by oak-dominated systems in Andean ecosystems in the Cundinamarca department using Illumina
montane forests with Quercus humboldtii as the primary host sequencing data. This author reports a trend for oak trees
and in Dipterocarpaceae/Fabaceae ectomycorrhizal systems growing in natural forests to host a higher species richness
in the Amazonian lowland forests (Peña-Venegas & Vasco- of EcM fungi than oaks growing in Bogotá. In addition, there
Palacios, 2019). The ecology of EcM fungi in Colombia has was a substantial change in EcM fungal species composition
been mostly studied on the basis of collections of fruiting between environments, with Russula and Lactarius being
bodies (e.g., Vargas & Restrepo, 2020). However, some dominant in natural ecosystems while Scleroderma and
recent studies have used metagenomic approaches (e.g., Laccaria dominate in urban trees. These changes in species
Vasco-Palacios et al., 2020). richness and composition could have been associated with
The best-studied systems in the country in terms of higher levels of nitrogen and other pollutants in urban versus
EcM ecology are the white sand and terra-firme Amazonian rural areas, as well as gardening practices applied to urban
forests, with several studies including molecular data and trees.
fruiting body surveys. Vasco-Palacios et al. (2018) studied Given the low number of native EcM host plant species,
EcM fungi associated with Dicymbe and Aldina trees in there has been a notorious the introduction of many exotic
white-sand forests in the department of Amazonas. Based EcM host species such as Pinus spp., Eucalyptus spp., and
on molecular data from fruiting body surveys and root tips, Acacia spp. to Colombia, all of which have been introduced
these authors found a high level of endemism and diversity with their associated EcM fungal species. Even though
in EcM fungi with a total of 48 species. They also found there are many potentially introduced and naturalised EcM
that some species reported for these hosts in Colombia fungal species in Colombia, the only species that has been
have already been reported as associated with congeneric studied in detail is Amanita muscaria. Vargas et al. (2019)
hosts in distant sites of Guyana, showing the possibility of documented A. muscaria in exotic tree species and the
long-distance dispersal of EcM fungal species. Also, Vasco- establishment of the species in native EcM host plants like
Palacios et al. (2020) studied fungal communities from Quercus humboldtii. These authors also reported that A.
terra-firme forests dominated by AMF trees (mixed forest) muscaria genotypes found in Colombia have Eurasian origin.
and EcM trees (Pseudomonotes tropenbosii) and white sand
forests with the presence of Dicymbe and Aldina trees. Saprotrophic fungi
These authors found, based on 454-pyrosequencing, a high Fungi, being heterotrophic organisms, use organic matter as
diversity of soil fungi, with the most important soil variables a source of energy for their development, participating in the
structuring the EcM fungal community being soil carbon degradation of organic matter, releasing simple compounds,
content and pH. They also found that soil type and not plant and contributing to carbon cycling in ecosystems (Robledo
community composition were the most important factors & Urcelay, 2009). Within these saprotrophic fungi are the
determining EcM fungal community composition changes. In wood - degrading fungi, composed mainly of the polyporoid
the same geographic area, Peña-Venegas & Vasco-Palacios group. One of the characteristics of this group is that they
(2019) evaluated the composition of AMF and EcM fungal are chemoorganotrophs, using organic compounds through
communities in mature and secondary terra-firme forests oxidation as suppliers of hydrogen to produce energy
and after cassava cultivation. They reported a high species (Schmidt, 2006). These fungi could generate a brown rot
diversity of both AMF and EcM fungi after abandoned (degradation of cellulose and hemicellulose) or a white
transitory cultivars of cassava, suggesting that disturbance rot (degradation of cellulose, hemicellulose, and lignin)
associated with this agricultural practice does not cause a (Schmidt, 2006). Given Colombia’s high ecosystem diversity,
permanent loss of soil fungal biodiversity. These authors many wood - degrading species have been reported to
also found that AMF and EcM fungi coexist in these forests. date. However, ecological studies regarding this group of
Still, this equilibrium could be altered after disturbance, organisms in Colombia are scarce. Studies on temperate
given that EcM host plants are primarily associated with ecosystems have reported that fungal species that cause
mature forests. brown rot are mainly associated with conifer - dominated
Oak forests are the most studied system in terms of forests. By contrast, species that generate white rot are
primarily associated with hardwoods (Gilbertson & Ryvarden, authors isolated 12 fungi belonging to the genera Curvularia,
1993). However, this hypothesis has not yet been tested in Fusarium, Stemphylium, and Rhizoctonia. Lizarazo-Medina et
tropical ecosystems, such as those from Colombia. al. (2015) analysed leaf and root tissues from Cattleya spp.
Wood fungi are not the only ones that recycle nutrients. growing under greenhouse conditions, finding differences
Several studies have shown that mycorrhizal fungi can in the composition of endophytes according to tissue type,
decompose organic nutrients (Hodge et al., 2001; Aristizábal with Aureobasidium, Botryotrichum, and Colletotrichum being
et al., 2004; Gui et al., 2017). In Colombia, this hypothesis the only genera in both tissues. Rodríguez & Lora (2016)
has been tested in two studies. Aristizábal et al. (2004), studied the endophytes of orchids in Quindío. These authors
studying fungal colonisation in the litterfall of Myrcia identified four genera of endophytic fungi from root isolates:
parvifolia, M. pubescens, and Paepalanthus sp. at different Tulasnella, Ceratobasidium, Sclerotium, and Colletotrichum.
soil depths in the Andean region of Colombia, reported the Pastures have also been studied to detect endophytes,
presence of hyphae and vesicles of AMF in the decomposing with Festuca spp. and Phalaris aquatica being studied by
litterfall from all the species and study sites, showing that Serrato (2019). This author found that Fusarium spp.,
this group of fungi could potentially obtain nutrients from Rhizoctonia sp., and Penicillium sp. are potentially non-
decomposing organic matter. In a similar study, Posada systemic endophytic fungi in grasses. Endophytic fungi
et al. (2012), showed that litterfall colonisation increases associated with grasses have also been shown to have
with soil depth in the Colombian Amazon using litterfall of the ability to solubilise phosphates. Pérez et al. (2012),
Eugenia sp. and Syzygium sp. However, saprotrophic fungi through the isolation of endophytes in grassroots in the
decrease with depth due to lower nutrient availability, department of Sucre cattle farms, isolated a total of
giving an advantage to AMF in colonising and decomposing 43 endophyte morphotypes, among which six species
the litterfall. Other studies on the decomposition of belonging to the genera Aspergillus, Penicillium, and
organic matter in the Colombian Amazon region identified Paecilomyces were able to solubilise phosphates. It is
Trichoderma sp. as an important species colonising late- also possible that endophytic fungi participate in nitrogen
stage organic matter, suggesting a primary lifestyle as a cycling, as reported by Saganome (2016), who studied
decomposer (López-Quintero et al., 2013). species isolated from the aquatic macrophyte species
Lemna sp. in the Cundinamarca department and reported
Endophytic fungi that isolates in the genera Penicillium sp. and Cladosporium
The richness of endophytic fungi is very high in the tropics, sp. were capable of oxidising ammonium and nitrite.
with at least one unique species of these endosymbionts per In addition, various studies have focused on finding
plant species (Arnold & Lutzoni, 2007; Lizarazo-Medina et al., possible applications of endophytic fungi, such as evaluating
2015). This richness has been little explored in Colombia, their antagonistic capacity against plant pathogens or
but it is expected to be high, given the great diversity of their antimicrobial activity (González, 2015; Trujillo, 2016;
plants reported for the country (Diazgranados et al., 2019). Montoya, 2019; Mosquera et al., 2020). However, we
Until now, the study of endophytic fungi for the country consider that these studies are mainly associated with
has been directed towards the knowledge of their species phytopathology and therefore they are not included in this
diversity and potential applications, especially in agriculture. chapter.
Vega et al. (2010) studied coffee plants’ leaves, stems, and
beans from Colombia, Hawaii, Mexico, and Puerto Rico. A Diversity and community structure
total of 267 endophytic fungi were isolated from Colombian Most specimens from the HUA herbarium collection
specimens, corresponding to 32% of all the species from all belonged to Basidiomycota (90%), given that this phylum
studied countries. Among the species identified through the includes most of the macromycete species. Polyporaceae,
analysis of the ITS region were Penicillium brevicompactum, Marasmiaceae, and Agaricaceae were the most abundant
P. brocae, and P. oxalicum, all these isolated from leaves. families in lowland and mixed montane forests, while in
Additionally, Salazar & Cepero de García (2005), in a study oak forests, the most abundant families were Russulaceae,
based on isolates from rose leaves (Rosa hybrida) collected Amanitaceae, and Cortinariacea. These differences between
in urban areas of Bogotá, were able to identify 31 isolates, montane forest types are probably due to their dominant
with Nigrospora oryzae, Xylaria spp., Aureobasidium spp., and mycorrhizal type. Saprotrophic fungi were prevalent in mixed
Acremonium sp. being the most isolated endophytes. montane forests compared with EcM fungi in oak forests.
The diversity of endophytes associated with orchids of The Amazonian lowland forests showed a higher proportion
the genera Vanilla, Stanhopea, and Maxillaria has also been of Ganodermataceae (9%) than all other ecosystem types.
studied. Ordóñez et al. (2012) isolated species from the In comparison, extra-Amazonian lowland forests showed a
roots of different Vanilla plants growing in the wild, reporting higher proportion of Xylariaceae (16%) than other ecosystem
a high diversity. In addition, when inoculating the Vanilla types (Figure 1a).
plants with the isolates, they found that some isolates Regarding fungal functional groups, the most important
impacted the aerial biomass, root length, and plant height. functional group in Amazonian lowland forests, extra-
In addition, Córdoba-Díaz et al. (2015) analysed the roots Amazonian lowland forests, and mixed montane forests
of the Stanhopea tricornis orchid in Valle del Cauca. These was “wood saprotroph” (36%, 52%, and 31%, respectively).
FIGURE 1. Number of specimens per ecosystem type for: A the 15 most abundant families and B fungal functional groups in Colombia.
FIGURE 2. Distribution by ecosystem type of fungial species reported as causing white rot, brown rot, or unknown decomposition type.
At the same time, the most important functional group and 1,360, respectively. Mueller et al. 2007 proposed that
in oak forests was, as expected, EcM (58%). The second there could be a 5:1 ratio of plant species to macrofungus
most important functional group for oak forests, mixed species in tropical regions. Based on that number, they
montane forests, and extra-Amazonian lowland forests was calculated an expected number of macrofungi for Costa
“litter saprotroph” (18%, 30%, and 28%, respectively). In Rica of 2,000 species and predicted that about half of the
comparison, EcM was the second most important group macrofungi from tropical South America had been recorded
for Amazonian lowland forests (22%) (Figure 1b). This high to date (6,595 out of 14,000 species). On the basis of
abundance of EcM fungi was expected for Amazonian these calculations, our estimated number of macrofungus
lowlands, given the presence of two important EcM host species for Colombia appears conservative. When
plants in this area. We also found a higher number of comparing the alpha diversity patterns among ecosystem
species that produce white rot (33%) compared with brown types based on the species-accumulation curves, the most
rot (2%; Figure 2) across all ecosystem types (Figure 2). diverse ecosystem was the Amazonian lowlands, with 524
With montane forests showing a lower proportion of white- species and an estimated number of species of 638 and
rot species compared with lowlands (Figure 2). These 975 based on the bootstrap and the Chao1 estimators,
findings are consistent with previous studies concluding respectively. This higher diversity is evidenced in the very
that most forests in Colombia comprise “hardwood” species steep slope shown in the species accumulation curve when
(132 gymnosperms versus 25,870 angiosperms, according compared to the other ecosystems (Figure 3b). The second
to SiB Colombia, 2020), but this hypothesis remains to be most diverse ecosystem type was oak forests (Figure 3b),
tested. However, it is striking that most species had no with 474 species and an estimated number of species
known decomposition type (65%). of 566 and 847 based on the bootstrap and the Chao1,
Regarding alpha diversity, a total of 1,138 species respectively estimators. The mixed montane forests and
were reported for the country. The species-accumulation extra-Amazonian lowland forests showed very similar alpha
curve, including all specimens, did not reach a plateau diversity patterns (Figure 3b) with 430 and 422 species,
(Figure 3a), meaning that there are still many species of respectively, and an estimated number of species between
macrofungi to discover. The potential number of species 518 and 856 based on the bootstrap and the Chao1
based on the Chao1 and bootstrap estimators were 1,867 estimates, respectively. The higher alpha diversity of
A B
FIGURE 3. Species accumulation curves of fungi and 95% confidence intervals for Colombia, based on a specimen dataset from the
Universidad del Antioquia Herbarium (HUA). A Curves for the total number of specimens, B Curves per ecosystem type. Abbreviations: OF-
Oak forests, MMF- Mixed montane forests, ALF- Amazonian lowland forests, EALF- Extra-Amazonian lowland forests.
FIGURE 4. Correlation matrix network graphic with the main ecosystems of Colombia represented by specimens from the Universidad del
Antioquia Herbarium (HUA). Abbreviations: OF- Oak dominated forest, MMF- Mixed montane forest, ALL- Amazonian lowland forests, EALF-
extra-Amazonian lowland forests.
the Colombian lowland Amazonian rainforest reveals Trichoderma Restrepo KJ, Montoya MI, Henao P, Gutiérrez LA, Molina LP (2019)
strigosellum sp. nov. and other species. Antonie Van Leeuwenhoek, Caracterización de hongos micorrízicos arbusculares de suelos
104(5), 657-674. doi: 10.1007/ s10482-013-9975-4. ganaderos del trópico alto y trópico bajo en Antioquia, Colombia. Idesia
López F (2009) Estudio de la micorriza arbuscular en Guadua angustifolia (Arica), 37(1), 35-44. doi: 10.4067/S0718-34292019005000301.
Kunth establecida a diferentes altitudes, en suelos de la zona Robledo G, Urcelay C (2009) Hongos de la madera en árboles nativos del
cafetera colombiana. Undergraduate thesis. Bogotá DC: Pontificia centro de Argentina. Córdoba: Ed. Univ. Nacional. 224p.
Universidad Javeriana. Rodríguez J, Lora F (2016) Micobiota endófita del género Maxillaria Ruiz &
Mahecha-Vásquez G, Sierra S, Posada R (2017) Diversity indices using Pav. (Orchidaceae) en dos reservas naturales del departamento del
arbuscular mycorrhizal fungi to evaluate the soil state in banana Quindío. Revista de Investigaciones Universidad del Quindío, 1, 151. doi:
crops in Colombia. Applied Soil Ecology, 109, 32-39. doi: 10.1016/j. 10.33975/riuq.vol28n1.32.
apsoil.2016.09. 017. Saganome GM (2016) Diversidad de microorganismos endófitos y su papel
Mata M, Halling RE, Mueller GM, Aragón-Quesada C (2003) Macrohongos en el ciclo de nitrógeno en macrófitas flotantes del Humedal Meandro
de Costa Rica Vol. 2. Santo Domingo: Editorial Santo Domingo de del Say. Undergraduate thesis. Bogotá DC: Universidad de la Salle. 28p.
Heredia. 240p. Salazar CS, Cepero de García MC (2005) Aislamiento de hongos endofitos
Montoya SHM (2019) Evaluación in vitro de la capacidad antagónica en rosa (Rosa hybrida) en Bogotá, Colombia. Revista Iberoamericana de
de hongos endófitos aislados de Moringa oleifera en el control Micología, 22(2), 99-101. doi: 10.1016/S1130-1406(05)70016-4.
de Pestalotia palmarum. Undergraduate thesis. Bucaramanga: Sanchez-Tello JD (2021) How ectomycorrhizal communities vary from natural
Universidad de Santander. 79p. to urban ecosystems: Quercus humboldtii as a study case in the tropical
Mosquera WG, Criado LY, Guerra BE (2020) Actividad antimicrobiana de Andes. Undergraduate thesis. Bogotá DC: Universidad del Rosario. 24p.
hongos endófitos de las plantas medicinales Mammea americana Sandoval KS, Ordoñez DAKM (2019) Presencia de hongos micorrízico
(Calophyllaceae) y Moringa oleifera (Moringaceae). Biomédica, 40(1), arbusculares en tres áreas de intervención antrópica en la vereda
55-71. doi: 10.7705/ biomedica.4644. Albania del municipio Tibacuy-Cundinamarca. Undergraduate thesis.
Mueller GM, Schmit JP, Leacock PR, Buyck B, Cifuentes J, Desjardin DE, Bogotá DC: Universidad Distrital Francisco José de Caldas. 108p.
Halling RE, Hjortstam K, Iturriaga T, Larsson KH, Lodge DJ (2007) Schmidt O (2006) Wood and tree fungi: biology, damage, protection, and use.
Global diversity and distribution of macrofungi. Biodiversity and Berlin: Springer. doi: 10.1007/3-540-32139-X.
Conservation, 16(1), 37-48. doi: 10.1007/s10531-006-9108-8. Serrato S (2019) Aislamiento y caracterización de hongos endofíticos
Nualart N, Ibánez N, Soriano I, López-Pujol, J (2017) Assessing the provenientes de pasto para uso agrícola. Undergraduate thesis. Bogotá
relevance of herbarium collections as tools for conservation biology. DC: Pontificia Universidad Javeriana. 57p.
The Botanical Review, 83(3), 303-325. doi: 10.1007/s12229-017- SiB Colombia (2020) Biodiversidad en Cifras, Sistema de Información sobre
9188-z. Biodiversidad de Colombia. Available at: https://cifras.biodiversidad.co/.
Ordóñez NF, Tupac Otero J, Díez GMC (2012) Hongos endófitos de Tirado-Ardila A (2017) Presencia de esporas de hongos micorricíco
orquídeas y su efecto sobre el crecimiento en Vanilla planifolia arbusculares en suelos del bosque alto andino, Parque Natural Chicaque.
Andrews. Acta Agronómica, 61(3), 282-290. Undergraduate thesis. Bogotá DC: Universidad Distrital Francisco José
Orjuela DM (2018) Microorganismos endófitos como alternativa para el de Caldas. 58p.
control de hongos patógenos asociados al cultivo de aguacate en Trujillo D (2016) Aislamiento de un hongo endófito de Piper crassinervium
Colombia. Master thesis. Bogotá: Universidad Nacional de Colombia. (Piperaceae) y su evaluación antibacteriana. Undergraduate thesis.
109p. Pereira: Universidad Tecnológica de Pereira. 90p.
Peña DY, Botía FL (2018) Presencia de esporas de hongos micorrízico Vargas N, Gonçalves SC, Franco-Molano AE, Restrepo S, Pringle A (2019) In
arbusculares en cinco estados sucesionales de bosque altoandino Colombia, the Eurasian fungus Amanita muscaria is expanding its range
Colombiano. Revista de Investigación Agraria y Ambiental, 9(2), 135- into native, tropical Quercus humboldtii forests. Mycologia, 111(5), 758-
147. doi: 10.22490/21456453.2194. 771. doi: 10.1080/00275514.2019.1636608.
Peña-Venegas C, Vasco-Palacios AM (2019) Endo- and ectomycorrhizas Vargas N, Restrepo S (2020) A checklist of ectomycorrhizal mushrooms
in tropical ecosystems of Colombia. In: Pagano MC, Lugo MA (eds). associated with Quercus humboldtii in Colombia. In: Perez-Moreno J,
Mycorrhizal Fungi in South America 1st edition. Berlin: Springer. Pp. Guerin-Laguette A, Flores Arzu R, Yu F (eds). Mushrooms, humans and
111-146. doi: 10.1007/978-3-030-15228-4_6. nature in a changing world: perspectives from ecological, agricultural and
Peña-Venegas C, Castro Torres D, León D (2020) Riqueza y abundancia de social sciences. Berlin: Springer. Pp. 425-450. doi: 10.1007/978-3-030-
organismos del suelo en el gradiente altitudinal Andino-Amazónico del 37378-8_16.
Camino Andakí, Caquetá (Colombia). Colombia Amazónica, 12, 145- Vasco-Palacios AM, Hernández J, Peñuela-Mora MC, Franco-Molano AE,
164. Boekhout T (2018) Ectomycorrhizal fungi diversity in a white sand
Peña-Venegas C, Sterling Cuéllar A, Andrade T (2021) Arbuscular forest in western Amazonia. Fungal Ecology, 31, 9-18. doi: 10.1016/j.
mycorrhization in Colombian and introduced rubber (Hevea funeco.2017. 10.003.
brasiliensis) genotypes cultivated on degraded soils of the Amazon Vasco-Palacios AM, Bahram M, Boekhout T, Tedersoo L (2020) Carbon
Region. Agriculture, 11 (4), 361. doi: 10.3390/agriculture11040361. content and pH as important drivers of fungal community structure in
Pérez A, De la Ossa J, Montes D (2012) Hongos solubilizadores de three Amazon forests. Plant and Soil, 450(1), 111-131. doi: 10.1007/
fosfatos en fincas ganaderas del departamento de sucre. Revista s11104-019-04218-3.
Colombiana de Ciencia Animal – RECIA, 4(1), 35-45. doi: 10.24188/ Vega FE, Simpkinsa A, Aimeb C, Posada F, Peterson SW, Rehner SA, Infante
recia.v4.n1.2012.263. F, Castillo A, Arnold AE (2010) Fungal endophyte diversity in coffee
Põlme S, Abarenkov K, Tedersoo L et al. (2020) FungalTraits: a user- plants from Colombia, Hawaii, Mexico, and Puerto Rico. Fungal Ecology,
friendly traits database of fungi and fungus-like stramenopiles. Fungal 3(3), 122-138. doi: 10.1016/j.funeco.2009.07.002.
Diversity, 105(1), 1-16. doi: 10.1007/s13225-020-00466-2. Wollan AK, Bakkestuen V, Kauserud H, Gulden G, Halvorsen R (2008)
Posada RH, Madriñan S, Rivera EL (2012) Relationships between the Modelling and predicting fungal distribution patterns using herbarium
litter colonisation by saprotrophic and arbuscular mycorrhizal fungi data. Journal of Biogeography, 35(12), 2298-2310. doi: 10.1111/j.1365-
with depth in a tropical forest. Fungal Biology, 116(7), 747-755. doi: 2699.2008.01965.x.
10.1016/j.funbio.2012. 04.003. Zanne AE, Abarenkov K, Afkhami ME, Aguilar‐Trigueros CA, Bates S,
Posada RH, Sánchez de Prager M, Heredia-Abarca G, Sieverding E Bhatnagar JM, Busby PE, Christian N, Cornwell WK, Crowther TW,
(2018) Effects of soil physical and chemical parameters, and farm Flores‐Moreno H (2020) Fungal functional ecology: bringing a trait‐based
management practices on arbuscular mycorrhizal fungi communities approach to plant‐associated fungi. Biological Reviews, 95(2), 409-433.
and diversities in coffee plantations in Colombia and Mexico.
Agroforestry Systems, 92(2), 555-574. doi: 10.1007/s10457-016-
0030-0.
R Core Team (2021) R: A language and environment for statistical Chlorociboria aeruginascens
computing. R Foundation for Statistical Computing, Vienna, Austria. [Robert Lücking]
Pleurotus sp.
[Nataly Gómez-Montoya]
150 CATALOGUE OF FUNGI OF COLOMBIA
Chapter 12
Useful Fungi of Colombia
Natalia Vargas1, Nataly Gómez-Montoya1,2, Rocío Peña-Cañón1 & Germán Torres-Morales1*
1
Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Bogotá, Colombia.
2
Laboratorio de Taxonomía y Ecología de Hongos – TEHO, Instituto de Biología, Facultad de Ciencias Exactas y Naturales, Universidad de Antioquia – UdeA,
Calle 70 No. 52-21, Medellín, Antioquia, Colombia.
ABSTRACT
Colombia is considered a megadiverse country with a substantial number of fungal species, many of them regarded as useful
throughout the history of the country. However, many of the uses for fungi have been lost due to the erosion of knowledge.
Making this knowledge visible and compiling it is of great importance as a potential resource for the bioeconomic development
of Colombia and as a part of the livelihoods of society. Therefore, the goal of this study was to discuss the main uses and to
categorise the useful fungi of Colombia based on a systematic review of secondary data. The taxonomy for the scientific names
was checked, the species’ uses were grouped by general and specific categories, and the geographical distribution was recorded
by Colombian regions. As a result, a list of 382 species of useful fungi was obtained and classified into eight general categories of
use. The general category with the highest number of uses is food, followed by medicinal use. Despite several useful fungi having
been reported for Colombia, most of them are underutilised in each category. From this analysis, it is possible to highlight that
Colombia has an important number of useful fungi, most of which are not yet widely used in the country and constitute a potential
resource of natural ingredients and livelihoods, especially for local communities and commercial initiatives.
RESUMEN
Colombia es considerado un país megadiverso con una gran cantidad de especies de hongos. Muchos de ellos se han
considerado útiles a lo largo de la historia del país; sin embargo, gran parte de los usos se han dejado de lado debido a la pérdida
del conocimiento. Visibilizar y recopilar este conocimiento es de vital importancia como fuente potencial para el desarrollo
bioeconómico del país, y como parte de los medios de vida de la sociedad. Por lo tanto, el objetivo de este trabajo fue discutir
los principales usos y categorizar los hongos útiles de Colombia a partir de una revisión sistemática de información secundaria
y literatura gris de referencias bibliográficas. Con base en esta revisión, se verificó la taxonomía de los nombres científicos, se
agruparon los usos de las especies por categorías generales y específicas y se registró la distribución geográfica por regiones
colombianas. Como resultado de esta metodología, se obtuvo una lista de 382 especies de hongos útiles, clasificadas en ocho
categorías generales de uso. Respecto a los usos, la categoría general con mayor número de usos es la de alimento, seguida del
uso medicinal. A pesar del uso de especies de hongos, la mayoría de ellas están infrautilizadas en cada categoría. A partir de
este análisis, es posible resaltar que Colombia cuenta con una cantidad importante de hongos útiles, de los cuales la mayoría
no son ampliamente utilizados en el país y constituyen un recurso potencial de ingredientes naturales y medios de subsistencia,
especialmente para comunidades locales e iniciativas comerciales. Las número de publicaciones de investigaciones realizadas
en el país y un incremento en la inversión en investigación básica y aplicada, permitirá descubrir nuevas alternativas de uso
de los hongos para resolver situaciones que afrontamos como sociedad. En este sentido, la conservación de los hongos debe
hacerse de la mano con las comunidades y acorde a su aplicación en diferentes campos, buscando hacer un uso sostenible y
prevenir los impactos negativos que puedan afectar las poblaciones de hongos y su supervivencia.
Wild macrofungi constitute an essential non-wood forest et al., 2021; see also Chapter 3). However, no study has ever
product that provide different ecosystem services and have categorised the different uses of fungi found in Colombia.
traditionally been used by many human cultures worldwide Many fungi reported as useful in other countries are not
(Gaya et al., 2021). They are used for many human activities, recognised as such in Colombia due to a lack of research
such as those in the biotechnology, food and pharmaceutical or erosion of traditional knowledge on this topic. This study
industries, and even in the rituals of different indigenous aimed to summarise information and categorise the useful
cultures (Gaya et al., 2021). Many local reports evidence the fungi of Colombia, contributing baseline information about
use of edible and medicinal macrofungi around the world. their uses, levels of use, and potential uses.
For example, areas with tendencies to use fungi include
southern and eastern Europe, Africa, Mexico, Turkey, and
MATERIALS AND METHODS
most of Asia (Boa, 2004). In the Americas, a high cultural
richness related to the collection, recognition, and use of We reviewed the literature related to Colombia’s useful
macrofungi has been widely documented in Mexican culture species of fungi by using the Scopus, Google Scholar, and ISI
(Ruan-Soto et al., 2014). Web of Knowledge portals. We searched for fungal species
Colombia’s wide range of climatic zones provides a high lists in Colombia and publications on the utility of fungal
diversity of habitats where organisms can grow (World Atlas, species based on the literature published in Colombia.
2021). Many papers and field guides have documented Information on the useful characters of some species
the funga occurring in Colombian ecosystems, and a high that occur in Colombia was included from publications in
number of species is estimated to occur in the country (Gaya other countries. All references related to uses, distribution,
ecology, and taxonomy of species included in this study can in the food category, followed by medicines, poisons,
be consulted in the ColFungi online portal (www.colfungi.org) environmental uses, biotechnology, social uses, biological
as the species dataset included in this study was included control, and materials. We will focus in the following sections
in that platform. Species use was systematised using a on the discussion: 1. the general aspects of useful fungi, 2.
modified version of Cook’s Economic Botany Standard research findings in Colombia, and 3. cases of fungi used by
categories of use (Cook, 1995) (Table 1). The search effort communities in Colombia.
was carried out for six months between 2020 and 2021
using English and Spanish keywords and modifiers, such as Useful fungi of Colombia
biofertilisers, biological control, biotechnology, edible fungal According to the literature review, 179 species were classified
species, esoteric, fungal enzymes, magic uses, medicinal in the food category. Many macrofungi are characterised by
fungi, pigments, social uses, and useful fungi. Graphics producing sporocarps with different morphologies, several
were made in Excel and the map of Colombian regions of them being edible useful species (Boa, 2004; Prescott
was modified from IGAC (1997). The terminology used in et al., 2018). Agaricales (Basidiomycota) is the order
this chapter is explained as follows: 1. Useful: a species
comprising most of the useful species in the food category,
of fungus categorised according to the modified Economic
edible subcategory (Figure 2), which coincides with the high
Botany Standards, regardless of whether it is used or not
number of reported edible fungi in this taxonomic order
in Colombia; 2. Used: reports of fungi species used by
(Prescott et al., 2018). Several yeast species involved in the
communities in Colombia; 3. Assessed use: research on
production of fermented beverages are classified in the order
the applicability on fungal species in laboratory or field
in Saccharomycetales (Ascomycota). Species in this order
conditions in Colombia; and 4. Underutilised: fungi species
have been consistently involved in the metabolic processes
occurring in Colombian ecosystems without reports of its
of brewing alcoholic beverages (Hyde et al., 2019).
use or assessment in Colombia, but their use is reported in
A total of 131 fungal species were classified in the
other countries.
medicines category. Agaricales (Basidiomycota) has the
highest number of useful species in this category, sometimes
RESULTS AND DISCUSSION
with important applications, including the treatment
A total of 382 species of useful fungi occurring in Colombia of digestive and blood disorders, and as the source of
were recorded. According to a modified version of the anticancer and antioxidants bioactive compounds (Table 1).
Economic Botany Data Collection Standards, the useful Some saprotrophic fungi in Polyporales (Basidiomycota)
fungi of Colombia are classified into eight general categories and entomopathogenic fungi (Hypocreales, Ascomycota)
(Figure 1, Table 1), with the highest number of species (Figure 2) also have great qualities as medicines, being
recognised as essential organisms in traditional medicine,
particularly in Asia (Boa, 2004). Saprotrophic fungal
species classified in this study as food, medicines, and
biotechnology have previously been reported in all regions
of Colombia, with saprotrophic fungi showing the highest
number of species in each region of the country (Figure 3).
A total of 26 species were assigned to the biological
control category, which comprises useful fungi mainly placed
in Hypocreales (Ascomycota) (Figure 2). In this order, most
entomopathogenic fungi are classified and can function as
natural insecticides. Entomopathogenic fungi infect their
hosts (arthropods) by penetrating the insect cuticles and
colonising their bodies with fungal microscopic structures
(Roy et al., 2006; Charnley & Collins, 2007; Wang & Leger,
2014). Most products used in biological control are based
on asexual stages of entomopathogenic fungi of Hypocreales
(Wang & Leger, 2014). The entomopathogenic species that
occur in Colombia found in this category have been reported
from all regions of Colombia, except the Caribbean. They are
mostly reported in the Andean region (Figure 3).
Regarding the social uses category, a total of 33
species were classified mostly in Agaricales, particularly
the hallucinogenic Psilocybe species, which are essential
sources of compounds, including psilocybin, the focus of
many antidepressant studies (Davis et al., 2020). Some
FIGURE 1. Percentage of useful fungi of Colombia by category Psilocybe species that occur in Colombia are used in Mexico
of use. as a healing element in religious ceremonies, as important
of crops (Castro-Toro & Rivillas-Osorio, 2012). Some studies Pseudallescheria boydii has potential in bioremediation
have also been carried out in native orchid mycorrhizae, because it presents gene families involved that are in
including species of Ceratobasidium (Mosquera-Espinosa et the degradation of hydrocarbons, such as chloroalkanes,
al., 2010; Ordóñez et al., 2012; Valadares et al., 2015) that chlorocyclohexanes, chlorobenzenes, benzoates, toluene,
provide plant protection or growth stimulation and have been xylene, and polycyclic aromatic hydrocarbons (Morales
reported to control fungal plant pathogens on rice seedlings et al., 2017).
(Mosquera-Espinosa et al., 2013). The potential use of fungal A high percentage of species in the biotechnology
species in pollution control has been assessed mainly in category has been the target of several research projects
bioaugmentation, biostimulation, biodegradation, and natural in laboratories and research centres in Colombia (Figure
attenuation. This topic is currently a promising strategy that 2, Table 2). Particularly the ability of some species to
oil companies are being encouraged to implement (Trujillo- decolourise Azure B and Coomassie Blue in solid media
Toro & Ramírez-Quirama, 2012; García & Arango, 2019). has been evaluated, and has a potential application in
decolourisation technology (Arboleda et al., 2008) and in 2012; Bolívar-Anillo et al., 2016). Among them are some
the removal of colourants from wastewater (Holguín-Múnera endophytic species isolated from Frailejón ecosystems
et al., 2017). The potential biotechnological application with antimicrobial activity against Botrytis cinerea
of the enzymes laccase and cellobiose dehydrogenase and Phytophthora infestans (Miles et al., 2012). Some
from Lentinus swartzii, Lentinus crinitus, and Pycnoporus commercial fungal species are the target of research in
sanguineus have been tested for their potential application medicine. For example, that show potential antioxidant and
in decolourisation technologies (Chaparro et al., 2009). The anticancer activities or potential as nutraceuticals have
laccase enzymes from the edible species Pleurotus ostreatus been isolated, identified and analysed from basidiocarps of
were evaluated for potential application in the textile industry Ganoderma lucidum (Moreno et al., 2011) and P. ostreatus
(Pérez-Higuera, 2013). Given the importance of laccase (Duarte-Trujillo et al., 2020), respectively.
in the industry, the effect of Cordyceps nidus extracts on
the laccase activity of P. ostreatus was evaluated, revealing Fungi used in Colombia
an improvement in the enzymatic activity. Cordyceps nidus According to our search, ca. 60 species of useful fungi are
extract is a stimulus for other fungi’s enzymatic activity and currently used in rural and urban regions of Colombia.
biochemical properties (Durán-Aranguren et al., 2020). On
the other hand, the remarkable capacity of Lentinus crinitus Wild macrofungi used by communities in Colombia
to transform agricultural residues increases the content of
proteins, fibre, and nitrogen of the spent substrate, providing Wild macrofungi reported to be used by communities in
different products of interest for agri-food (animal feed) and Colombia (Franco-Molano et al., 2005; Ruíz & Henao, 2006;
agroindustry (Dávila et al., 2020). In addition, some Mucor Vasco-Palacios, 2002; Peña-Cañón & Henao-Mejía, 2014;
species produce proteases with potential applications in the Villalobos et al., 2017) were classified in the food, materials
food industry (Alves et al., 2005). (cosmetics), or medicines categories (Table 3). The food
Novel strains of the yeast Saccharomyces cerevisiae have category had the highest number of useful species (Table
been evaluated as candidates to improve ethanol production 3), with 36 wild edible species used by rural communities in
due to their ability to consume xylose in the fermentation Colombia, mainly in the departments of Boyacá, Amazonas,
of hydrolysates (Cifuentes-Triana, 2016). Also, the ability of and Putumayo. In addition, six commercial edible species
the yeast Meyerozyma guilliermondii to produce oil as raw (Agaricus bisporus, Auricularia auricula-judae, Lentinula
material for biodiesel has potential as a biodiesel feedstock edodes, Pleurotus ostreatus, P. eryngii, and Flammulina
(Ramírez-Castrillón et al., 2017). Finally, the growth and velutipes) are also used in urban areas.
enzyme production of the anaerobic fungus Neocallimastix In some regions of Colombia, rural communities have
frontalis, isolated from the rumen of a sheep from a tropical passed on traditional knowledge on the use of edible and
ecosystem, was evaluated in a range of carbon sources and medicinal macrofungi (Peña-Cañón & Henao-Mejía, 2014;
by varying the form of inoculation (mobile and vegetative Pérez & Piragauta, 2006; Ruíz-Roa et al., 2008; Sanjuan,
states) (Mayorga et al., 2005). 1999; Vasco-Palacios, 2006). Wild edible macrofungi are
Although several entomopathogenic species are used by rural communities in some municipalities in the
currently applied to control animal pests in Colombian department of Boyacá (Table 3). During the rainy season,
crops, some species are being assessed in field and several communities in this region harvest macrofungi for
laboratory conditions. For example, a strain of the species personal use thanks to the oral tradition that transmits
Cladosporium cladosporioides was evaluated against the knowledge from their elders (Ruíz & Henao, 2006; Peña-
spider mite Tetranychus urticae, an important polyphagous Cañón & Henao-Mejía, 2014; Pérez & Piragauta, 2006;
pest worldwide (Gámez-Guzmán et al., 2019) The biological Ruíz-Roa et al., 2008). Macrolepiota colombiana, known
control of colonies of Atta cephalotes with baits prepared as lechucitas or caicas, and species of Ramaria, known as
with sporulated cultures of Metarhizium anisopliae and manitas, guaras, or pericocas, have a cultural significance
Trichoderma viride has also been evaluated (López & for rural communities. Among other species used by rural
Orduz, 2002). Akanthomyces lecanii has been used for the communities in Boyacá are the ectomycorrhizal Porphyrellus
biological control of cattle ticks (Beltrán-Alzate et al., 2008). indecisus and some saprotrophic fungi such as Crepidotus
Likewise, conidia of M. anisopliae and Metarhizium rileyi have palmarum, Auricularia fuscosuccinea, Hericium erinaceus,
been tested as control agents against Spodoptera frugiperda Lentinula boryana, L. crinitus, Oudemansiella canarii, and
(Grijalba et al., 2018), Beauveria bassiana, Purpureocillium Cantharellus lateritius. All species are given common names
lilacinum, and Akanthomyces dipterigenus has been tested and are cooked according to local culinary preferences (Ruíz
in the control of the poultry red mite Dermanyssus gallinae & Henao, 2006; Peña-Cañón & Henao-Mejía, 2014).
(Machado-Agudelo, 2019) and B. bassiana. Cordyceps Peña-Cañón and Henao-Mejía (2014) contributed to the
fumosorosea and P. lilacinum were also tested for their knowledge of these fungi in the department of Boyacá,
ability to control Leptopharsa gibbicarina, an economically recording the use of 16 species by rural communities, of
important pest of oil palm plantations (Barrios et al., 2016). which 11 are reported as useful in the municipalities of
In the poisons category, some filamentous fungi with Pauna and Chiquinquirá. Pérez & Piragauta (2006) recorded
antimicrobial activity have been tested as antagonists of 23 species used by the communities of Arcabuco in the
other fungal plant pathogens in Colombia (Miles et al., department of Boyacá: 14 species used as food, four as
medicines, and three as insecticides. In the municipality of commercialise them in the town centre of the municipality
Moniquirá (Boyacá), 22 species were recorded, with 16 being of Gachantivá (N. Vargas, pers. observ.).
used as food, four as medicinal, and four as insecticides. Several studies have reported the use of fungi by
Ethnomycological explorations continue to document indigenous communities in the Amazon region. Since the
the traditional use of macrofungi among Colombian rural 1980s, the use of fungal species has been reported in
communities, expanding the list of municipalities in the indigenous ethnic groups. Sanjuan (1999) found that Inganos
department of Boyacá with a tradition of macrofungi use for indigenous communities from Caquetá use six species of
food and/or medicine. In fact, in a recent expedition to the fungi, known by them as kallambas, including Oudemansiella
municipality of Otanche (west of Boyacá), two edible species, canarii, Schizophyllum commune, and Favolus tenuiculus. At
Macrocybe titans and Favolus tenuiculus, were recorded as the beginning of the 2000s, Vasco-Palacios (2002), Franco-
useful. The abovementioned cases are examples of the Molano et al. (2005), and Vasco-Palacios (2006) elaborated a
personal use of native species, but fewer cases of direct review of the fungal species used by the Uitoto, Muiname, and
marketing by collectors are reported in Colombia (Peña- Andoke indigenous people of the middle Caquetá, reporting
Cañón & Henao-Mejía, 2014; Pérez & Piragauta, 2006). their edible and medicinal uses (Table 3). Species such as
Wild macrofungi are shared among people or relatives as Lentinula raphanica, Auricularia delicata, A. fuscosuccinea,
a gift and are exchanged for other products such as wheat Favolus tenuiculus, and three species of Lentinus were
(Peña-Cañón & Henao-Mejía, 2014). In the municipality reported as edible by the Amazonian indigenous people
of Gachantivá (Boyacá), some farmers collect macrofungi (Franco-Molano et al., 2005; Vasco-Palacios, 2002; Sanjuan,
such as Agaricus spp. near their homes in rural areas and 1999; Vasco-Palacios, 2006).
For example, in the medicines category, the species sawdust with wood shavings and rice or sorghum bran, dry
Pycnoporus sanguineus is reported to treat oral mycosis, wood from plantations of Quercus trees (oaks) (Rodríguez &
and Daldinia concentrica is used to treat skin sores (Franco- Jaramillo, 2005), and a mixture of sawdust from the coffee
Molano et al., 2005). Other species such as Bovista plumbea stem, coffee grounds, wheat bran, carbonate, gypsum, and
and Psilocybe cubensis are used to prevent and cure sugar (Varón, 2004) have been used in Colombia. To reduce
dermatological diseases (Villalobos et al., 2017). In the social production time and costs, and to increase the production
uses category, women in Wayuu communities (department of compounds of interest such as beta-glucans, different
of La Guajira) use the spores of some fungi species as studies have been carried out in liquid media, allowing the
sunscreen in religious rituals and dances (Villalobos et fungus and its compounds to be more easily absorbed by the
al., 2017). In the Andean region, rural communities use human body, and also making them soluble and thermostable
the term “dangerous” to refer to the mushroom species (Puerta-Marín & Zapata-Cardona, 2006). Lentinula edodes is
Psilocybe cubensis (Ruíz & Henao, 2006), reporting that originally from Asia, and the strains used in Colombia come
they are sought after by visitors with great interest in their from imported strains in international stocks, mainly from
hallucinogenic effects. Despite this, some local experts the United States, Japan, and China. Some studies have
refer to their medicinal use in hot compresses to treat boils tested its cultivation with substrates that produce optimal
(Pérez & Piragauta, 2006). biological efficiencies (Arias-García et al., 2008; Arredondo-
In the biological control category, several entomo- Peña & Pérez-Martínez, 2007).
pathogenic species are available in markets as agents to In Colombia, some entities help to enhance the cultivation
control pathogens in crops. Colombia has a strong tradition of macrofungi. For example, the National Federation of
in this research field, and companies and research institutes Fungi Growers of Colombia (FEDEFUNGICOL), a non-profit
are developing technologies to produce commercial products organisation, seeks to provide better support to its members
based on entomopathogens (Espinel-Correal et al., 2018). and more opportunities for human communities, sheltering
For example, different products based on Beauveria bassiana different branches of the fungiculture, such as producers of
control the coffee berry borer (Alves et al., 2002). Cordyceps oyster mushrooms, Ganoderma, and shiitake.
fumosorosea is used to control mites, thrips, whiteflies,
and Metarhizium anisopliae is formulated to control beetle Underutilised fungal species
larvae, leaf cutter ants, and mosquito larvae (Espinel-Correal The number of used and/or assessed useful species in
et al., 2018). Colombia compared to the underutilised species (reports
made in other countries) are shown in Figure 4. Regarding
Commercial macrofungi the fungal species present in Colombia and their utility, most
Regarding non-native macrofungi, a few well-known species categories, particularly food, materials, medicines, poisons,
are commercialised in markets or consumed in restaurants and social uses, have a higher number of underutilised
in urban cities. These species are distributed mainly in cool species than the number of used or assessed species.
climates, and their strains have been brought from other
countries for cultivation in Colombia. Agaricus bisporus, the
common champiñón, is the most commercialised species
in urban centres in Colombia, being sold in supermarkets,
squares and small groceries, and dairy and meat stores. The
species with the most reports of use in small businesses
and entrepreneurship in urban areas is the oyster mushroom
Pleurotus ostreatus. Different entities, small businesses
and research groups have developed studies to optimise
its cultivation, the development of value-added products,
and the analysis of supply chains (Rodríguez-Valencia &
Jaramillo, 2004; Rodríguez-Valencia et al., 2006; Arias-
García et al., 2008; López-Rodríguez et al., 2008; Montoya-
Barreto et al., 2009; Romero et al., 2010; Suárez-Arango,
2010; Jaramillo-Ruiz et al., 2011; Fernández-Uribe, 2014;
Vargas et al., 2019).
On the other hand, one of the most used and studied
species at an industrial level is Ganoderma lucidum, a species
commonly known as the king of fungi and longevity in Asia. In
Colombia, it is estimated that its research began 12 years
ago, and its industrial cultivation started approximately six
years ago, due to the interest generated by its medicinal
properties (Giraldo, 2013). Among the culture media used to FIGURE 4. Number of fungal species used and assessed in
grow G. lucidum, mixtures of substrates such as hardwood Colombia vs underutilised species.
CONCLUSIONS Barrios TCE, Bustillo PAE, Ocampo RKL, Reina CMA, Alvarado MHL (2016)
Eficacia de hongos entomopatógenos en el control de Leptopharsa
In Colombia, the use of wild fungal species is limited, gibbicarina (Hemiptera: Tingidae) en palma de aceite. Revista
and it is an infrequent practice, probably given that their Colombiana de Entomología, 42(1), 22. doi: 10.25100/socolen.
recognition and use has been forgotten in some areas of v42i1.6665
Bautista-Gonzalez JA, Moreno-Fuentes A (2014) Los hongos medicinales
the country, even though fungi were part of pre-Hispanic de México. In: Moreno-Fuentes A, Garibay-Orijel R (eds). La
cultures (Velandia et al. 2008). Indeed, with the increase in Etnomicología en México. Estado del Arte-Red de Etnoecología y
research involving fungi and the documentation of its uses Patrimonio Biocultural (CONACyT). México DF: Universidad Autónoma
in unexplored regions, we will understand and elucidate del Estado de Hidalgo. Pp. 145-176.
Beltrán-Alzate C, Gutiérrez AI, Saldarriaga Y (2008) Patogenicidad de
whether Colombian citizens fit into one of the two differential Lecanicillium lecanii (Fungi) sobre la garrapata Boophilus microplus
attitudes towards fungi, understood as mycophilia and (Acari: Ixodidae) en laboratorio. Revista Colombiana de Entomología,
mycophobia (Ruan-Soto et al., 2014). We showed data on 34(1), 90-97.
the useful fungi of Colombia, highlighting that a few species Boa E (2004) Wild edible fungi: a global overview of their use and
importance to people. Roma: FAO. 149p.
are used and some others have been assessed, but most of Bolívar-Anillo HJ, Orozco-Sanchez C, da Silva-Lima G, Franco dos Santos
them are still underutilised. Thus, different sectors, such as G (2016) Endophytic microorganisms isolated of plants grown
academia, research institutions, industry, and entrepreneurs in Colombia: a short review. Journal of Microbial & Biochemical
must further explore the potential uses and properties of the Technology, 8(6), 509-513. doi: 10.4172/1948-5948.1000335
Brough SG (1988) Navajo lichen dyes. The Lichenologist, 20(3), 279-290.
Colombian funga. The increase in investment in basic and doi: 10.1017/S0024282988000313.
applied research is an opportunity to explore and discover Castro-Toro AM, Rivillas-Osorio CA (2012) Trichoderma spp. Modos de
these properties. Finally, it is essential to highlight that the acción, eficacia y usos en el cultivo de café. Boletín Técnico Cenicafé
conservation of fungi should be done hand in hand as society (Issue 38). Chinchiná: FNC- Cenicafé. 31p.
Chaparro DF, Rosas DC, Varela A (2009) Aislamiento y evaluación de
starts to use or apply fungi in different fields. Planning to la actividad enzimática de hongos descomponedores de madera
prevent the impact of humans on fungal populations (e.g., (Quindío, Colombia). Revista Iberoamericana de Micologia, 26(4),
excessive macrofungi harvesting) is a key to their sustainable 238-243. doi: 10.1016/j.riam.2009. 03.005
use (Gibbens, 2021). Charnley AK, Collins SA (2007) Entomopathogenic fungi and their role
in pest control. In: Kubicek CP, Druzhinina IS (eds). The Mycota IV,
Environmental and Microbial Relationships. Berlin: Springer. Pp. 159-
188.
Acknowledgements Cifuentes-Triana YA (2016) Secuenciación y caracterización del genoma
de la cepa Saccharomyces cerevisiae 202-3 con potencial para la
The authors would like to thank Mauricio Diazgranados and Mark producción de etanol de segunda generación. Master thesis. Bogotá
Nesbitt at Royal Botanic Gardens, Kew, for their support and DC: Universidad Nacional de Colombia. 165p.
advice in categorising uses; and the reviewer Adriana Corrales for Cook F (1995) Economic Botany Data Collection Standard Prepared for
her insightful comments. This project was supported by a grant the International Working Group on Taxonomic Databases for Plant
under the Newton-Caldas Fund partnership. The grant is funded by Sciences (TDWG). Richmond: Royal Botanic Gardens, Kew. 146p.
Dávila GLR, Murillo AW, Zambrano FCJ, Suárez MH, Méndez AJJ (2020)
the UK Department for Business, Energy, and Industrial Strategy
Evaluation of nutritional values of wild mushrooms and spent
(BEIS) and the Colombian Ministry of Sciences, Technology, and substrate of Lentinus crinitus (L.) Fr. Heliyon, 6(3), 0-4. doi: 10.1016/j.
Innovation (Minciencias) and delivered by the British Council. heliyon.2020.e03502
Davis AK, Barrett FS, May DG, Cosimano MP, Sepeda ND, Johnson MW,
Finan PH, Griffiths RR (2020) Effects of Psilocybin-Assisted Therapy
References on Major Depressive Disorder: A Randomised Clinical Trial. JAMA
Psychiatry, 1-9. doi: 10.1001/jama psychiatry.2020.3285
Alves M, De Campos-Takaki G, Okada K, Ferreira Pessoa I, Milanez A Deak T (2009) Ecology and biodiversity of yeasts with potential
(2005) Detection of extracellular protease in Mucor species. Revista value in biotechnology. In: Satyanarayana T, Kunze G (eds). Yeast
Iberoamericana de Micologia, 22(2), 114-117. doi: 10.1016/S1130- Biotechnology: Diversity and Applications. Berlin: Springer. Pp. 151-
1406(05)70020-6 168.
Alves S, Pereira R, Lopes R, Tamai M (2002) Use of entomopathogenic de Mattos-Shipley KMJ, Ford KL, Alberti F, Banks AM, Bailey AM,
fungi in Latin America. In: Upadhyay R (ed). Advances in Microbial Foster GD (2016) The good, the bad and the tasty: the many roles
Control of Insect Pests. Boston: Kluwer Academic/Plenum Publishers. of mushrooms. Studies in Mycology, 85, 125-157. doi: 10.1016/j.
Pp. 193-211. simyco.2016.11.002
Aly AH, Debbab A, Kjer J, Proksch P (2010) Fungal endophytes from Demain AL, Martens E (2017) Production of valuable compounds by
higher plants: a prolific source of phytochemicals and other bioactive moulds and yeasts. Journal of Antibiotics, 70(4), 347-360. doi:
natural products. Fungal Diversity, 41, 1-16. doi: 10.1007/s13225- 10.1038/ja.2016.121
010-0034-4 Díaz-Escandón D, Soto-Medina E, Lücking R, Silverstone-Sopkin PA (2016)
Arboleda C, Mejía AI, Franco-Molano AE, Jiménez GA, Penninckx MJ (2008) Corticolous lichens as environmental indicators of natural sulphur
Autochthonous white rot fungi from the tropical forest of Colombia emissions near the sulphur mine El Vinagre (Cauca, Colombia). The
for dye decolourisation and ligninolytic enzymes production. Sydowia, Lichenologist, 48(2), 147159. doi: 10.1017/S0024282915000535
60(2), 165-180. Duarte-Trujillo AS, Jiménez-Forero JA, Pineda-Insuasti J, González-Trujillo
Arias-García G, Gutiérrez-Clavijo C, Ospina-Quintero CA (2008) Propuesta CA, García-Juárez M (2020) Extraction of bioactive substances from
del cultivo de hongos Pleurotus y Lentinula edodes a partir de la Pleurotus ostreatus (Pleurotaceae) by dynamic maceration. Acta
biomasa del café en las fincas cafeteras de Manizales para el Biologica Colombiana, 25(1), 61-74. doi:10.15446/abc.v25n1.72409.
fortalecimiento de los programas de desarrollo alternativo. Cuadernos Durán-Aranguren D, Chiriví-Salomón JS, Anaya L, Durán-Sequeda D, Cruz
Latinoamericanos de Administración, 4(6), 35-67. LJ, Serrano JD, Sarmiento L, Restrepo S, Sanjuan T, Sierra R (2020)
Arredondo-Peña CA, Pérez-Martínez AM (2007) Evaluación del crecimiento Effect of bioactive compounds extracted from Cordyceps nidus
del hongo Lentinula edodes Berk Pegler en residuos agroindustriales. ANDES-F1080 on laccase activity of Pleurotus ostreatus ANDES-F515.
Undergraduate thesis. Medellín: Universidad EAFIT. 132p. Biotechnology Reports, 26, 1-9. doi: 10.1016/j.btre.2020.e00466
Erjavec J, Kos J, Ravnikar M, Dreo T, Sabotič J (2012) Proteins of higher Suryanarayanan TS, Sujarit K, Suwannarach N, Suwunwong T, Thongbai
fungi - from forest to application. Trends in Biotechnology, 30(5), 259- B, Thongklang N, Wei D, Wijesinghe SN, Winiski J, Yan J, Yasanthika
273. doi: 10.1016/j.tibtech.2012.01.004 E, Stadler M (2019) The amazing potential of fungi: 50 ways we can
Espinel-Correal C, Torres Torres L, Villamizar-Rivero L, Bustillo- exploit fungi industrially. Fungal Diversity, 97(1), 1-136. doi: 10.1007/
Pardey A, Zuluaga-Mogollón MV, Cotes-Prado AM (2018) Hongos s13225-019-00430-9
entomopatógenos en el control biológico de insectos plaga. In: Cotes IGAC - Instituto Geográfico Agustín Codazzi (1997) Mapa de regiones
AM (ed). Control biológico de fitopatógenos, insectos y ácaros, Vol. 1. naturales de Colombia.
Mosquera: Corporación Colombiana de Investigación Agropecuaria - Jakubczyk D, Dussart F (2020) Selected fungal natural products with
Agrosavia. Pp. 334-367. antimicrobial properties. Molecules, 25(4), 1-18. doi: 10.3390/
Fernández-Uribe Y (2014) Cultivo de orellanas (Pleurotus ostreatus) en molecules25040911
cinco sustratos generados en los procesos productivos agropecuarios, Jaramillo-Ciro M, Botero-Botero R (2010) Comunidades liquénicas como
en dos épocas de siembra, en el municipio de Ituango. Undergraduate bioindicadores de calidad del aire. Gestión y Ambiente, 13(1), 97-110.
thesis. Medellín: Universidad Nacional Abierta y a Distancia de Jaramillo-Padilla SP, Silva-Benjumea JP, Osorio N (2004) Potencial
Colombia. 92p. simbiótico y efectividad de hongos micorrizo arbusculares de tres
Franco-Molano AE, Vasco-Palacios AM, López-Quintero CA, Boekhout T suelos sometidos a diferentes usos. Revista Facultad Nacional de
(2005) Macrohongos de la región del Medio Caquetá - Colombia: guía de Agronomía Medellín, 57(1), 2205-2217.
campo. Medellín: Universidad de Antioquia. 211p. Jaramillo-Ruiz D, Yepes L, Hincapié-Llanos G, Velasquez A, Vélez AL
Fungifert (2020) Ectomicorrizas hidrosolubles (inoculante ectomicorricico (2011) Desarrollo de productos a partir de la orellana (Pleurotus
para pinos y cipreses)- El semillero su aliado local. Available at: http:// ostreatus). Investigaciones Aplicadas, 5(2), 82-91.
elsemillero.net/enviar-pqr/ Kadri T, Rouissi T, Kaur Brar S, Cledon M, Sarma S, Verma M (2017)
Gámez-Guzmán A, Gaigl A, Torres-Rojas E (2019) Potential of a Cladosporium Biodegradation of polycyclic aromatic hydrocarbons (PAHs) by fungal
cladosporioides strain for the control of Tetranychus urticae Koch (Acari: enzymes: a review. Journal of Environmental Sciences, 51, 52-74. doi:
Tetranychidae) under laboratory conditions. Agronomía Colombiana, 10.1016/j.jes.2016.08.023
37(1), 21-26. doi: 10.15446/agron.colomb.v37n1.73353 López EL, Orduz S (2002) Metarhizium anisopliae y Trichoderma viride
García DAR, Arango MAR (2019) Estudios sobre la biorremediación en controlan colonias de Atta cephalotes en campo mejor que un
Colombia. Hechos Microbiológicos, 10(1), 39-48. doi: 10.17533/udea. insecticida químico. Revista Colombiana de Biotecnología, 4(1),
hm.v10n1a05 71-78.
García DA, Bustillo PAE, Castro VU, Arenas BY (2012) Selección de hongos López-Arboleda A, Ramírez-Castrillón M, Mambuscay-Mena LA, Osorio-
entomopatógenos para controlar salivazos (hemiptera: Cercopidae) de Cadavid E (2010) Diversidad de levaduras asociadas a chichas
la caña de azúcar en Colombia. Revista Colombiana de Entomologia, tradicionales de Colombia. Revista Colombiana de Biotecnología
38(2), 252-259. 12(2), 176-186.
Gaya E, Vasco-Palacios AM, Vargas-Estupiñán N, Lücking R, Carretero López-Rodríguez C, Hernández-Corredor R, Suárez-Franco C, Borrero M
J, Sanjuan T, Moncada B, Allkin B, Bolaños-Rojas AC, Castellanos- (2008) Evaluación del crecimiento y producción de Pleurotus ostreatus
Castro C, Coca LF, Corrales A, Cossu T, Davis L, dSouza J, Dufat A, sobre diferentes residuos agroindustriales del departamento de
Franco-Molano AE, García F, Gómez-Montoya N, González-Cuellar FE, Cundinamarca. Universitas Scientiarum, 13(2), 128-137.
Hammond D, Herrera A, Jaramillo-Ciro MM, Lasso-Benavides C, Mira Machado-Agudelo D (2019) Efecto in vitro de un bioformulado fúngico
MP, Morley J, Motato-Vásquez V, Niño-Fernández Y, Ortiz-Moreno ML, sobre el ácaro rojo de las aves Dermanyssus gallinae. Undergraduate
Peña-Cañón ER, Ramírez-Castrillón M, Rojas T, Ruff J, Simijaca D, thesis. Medellín: Universidad de Antioquia.
Sipman HJM, Soto-Medina E, Torres G, Torres-Andrade PA, Ulian T, Maldonado MC, Ibarra LV (2016) Colorantes orgánicos de hongos y
White K, Diazgranados M (2021) ColFungi: Colombian resources for líquenes. Scientia CUCBA, 8(2), 141-161.
Fungi Made Accessible. Richmond: Royal Botanic Gardens, Kew. 36p. Masi M, Nocera P, Reveglia P, Cimmino A, Evidente A (2018) Fungal
Gibbens S (2021) Fungi are key to our survival. Are we doing enough to metabolites antagonists towards plant pests and human pathogens:
protect them? National Geographic. Available at: https://www.national structure-activity relationship studies. Molecules, 23(4), 834. doi:
geographic.com/ environment/article/fungi-are-key-to-survival-are- 10.3390/molecules23040834
we-doing-enough-to-protect-them Mayorga OL, López E, Díaz TE, Barahona R (2005) Efecto de la
Giraldo, B. (2013). La manizaleña de los champiñones. Available at: https:// fuente de carbono y el tipo de inóculo sobre la producción de
www.lapatria.com/en-domingo/carmenza-jaramillo-la-manizalena-de- enzimas hidrolíticas del hongo anaeróbico ruminal Neocallimaxtis
los-champinones-45309. frontalis NFT 101. Revista CORPOICA, 6(1), 12-19. doi: 10.15797/
Grijalba EP, Espinel C, Cuartas PE, Chaparro ML, Villamizar LF (2018) concom.2019..23.009
Metarhizium rileyi biopesticide to control Spodoptera frugiperda: Miles LA, Lopera CA, González S, de García MCC, Franco AE, Restrepo S
stability and insecticidal activity under glasshouse conditions. Fungal (2012). Exploring the biocontrol potential of fungal endophytes from
Biology, 122(11), 1069-1076. doi: 10.1016/j.funbio. 2018.08.010 an Andean Colombian Paramo ecosystem. BioControl, 57(5), 697-
Guzmán G, Varela L (1978) Hongos de Colombia. III. Observaciones 710. doi: 10.1007/s10526 -012-9442-6
sobre los hongos, líquenes y mixomicetos de Colombia. Caldasia, Montoya S, Sanchez O, Levin L (2014) Evaluation of endoglucanase,
12, 309-338. exoglucanase, laccase, and lignin peroxidase activities on ten white-
Guzmán G, Torres MG, Ramírez-Guillén F, Ríos-Hurtado A (2004) rot fungi. Biotecnología en el Sector Agropecuario y Agroindustria,
Introducción al Conocimiento de los Macromicetos del Chocó, 12(2), 115-124.
Colombia. Revista Mexicana de Micología, 19, 33-43. Montoya-Barreto S, Restrepo GM, Tabares LA (2009) Rendimientos en
Holguín-Múnera JF, Esneider A, Oquendo E, Del Pilar R, Rodríguez M, Margely el cultivo de los hongos Pleurotus ostreatus y Pleurotus pulmonarius
G, Marín M (2017) Remoción de colorantes reactivos empleando el utilizando como sustratos diversos residuos agroindustriales.
hongo Bjerkandera adusta. Informador Técnico (Colombia), 81(2), 11- Revista de Investigaciones, 14, 16-26.
19. doi: 10.23850 /22565035.877 Morales LT, González-García LN, Orozco MC, Restrepo S, Vives MJ (2017)
Hyde KD, Xu J, Rapior S, Jeewon R, Lumyong S, Niego AGT, Abeywickrama The genomic study of an environmental isolate of Scedosporium
PD, Aluthmuhandiram JVS, Brahamanage RS, Brooks S, Chaiyasen apiospermum shows its metabolic potential to degrade hydrocarbons.
A, Chethana KWT, Chomnunti P, Chepkirui C, Chuankid B, de Silva Standards in Genomic Sciences, 12(1), 1-12. doi: 10.1186/s40793-
NI, Doilom M, Faulds C, Gentekaki E, Gopalan V, Kakumyan P, 017-0287-6
Harishchandra D, Hemachandran H, Hongsanan S, Karunarathna Moreno HP, Martínez AM, Fujimoto Y (2011) Aislamiento e identificación
A, Karunarathna S, Khan S, Kumla J, Jayawardena RS, Liu J, Liu de dos esteroles y un triterpenoide del cuerpo fructífero de
N, Luangharn T, Macabeo A, Marasinghe DS, Meeks D, Mortimer Ganoderma lucidum cultivado en Colombia. Vitae, 18(1), 11-15.
P, Mueller P, Nadir S, Nataraja K, Nontachaiyapoom S, O'Brien M, Mosquera-Espinosa AT, Bayman P, Otero JT (2010) Ceratobasidium como
Penkhrue W, Phukhamsakda C, Ramanan U, Rathnayaka AR, Sadaba hongo micorrízico de orquídeas en Colombia. Acta Agronomica, 59,
RB, Sandargo B, Samarakoon BC, Tennakoon DS, Siva R, Sriprom W, 316-326.
Mosquera-Espinosa AT, Bayman P, Prado GA, Gómez-Carabalí A, Otero JT Roy HE, Steinkraus DC, Eilenberg J, Hajek AE, Pell JK (2006) Bizarre
(2013) The double life of Ceratobasidium: orchid mycorrhizal fungi and interactions and endgames: entomopathogenic fungi and their
their potential for biocontrol of Rhizoctonia solani sheath blight of rice. arthropod hosts. Annual Review of Entomology, 51, 331-357. doi:
Mycologia, 105(1), 141-150. doi:10.3852/12-079 10.1146/annurev.ento.51.110104.15 0941
Moya-Álvarez L (2012) Determinación de la capacidad celulolítica de Ruan-Soto FC, Caballero-Nieto J, Cifuentes J, Garibay-Orijel R (2014)
consorcios de hongos provenientes de bosque Altoandino in vitro en Micofilia y micofobia: revisión de los conceptos, su reinterpretación e
tusas de palma de aceite. Master thesis. Nueva Granada: Universidad indicadores para su evaluación. In: Moreno-Fuentes A, Garibay-Orijel R
Militar Nueva Granada. 89p. (eds). La Etnomicología en México. Estado del Arte-Red de Etnoecología
Moya-Alvarez LA, Torres RE (2012) Hydrolysis of cellulose and oil palm y Patrimonio Biocultural (CONACyT). México DF: Universidad Autónoma
empty fruit bunches by using consortia of fungi isolated from the soil del Estado de Hidalgo. Pp. 17-31.
of Colombian high Andean Forest. Agronomía Colombiana, 30(3), 411- Rubiano LJ (1988) Líquenes como indicadores de contaminación en
418. el complejo industrial de Betania y la termoeléctrica de Zipaquirá,
Ordóñez CN, Tupac-Otero J, Díez GM (2012) Hongos endófitos de orquídeas Cundinamarca Líquenes. Acta Biológica Colombiana, 1(4), 95-125.
y su efecto sobre el crecimiento en Vanilla planifolia Andrews. Acta Ruíz A, Henao LG (2006) Hongos comestibles de Iguaque, Serie Especies
Agronómica, 61(3), 282-290. Colombianas (14). Bogotá: Instituto de Investigación de Recursos
Osorio-Cadavid E, Chaves-Lopez C, Tofalo R, Paparella A, Suzzi G (2008) Biológicos Alexander von Humboldt. 8p.
Detection and identification of wild yeasts in champús, a fermented Ruíz-Roa W, Henao-M LG, Peña-Cañón ER, Cipatoca LA, Cipamocha J
Colombian maise beverage. Food Microbiology, 25(6), 771-777. (2008) Reconocimiento y valoración de hongos silvestres comestibles
Palacio-Barrera AM, Areiza D, Zapata P, Atehortúa L, Correa C, Peñuela- de bosques de roble (Quercus humboldtii) en Paipa, Boyacá.
Vásquez M (2019) Induction of pigment production through media Undergraduate thesis. Tunja: Universidad Pedagógica y Tecnológica
composition, abiotic and biotic factors in two filamentous fungi. de Colombia.
Biotechnology Reports, 21, e00308. doi: 10.1016/j.btre.2019.e00308 Sanjuan T (1999) La diversidad del género Cordyceps en hormigas del
Peña-Cañón ER, Henao-Mejía LG (2014) Conocimiento y uso tradicional bosque húmedo tropical de Colombia. Undergraduate thesis. Bogotá
de hongos silvestres de las comunidades campesinas asociadas a DC: Universidad Nacional de Colombia.
bosques de roble (Quercus humboldtii) en la zona de influencia de la Shukla P, Upreti DK, Nayaka S, Tiwari P (2014) Natural dyes from
Laguna de Fúquene, Andes Nororientales. Etnobiología, 12(3), 28-40. Himalayan lichens. Indian Journal of Traditional Knowledge, 13(1),
Pérez E, Piragauta M (2006) Estudio etnomicológico entre los campesinos 195-201.
de los municipios de Arcabuco y Moniquirá, departamento de Boyacá. Simijaca-Salcedo DF, Morales-Puentes ME, Díaz-Pérez CN (2011)
Undergraduate thesis. Tunja: Universidad Pedagógica y Tecnológica de Líquenes y contaminación atmosférica en la Universidad Pedagógica
Colombia. y Tecnológica de Colombia, Tunja, Boyacá, Colombia. Revista Ciencia
Pérez UA, Ramírez MM, Serralde DP, Peñaranda AM, Wilches WA, Ramírez En Desarrollo, 3, 69-88.
L, Rengifo GA (2019) Hongos formadores de micorrizas arbusculares Smith A, Beltrán CA, Kusunoki M, Cotes AM, Motohashi K, Kondo T,
(HFMA) como estrategia para reducir la absorción de cadmio en Deguchi M (2013) Diversity of soil-dwelling Trichoderma in Colombia
plantas de cacao (Theobroma cacao). Revista Terra Latinoamericana, and their potential as biocontrol agents against the phytopathogenic
37(2), 121-131. doi: 10.28940/terra.v37i2.479 fungus Sclerotinia sclerotiorum (Lib.) de Bary. Journal of General Plant
Pérez-Higuera MV (2013) Estudio de la producción de lacasa por un Pathology, 79(1), 74-85. doi: 10.1007/s10327-012-0419-1
aislado nativo de Pleurotus ostreatus en cultivo sólido mediante el Suárez-Arango C (2010) Obtención in vitro de micelio de hongos
uso de cartón, cáscara de café y cascarilla de arroz bajo diversas comestibles, shiitake (Lentinula edodes) y orellanas (Pleurotus
concentraciones de cobre. Bogotá DC: Universidad de Los Andes. 21p. ostreatus y Pleurotus pulmonarius) a partir de aislamientos de cuerpos
Prescott T, Wong J, Panaretou B, Boa E, Bond A, Chowdhury S, Davies L, fructíferos, para la producción de semilla. Undergraduate thesis.
Østergaard L (2018) Useful Fungi. State of the World’s Fungi. Pp. 24-31. Bogotá DC: Universidad Nacional de Colombia. 86p.
doi: 10.1038/180936a0 Sun S, Zhang X, Chen W, Zhang L, Zhu H (2016) Production of natural edible
Primer Coloquio Colombiano de Micología, 2º día (2020) Asociación melanin by Auricularia auricula and its physicochemical properties.
Colombiana de Micología. Available at: https://www.youtube.com/ Food Chemistry, 196, 486-492. doi: 10.1016/j.foodchem.2015
watch?v=qzn4rhrEhQE .09.069
Puerta-Marín J, Zapata-Cardona PA (2006) Diseño conceptual de un Trujillo-Toro MA, Ramírez-Quirama JF (2012) Biorremediación en suelos
proceso para elaboración de un producto funcional a partir de contaminados con hidrocarburos en Colombia. Revista de Investigación
Ganoderma lucidum. Undergraduate thesis. Medellín: Universidad Agraria y Ambiental, 3(2), 37-62. doi: 10.22490/21456453.952
EAFIT. 115p. Usuga-Osorio CEU, Sánchez DAC, Molano AEF, Velásquez FAG, Agudelo
Ramírez-Castrillón M, Jaramillo-Garcia VP, Rosa PD, Landell MF, Vu D, CAL (2008) Efecto de la micorrización y la fertilización en la
Fabricio MF, Ayub MAZ, Robert V, Henriques JAP, Valente P (2017) acumulación de Biomasa en plantas de banano (Musa AAA cv. Gran
The oleaginous yeast Meyerozyma guilliermondii BI281A as a Enano) (Musaceae). Revista Facultad Nacional de Agronomía, 61(1),
new potential biodiesel feedstock: selection and lipid production 4269-4278.
optimisation. Frontiers in Microbiology, 8(1776) 1-11. doi: 10.3389/ Valadares R, Otero J, Pereira M, Cardoso E (2015) The epiphytic orchids
fmicb.2017.01776 Ionopsis utricularioides and Psygmorchis pusilla associate with different
Ramírez-Gil JG (2019) Dependencia, colonización y crecimiento en Ceratobasidium lineages at Valle del Cauca, Colombia. Acta Botanica
Gmelina arborea inoculada con cinco cepas de hongos micorrízicos Brasilica, 29(1), 40-44. doi: 10.1590/0102-33062014abb3397
arbusculares. Revista Facultad Nacional de Agronomía Medellín, 72(2), Vanegas-Córdoba I, Yepes-Pérez M, Ruiz-Villadiego O (2004) Producción de
8775-8783. doi: 10.15446 /rfnam.v72n2.74691 xilitol a partir de levaduras nativas colombianas. Revista Colombiana
Rodríguez N, Jaramillo C (2005) Cultivo de hongos medicinales en residuos de Biotecnología, 6(2), 31-36.
agrícolas de la zona cafetera. Boletín técnico Cenicafe. 73p. Vargas N, Gutiérrez C, Restrepo S, Velasco N (2019) Oyster mushrooms
Rodríguez-Valencia NR, & Jaramillo C (2004) Cultivo de hongos comestibles cultivation as an economic and nutritive alternative for rural low-
del género Pleurotus sobre residuos agrícolas de la zona cafetera. income women in Villapinzón-Colombia. In: Smith A (ed). Women in
Boletín técnico Cenicafe. 66p. Industrial and Systems Engineering: Key Advances and Perspectives on
Rodríguez-Valencia NR, Araque M, Perdomo F (2006) Producción de los Emerging Topics. Switzerland: Springer Nature. Pp. 561-587.
hongos comestibles Orellanas y Shiitake. Boletín técnico Cenicafe. Varón L (2004) Cultivo de hongos tropicales en residuos agroindustriales
33p. del departamento del Tolima. Undergraduate thesis. Tolima:
Romero O, Huerta M, Damián MA, Macías A, Tapia A, Parraguirre J, Juárez J Universidad del Tolima.
(2010) Evaluación de la capacidad productiva de Pleurotus ostreatus Vasco-Palacios AM (2002) Estudio etnobiológico de los hongos
con el uso de hoja de plátano (Musa paradisiaca l., CV. Roatan) macromicetes entre los Uitoto de la región de Araracuara (Amazonía
deshidratada, en relación con otros sustratos agrícolas. Agronomía Colombiana). Undergraduate thesis. Bogotá DC: Universidad Nacional
Costarricense, 34(1), 53-63. de Colombia.
A Colombian fungi producer in the region of Cundinamarca. The Agripina Kuecomuay of the indigenous Uitoto of Amazonas,
species, Lentinula edodes (Berk.) Pegler, commonly known as Colombia, showing her mushroom harvest.
shiitake mushroom, is an edible fungus grown in eucalyptus and [Tatiana Sanjuan]
acacia sawdust.
[Tatiana Sanjuan]
Panus neostrigosus
[Martha Ortiz-Moreno]
164 CATALOGUE OF FUNGI OF COLOMBIA
Chapter 13
Biotechnology of Fungi of Colombia
Martha L. Ortiz-Moreno1*, Rafael F. de Almeida2, Martha Cárdenas3, Marcela Guevara-Suarez4, Angelis Marbello3, Natalia Bolaños3, Pedro
Jiménez5 & Silvia Restrepo3
1
Grupo de Investigación en Sustentabilidad Ambiental (SUSA), Facultad de Ciencias Básicas e Ingenierías, Universidad de los Llanos.
2
Royal Botanical Gardens Kew, Ecosystem Stewardship, Sustainable Use, Seeds and Solutions, Richmond, TW9 3AE, Surrey, UK.
3
Laboratorio de Micología y Fitopatología (LAMFU), Departamento de ingeniería Química y de alimentos, Universidad de los Andes.
4
Applied Genomics Research Group, Vicerrectoría de Investigación y Creación, Universidad de los Andes.
5
Facultad de ciencias básicas y aplicadas, Universidad Militar Nueva Granada, Bogotá, Colombia.
ABSTRACT
The biotechnological use of fungi has been walking side by side with the development of human well-being since the dawn
of civilisation, enriching our diet with new products, increasing the assimilation of plant sources, and even contributing to
the conservation of protein sources. A qualitative leap in fungi manipulation was made possible with the development of
microbiology, mycology, and the use of molecular tools, laying the foundations for the development of fungal biotechnology.
These advances allowed the better identification of promising strains, understanding the relationship between genes and
phenotypes, and optimising their culture conditions to maximise the obtention of biomass or metabolites of interest to the
industry. In the global context, multiple studies have focused on the characterisation, bioprospecting, and development
of fungal biotechnological products, leading to patents and innovative, high-efficiency industrial processes with reduced
environmental impact. However, despite the great fungal diversity in Colombia, there is still much to investigate in order to
transform the dream of exploiting its funga as a source of biotechnological products that generate social well-being and allow
the positioning of technological innovation within the national productive sectors. Therefore, researchers, companies, and
consumers are urged to support the development of a Colombian fungal biotechnology strategy as an engine for sustainable
economic change.
RESUMEN
El uso de hongos ha acompañado el desarrollo del bienestar humano desde los albores de la civilización enriqueciendo la dieta
con nuevos productos, aumentando la asimilación de fuentes vegetales e inclusive contribuyendo a la conservación de las
fuentes de proteína. Pero es con el desarrollo de la microbiología, la micología y el uso de herramientas de biología molecular
que se dió un salto cualitativo en la manipulación de los hongos para sentar las bases del desarrollo de la biotecnología
fúngica, al lograr una mejor identificación de las cepas promisorias, comprender la relación entre los genes, los fenotipos
y cómo optimizar sus condiciones de cultivo para maximizar la obtención de biomasa o de metabolitos de interés para la
industria. En el contexto mundial son múltiples los estudios que se han enfocado en la bioprospección, caracterización y
desarrollo de productos biotecnológicos fúngicos conducentes al desarrollo de patentes e innovadores procesos industriales
de alta eficiencia y con impacto ambiental reducido. Sin embargo, en Colombia a pesar de su amplia diversidad fúngica aún
falta mucho por investigar para transformar en realidad el sueño de explotar la funga como fuente de productos biotecnológicos
que generen bienestar social y permitan posicionar la innovación tecnológica dentro de los sectores productivos nacionales.
Por ende, se insta a los investigadores, empresas y consumidores a apoyar el desarrollo de la biotecnología fúngica nacional
como un motor de cambio económico sustentable.
of environmental stresses, producing metabolites with alcohol for the pharmaceutical industry, and more recently,
possible applications in industry. Additionally, fungi have for the biofuel industry (Nandy & Srivastava, 2018). Starchy
rapid growth and the ability to adapt to the industrial and sugary substrates are used in these fermentation
production of biomass (Bekirogullari et al., 2020). They processes. Still, with the development of third-generation
can also be sources of genes of great interest that can be fuels, the possibility of saccharifying harvest waste or other
transferred to model organisms (Behera et al., 2017). Some substrates to incorporate enzymes of filamentous fungi in
of the most studied organisms in biotechnology are fungi fuel ethanol production have been explored (Nanda et al.,
due to their unique biological characteristics (Nevalainen 2018). Fungal strains are also used to produce citric, acetic,
et al., 2017). gluconic, gallic, kojic, and itaconic acids, fats, and glycerine,
Fungi are ubiquitous organisms that are adapted to among other chemical products (Herrera & Ulloa, 1998;
various ecosystems and habitats. Their enzymes participate Money, 2016). Dehydrated yeasts are also an important
in the decomposition of biomass, playing a fundamental role industrial product, used as nutritional supplements in human
in the development of cultivated plants, and collaborating and animal food, as they are rich in proteins and B vitamins
in the maintenance of soil fertility by decomposing complex (Shurson, 2018).
molecules (i.e., starch, cellulose, chitin, fats, and proteins) On the other hand, the maturation and fermentation
into simple elements that can be absorbed by plants and the of cheeses such as Roquefort, Camembert, and similar
soil microbiota (Herrera & Ulloa, 1998; Cepero et al., 2012). types depend on the metabolic activities of certain
Fungi participate in the humification and ammonisation moulds (Canellada et al., 2018). Likewise, the enzyme
phases within the nitrification process, transforming manufacturing industry has only flourished using various
proteins into amides, amines, amino acids, ammonia, and fungi to produce amylases, diastases, invertases, lipases,
ammonia salts (Herrera & Ulloa, 1998). Their role in soil proteases, pectinases, and other enzymes (Wösten, 2019).
fertility is especially important in acidic soils (Herrera & These enzymes are used to produce textiles, paper, leather,
Ulloa, 1998; Hoffland et al., 2004), such as those found in gums, syrups, and fruit juices (Wösten, 2019). The taka-
vast areas of Colombia (Fonte et al., 2019). Fungi are also diastase obtained from various Aspergillus species is used
important agents that participate in the ecological balance, for industrial and therapeutic purposes, as it consists of an
establishing symbiotic, mutualistic, synergistic, parasitic, enzyme complex of sucrase, maltase, lactase, dextrinase,
and even predatory interactions with other organisms amylase, cellulase, pectinase, lipase, peptase, ereptase,
(Herrera & Ulloa, 1998; Cepero et al., 2012). renin, trypsin, catalase, inulase, emulsin, tanases and
Currently, industrialised countries highlight the role of proteases, among others (Herrera & Ulloa, 1998; Kitagaki,
mycology in the human economy due to its great potential 2021).
in biotechnological applications derived from the metabolic Several antibiotics have been obtained from different fungi,
diversity of fungi (Meyer et al., 2020). With sufficient especially from Penicillium and Aspergillus, such as penicillin,
attention and resources, fungal biotechnology, in terms of produced by Penicillium notatum and P. chrysogenum (García-
the production of food and important bioactive products, Estrada et al., 2020). Likewise, an alkaloid called ergotine
including enzymatic, medicinal, and pharmacological ones, is extracted from ergot (Claviceps purpurea), and used in
can in the future contribute to the economy of developing medicine to combat bleeding associated with childbirth,
countries (Challa et al., 2019), such as Colombia. to induce uterine contractions (Liabsuetrakul et al., 2018),
and in the treatment of migraines due to its peripheral
USES OF FUNGI vasoconstrictive effect (Tfelt-Hansen & Koehler, 2008).
Fruiting bodies of certain Basidiomycota and Ascomycota These are some examples of the wide potential of fungi as
can be used as food by humans (Herrera & Ulloa, 1998; De a biotechnological resource. Thus, the goal of this chapter
Román & Boa, 2004). During certain seasons of the year, they was to review the state of knowledge of the biotechnological
constitute a food alternative for rural populations and even applications of the fungi of Colombia.
a source of income from the commercialisation of sporomes
as culinary delicacies (Christensen et al., 2008). Some of MATERIAL AND METHODS
these edible mushrooms, such as Agaricus bisporus, have To elucidate the biotechnological potential of the fungi
been cultivated for many years in various countries, giving of Colombia, we performed systematic searches on the
rise to an industrial branch of great economic importance reports of biotechnological applications of the genera or
(Carrasco et al., 2020). These mushrooms are sold fresh, species listed in the ColFungi portal during July and August
dried, pickled, or canned (Pardo-Giménez et al., 2017). 2021, with a temporal coverage of studies from between
However, this is not the only use of fungi as they are also used 1990 and 2021. Also, a brief review of studies conducted
in industrial processes to obtain many products, including in Colombia that have described endophyte fungi biology
food, beverages, and important drugs, with a huge impact was also performed. Most of these studies aimed to
on the economy of most countries (Pombeiro-Sponchiado et describe the diversity and, in some cases, tried to uncover
al., 2017; Singh, 2019). the potential of endophytic fungi as biocontrol agents or as
Since ancient times, yeasts have been used to produce producers of interesting metabolites. We also performed
bread, distilled beverages (i.e., beer and wine), and ethyl a search in Google Scholar, without time restriction, to
look for studies related to Colombian fungal diversity with alkaloids, terpenoids) to discourage the consumption of
the keyword’s modifiers “endophytes” AND “Colombia”, their biomass, develop pigments with antioxidant activity,
“Micromycetes diversity” AND “Colombia”; “Microfungal bioaccumulate metals, and produce other biomolecules with
diversity” AND “Colombia”; “Microfungi diversity” AND biological activities that are of great biotechnological interest
“Colombia”, excluding lichens, and the same terms for (Souza et al., 2015; Hyde et al., 2019). In this sense, it is a
Macromycetes. We also searched for metagenomics and priority to advance the knowledge of Colombian funga as it has
metabarcoding studies using the modifiers “Metagenomics” great potential to contribute to the sustainable development
AND “Microfungi” AND “Colombia”. of the country. From the data from ColFungi, we identified
287 plant symbiotrophs that have potential as promoters
RESULTS AND DISCUSSION of plant growth that could be used in forest production and
Biotechnological potential of fungi of Colombia annual and perennial agriculture. A total of 3,161 species
of saprotroph-symbiotrophs could be the source of genes,
The ecological role of fungi is very important as they enzymes, and biomolecules of great industrial importance, to
participate in nutrient cycling, playing a fundamental role in name a few example uses. To illustrate the biotechnological
diverse ecosystem services, especially in regulatory ones. potential of some of the genera recorded in Colombia, some
They can also act as predators and parasites that regulate examples will be presented below.
populations of other biological groups. On the basis of
the data from the ColFungi portal from the Royal Botanic Filamentous fungi: Trametes and Xylaria
Gardens, Kew, we identified eight main trophic modes
described for their species: pathogenic fungi feeding only One of the most-studied genera of lignin and cellulose-
on vascular plants (1,073), on animals (154 species), on degrading fungi is Trametes (Basidiomycota, Polyporales;
animals and vascular plants (2), only on bryophytes (1), on Figure 1). Several species (T. pubescens, T. elegans, T.
plant endophytes (2), other fungi excluding lichens (23), and versicolor, and T. villosa) are used for industrial applications
lichens (110). There are also saprotroph-pathotroph fungi, such as the degradation of agricultural waste (Osma et al.,
which can go from consuming organic matter to parasitising 2007; Gonzalez et al., 2013; Velásquez-Riaño et al., 2018),
other organisms under certain conditions. These are to biocatalyze the production of flavours (Mendez et al.,
represented by 176 species that can infect animals, other 2018; Jaramillo et al., 2020), and to bioremedy aromatic
fungi, bryophytes, and vascular plants. There is also a substances and wastewater (Osma et al., 2007; Ortiz-
group of species that can be consumers of organic matter, Monsalve et al., 2019; Mejía-Otálvaro et al., 2021). Degrading
pathogens, and mutualists depending on environmental activity on plant biomass has also been described for native
conditions. These are represented by 135 species, which isolates from Quindío corresponding to the genera Cookeina,
can be endophytes, epiphytes, and ectomycorrhizal. Xylaria (Ascomycota), and Earliella (Basidiomycota; Figure 1)
Another important functional group is the (Chaparro et al., 2009), showing that there are many genera
pathosymbiotrophs, made up of 119 species, mostly that have potential to degrade organic matter that are in
endophytes, although they can eventually parasitise other need of additional studies in Colombia.
organisms. In turn, saprotroph-symbiotrophs can go from
a free lifestyle to establishing symbiotic associations with Yeasts: Saccharomyces and Meyerozyma
different organisms. These are represented by 44 species Yeasts of the genus Saccharomyces (Ascomycota; Figure
that have been obtained from manure, litter, and roots. The 1) have been used in multiple applications as antioxidants
saprotrophs or consumers of organic matter are one of the and anti-topoisomerase I and II for the prevention of
most diverse groups with 1,430 species. They have been cardiovascular diseases and cancer (Correa Soto & Gaviria
obtained from manure, leaf litter and decomposing wood, Mendoza, 2010), in ethanol production from native strains
and the soil. (Garzón Castaño & Hernández Londoño, 2009; Peña-Serna
The symbiotrophs that grow in association with organisms et al., 2012; Walteros Pinzón, 2020), and in the processing
in the form of arbuscular mycorrhizae, ectomycorrhizae, of by-products such as glycerine (Sierra Arenas et al., 2015)
endophytes, epiphytes, and lichenised fungi represent 3,074 and surplus banana pulp (Oviedo Zumaqué et al., 2009).
species. Whereas the symbiotroph-saprotrophs (that is, fungi Likewise, S. cerevisiae has been used for lead bioadsorption
that predominantly live associated with other organisms but, (Pauro Roque et al., 2009) and animal supplementation
under certain conditions, can adopt free lifeforms through (Ramírez et al., 2007). Another widely studied yeast genus
the consumption of organic matter) were represented by 53 is Meyerozyma, which has been used as a probiotic in
species associated with wood (49) and lichens (4). ruminants (M. guilliermondii) (Chaparro et al., 2017; Castillo
Fungi have developed diverse mechanisms to persist in Saldarriaga et al., 2018) and to produce biodiesel (Ramírez-
different types of environments, such as the production of Castrillón et al., 2017). It has also been used for the
lytic enzymes adapted to different environmental conditions, fermentation of cocoa beans (Fonseca Blanco et al., 2020;
which can even degrade xenobiotics (chemical substances Lozano Tovar et al., 2020), and in plant protection against
not normally present in the environment of living organisms). anthracnose in fruits (M. caribbica) (Bautista Rosales et al.,
They can also release antibiotics to compete with other 2013; Ramírez-Castrillón et al., 2019; Navarro-Herrera &
microorganisms, synthesise complex molecules (e.g., Ortiz-Moreno, 2020).
Edible fungi: Auricularia, Favolus, Morchella, and Hericium because: 1. they have been found in every plant species
Edible mushrooms are of great interest to biotechnology examined to date, 2. researchers keep discovering new
due to their nutritional value and the possibility of offering species of fungi inhabiting plants, and 3. these fungi produce
the consumer biomolecules that can improve or maintain novel bioactive metabolites that can be used in agricultural
their health over time (Kües & Badalyan, 2017). Among the or industrial processes (Strobel & Daisy, 2003).
native fungi with great potential to be explored in Colombia Several problems with the taxonomy of endophytes make
are species of Auricularia, Favolus, Morchella, and Hericium their study and proper characterisation difficult (Miles et
(Figure 1). In Auricularia, Wu et al. (2014) have reported the al., 2012). First, many endophytic fungi do not sporulate
cultivation of A. auricula-judae, A. heimuer, A. americana, and under culture conditions. Thus, they are described as sterile
A. villosula as human food in China. These authors highlight mycelium or listed as morphospecies. In some cases,
that the consumption of these fungi dates back to Imperial molecular identification can help to clarify their taxonomy
China as a culinary delicacy reserved for emperors. By 2013, or at least helps to group some morphospecies. Second,
approximately 4.75 billion kg (fresh weight) of A. auricula- many represent new species to science or belong to novel
judae and other species known as “Heimuer” were grown, groups and do not fit within previously described taxa.
representing a 4 billion USD/year market. This high demand Third, most microorganisms do not grow in culture. Keeping
is due to its attributed anti-ageing properties, and the ability these problems in mind, we reviewed the diversity of these
of these foods to reduce sugar and fat in the blood, improve fungi in Colombia, focusing mainly on those that could be
immunity, and prevent and treat cancer (Wu et al., 2014). taxonomically identified.
In Colombia, Niño et al. (2017) analysed the cultivation Several authors, notably Elizabeth Arnold from Arizona
potential of Auricularia fuscosuccinea and Crepidotus University, have extensively studied fungal endophytes in the
tropics, resulting in the publication of more than 15 studies
palmarum, species with traditional use reported in rural
by her research group on the diversity of these organisms in
communities of Boyacá, obtaining positive results regarding
several tropical plants (Arnold et al., 2000), postulating that
their isolation and growth in substrates such as potato
tropical endophytes might be hyper-diverse. Although the
dextrose agar and malt extract agar. The consumption of
diversity of endophytes has been explored in several tropical
species of Auricularia, as well as Favolus, Lentinus, Lentinula,
countries and different plants, the diversity of native Colombian
Panus, and Pleurotus (Figure 1), has been reported in
microorganisms has not yet been explored extensively.
Amazonian indigenous communities. Thus, it is necessary
The first study of endophytes in Colombia was probably
to rediscover these traditional uses in order to find new food
performed by Dreyfuss & Petrini (1984). They isolated a
products for future world food sovereignty in a context where
few fungi from the plant families Araceae, Bromeliaceae,
access to quality protein and vitamins is increasingly limited
and Orchidaceae in Colombia, Brazil, and France. These
(Vargas-Isla et al., 2013). In addition, Murillo & Suárez
authors described several species of Ascomycota from
(2020) showed that the cultivation of Lentinus crinitus and
orchid species of Epidendrum and Maxillaria. One of the
its use on a laboratory scale is feasible. Meanwhile, Flórez-
most ubiquitous genera reported was Fusarium and its
Sampedro et al. (2016) found that Lentinus and Ganoderma
teleomorphs (Sordariomycetes). Orchids are essential
have anti-HIV activity.
plants in Colombia, encompassing an important commercial
On the other hand, Morchella (Ascomycota) is an edible
crop and the national flower (Cattleya trianae) (https://
fungus highly appreciated in European cuisine. This genus www.cancilleria.gov.co/simbolos-patrios-y-otros-datos-
has been reported in Colombia (Pinzón-Osorio et al., 2017; interes). The study of endophytes in orchid roots has
Baroni et al., 2018) to have great biotechnological potential had exciting potential as the limits between endophytes,
to be explored as it contains immunomodulatory proteins mycorrhizae, and parasites have been difficult to draw for
(Wu et al., 2020), as well as antioxidant and antimicrobial the microorganisms living in those tissues. For example, it
activities (Heleno et al., 2013). In turn, the genus Hericium has not been conclusively demonstrated that either partner
has also been recorded in Colombia (Murillo & Suárez, benefits from their interaction (Bayman & Otero, 2006).
2020), and shows a pleasant taste, high nutritional value, In Colombia, a significant number of altitudinal climatic
and hypoglycaemic activity. In addition, this genus has regions are likely to harbour many diverse microbial
cytoprotective properties for gastric cells (Liao & Huang, communities. According to Strobel & Daisy (2003), the
2019), and it is also neuroprotective (Brahmachari, 2017). number of plant species is so great that researchers looking
Hericium cultivation conditions have been evaluated by for microorganisms that produce novel bioactive products
Buchalo et al. (2005). Pre-treatment with Hericium has even should use creative and imaginative sampling strategies.
been proposed as a strategy to increase the digestibility of For example, one should look for endophytes in plants living
corn stover in ruminants (Moreno et al., 2019). in unique environmental sites that show different strategies
to survive extreme conditions (Strobel & Daisy, 2003).
Endophytic fungi: the hidden diversity Following Strobel’s rationale for plant selection (Strobel
Fungal endophytes colonise internal plant tissues without & Daisy, 2003), Miles et al. (2012) selected the endemic
causing any disease symptom, tissue damage, or apparent Andean giant rosettes (Espeletia complex: Asteraceae) from
damage in their hosts (Kogel et al., 2006; Schulz & Boyle, the paramo ecosystem to look for endophytes, describe
2005). These fungi have captured the interest of researchers their diversity and study their potential as biocontrol agents.
Their results showed that endophyte communities were P. brevicompactum. In this case, no species of Penicillium
dominated by organisms belonging to the Ascomycota, have ever been reported as pathogens of species of Coffea.
with Sordariomycetes predominating over other classes of The authors mentioned that this might imply that these
fungi. An exciting result of this study was that the taxonomic endophytes are not latent pathogens, suggesting that these
composition at the class level resembled that from tropical fungi can protect the plants against insects producing
forests more closely than that from temperate regions. This ochratoxin A (Vega et al., 2006). A more diverse group of
result was surprising as paramo conditions are more similar fungi was found in a posterior study using green coffee
to temperate locations than to tropical forests. Still, it might seeds (Vega et al., 2008). In this second study, fungi from
reflect the close geographical proximity of the paramos with the following genera and species were identified in Colombia:
Colombian tropical forests (pers. observ.). Gibberella sp., Aspergillus tubingiensis, and Penicillium
In Colombia, endophytes have also been studied in olsonii (Vega et al., 2008). In a third study, the same and
cultivated plants (e. g., coffee, Tahiti lime, tomatoes), mainly additional genera and species were discovered: Aspergillus
to find alternatives to control the causal agents of plant sp., A. fumigatus, A. oryzae, A. pseudodeflectus, Beauveria
diseases. In a first study, 15 morphospecies were isolated bassiana, Beauveria sp., Botryosphaeria sp., Cercospora
from several tissues (branches, leaves, and fruits) of the sp., Cladosporium sp., Clonostachys cf. rosea, Colletotrichum
Tahiti lime (Citrus x latifolia), all belonging to the Ascomycota, sp., Fusarium sp., Penicillium sp., Phomopis sp. Xylaria sp.,
except for two classified as sterile mycelium (Muñoz- among others (Vega et al., 2010).
Guerrero et al., 2021). Two isolates showing biocontrol The host specificity of endophytes is another wide-
activity were identified using the ITS and EF genetic markers open question. In Colombia, one study addressed this
as Xylaria adscendens and Trichoderma atroviride, again, two issue with the hypothesis that one endophyte from the
Sordariomycetes (Muñoz-Guerrero et al., 2021). plant family Melastomataceae, Chrysoporthe cubensis
The genus Fusarium is frequently found in endophyte (Cryphonectriaceae), had a change of host to infect species
studies (Kuldau & Yates, 2000). Isolating endophytes from of Eucalyptus (Myrtaceae) in Colombia. Six plant species
tomato roots, Andrade-Linares et al. (2011) obtained 51 from three genera of Melastomataceae were sampled,
fungal isolates with 20 isolates corresponding to Fusarium, with three species of the fungal family Cryphonectriaceae
eight representing sterile mycelia and seven with dark (Aurapex penicillata, C. cubensis, and C. inopina) being
septate hyphae. The dual nature of Fusarium as a pathogen identified using morphological and molecular techniques
and an endophyte is intriguing, and several hypotheses have (Granados et al., 2020). However, more studies are needed
been proposed in the literature. Many questions still need to determine whether host switches are common among
to be properly addressed by further studies: could there be endophyte species.
avirulent variants? Could they be living in a protected habitat Finally, a substantial amount of information on mycological
as long as the plant is healthy? Do they show different diversity is reported in documents that frequently are
stages of development? in university library repositories and are on either partly
Other studies have shown the recurrent isolation of available or not available at all to the public. These include
Fusarium from crops. Parsa et al. (2016) described endophytes honour theses (Montoya Sánchez, 2019), master's theses
in germinated seeds of the common bean, Phaseolus vulgaris (Hurtado Clopatosky, 2020), internal university journals
(Leguminosae). Although Aureobasidium pullulans was the (Melo & Suarez, 2016), and research project final reports.
dominant species, Fusarium oxysporum, Xylaria sp. and The species lists reported are similar to those in the
Cladosporium cladosporioides, among others, were also abovementioned papers, including frequent reports on
isolated (Parsa et al., 2016). Interestingly some of the most genera such as Aspergillus, Colletotrichum, Fusarium, and
important pathogens of the common bean were also found Trichoderma (Hurtado Clopatosky, 2020). Remarkably, many
as endophytes: Colletotrichum lindemuthianum, Fusarium of the reports were not directed to the study of biodiversity.
solani (currently Neocosmospora solani), and Macrophomina Instead, they were mainly focused on the search for practical
phaseolina (Parsa et al., 2016). A more recent study (Henao uses for endophytic fungi, such as the production of
et al., 2019) isolated 143 endophytes from leaves, roots, metabolites (Charria et al. 2021), plant growth stimulation
pseudostems, and corms obtained from healthy bananas (Ordóñez et al., 2012) or biological control of economic
and bananas affected by Fusarium oxysporum f. sp. cubense important fungal pathogens (Hurtado Clopatosky, 2020).
race 1. The results showed that Fusarium sp. was the most
common fungal morphotype in healthy (18.67%) and diseased Fungal culture collections
(25.53%) plants (Henao et al., 2019). Authors found 14 genera Research related to fungal bioprospection begins with
belonging to the Ascomycota, distributed in three classes: traditional studies on fungal biodiversity, including assertive
Sordariomycetes, Dothideomycetes, and Eurotiomycetes. polyphasic identification and the elaboration of a living
Coffee is probably the most important or known export (culture) fungi collection (Hyde et al., 2019). This approach
product from Colombia. Different tissues sampled from has discovered unique features of Micromycetes, showing
healthy plants belonging to Coffea arabica, C. congensis, the potential of these microorganisms in biotechnology
C. liberica, and C. dewevrei were used to isolate fungi. (Hyde et al., 2019). The cultures placed in the microbiological
Interestingly, the species identified among the isolates collections will then be used in applied research to exploit all
belonged to the genus Penicillium: P. oxalicum, P. brocae, and their industrial and biotechnological properties (Hyde et al.,
2019). One of Colombia’s most important microbiological Fitotripen (Trichoderma harzianum, T. koningii and T. viride),
culture collections is in the Pontificia Universidad Javeriana Mycorrhizagro (Glomus, Acaulospora, Scutellospora and
at Bogota [Member of the World Federation for Culture Entrophospora), and Safelomyces (Purpureocillium lilacinum
Collections (WFCC)]. However, only 130 fungal strains are and Cordyceps fumosorosea), used as bio controllers, plant
deposited in this collection, representing around 33 genera growth promoters, plant protectors and soil improvers
and 35 species, with no deposits of new fungal taxa (e.g., applicable to different crops. For other companies, such as
isotypes or holotypes). This fact is a worrying reality for Orius Biotech and Bio-Crop, with products oriented to the
future applied studies in Colombia. Therefore, strengthening agricultural sector based on fungi, it was not possible to
microbiological collections in countries like Colombia that obtain information on the origin of their strains.
have high biodiversity should be a national priority to
better represent fungal material collected and to support CONCLUSIONS
bioprospecting and bioeconomy studies based on fungi. The Colombian funga has an immense biotechnological
potential of which much remains unexplored, in addition
Fungal biotechnology in Colombian industry to building clusters of university research centres and
According to specialists consulted from the Asociación companie that allow the transfer of scientific knowledge
Colombiana de Micología (ASCOLMIC), there is a broad into commercial products, strengthening the Colombian
interest in innovation in Colombian industry, which is why productive sector, and generating a true transfer of social
fungal biotechnology processes have been used in the sugar- well-being with the use of fungal diversity.
alcohol, food, chemical, and agricultural products industries.
However, many of these bioprocesses are based on the use
of strains of fungi imported from other latitudes. This may Acknowledgements
be related to several factors: the political and legal difficulty
The authors are grateful for the support of their respective
of bringing a native fungal strain to the commercialisation institutions of affiliation: Universidad de los Llanos; Royal Botanic
phase as this implies costly and cumbersome bureaucratic Gardens, Kew; Universidad de los Andes and Universidad Militar
processes that universities, research centres, or business Nueva Granada. Also, the authors express their gratitude to the
consortia can hardly take on. There is also the difficulty in specialists of the Asociación Colombiana de Micología (ASCOLMIC):
legally accessing biological and genetic resources of fungi, Aida Vasco, Paola Zapata, Mauricio Ramírez, and Carolina
which implies a need to maintain reference collections and Chegwin. The authors also thank Ing. Nubia Moreno Sarmiento
to obtain permits for the study of fungi to commercialise from the company Biocultivos S.A. and the Microbiologist Alba
knowledge. At the same time, many resources are needed Lucia Calderón Jiménez from the company Natural Control S.A. for
to finance high-level research to characterise the native and supplying information on their fungal strains. We also thank the
promising strains of fungi of Colombia and to finance the reviewer Tom Prescott for his comments and suggestions.
processes of protecting intellectual property rights for the
innovations that are obtained, just as it is complex to involve
communities in order to transfer the benefits of biodiversity
References
to them in non-extractive exploitation schemes (Álvarez et Álvarez CF, Maldonado-Ocampo JA, Sanjuan T (2019) Evaluación nacional de
biodiversidad y servicios ecosistémicos. Bogota DC.: Instituto Alexander
al. 2019). von Humboldt. 330p.
The most successful experiences in national fungal Andrade-Linares DR, Grosch R, Franken P, Rexer KH, Kost G, Restrepo S,
biotechnology have been in agricultural applications, de Garcia MCC, Maximova E (2011) Colonisation of roots of cultivated
Solanum lycopersicum by dark septate and other ascomycetous
highlighting the development of products such as Trichotec endophytes. Mycologia, 103(4), 710-721.
from Agrosavia based on Trichoderma koningiopsis Th003 Arnold AE, Maynard Z, Gilbert GS, Coley PD, Kursar TA (2000) Are tropical
for the biocontrol of Fusarium oxysporum, Rhizoctonia solani, fungal endophytes hyperdiverse? Ecology Letters, 3(4), 267-274.
Sclerotinia sclerotium, S. minor and Botrytis cinerea, in Baroni TJ, Beug MW, Cantrell SA, Clements TA, Iturriaga T, Læssøe T, O’Donnell
K (2018) Four new species of Morchella from the Americas. Mycologia,
tomato, rice, lettuce and red fruit crops; this product also 110(6), 1205-1221. doi: 10.1080/00275514.2018.1533772
has a plant growth-promoting action. In addition to native Bautista-Rosales PU, Calderon-Santoyo M, Servín-Villegas R, Ochoa-
vesicular-arbuscular mycorrhizae included in their fertilisation Álvarez NA, Ragazzo-Sánchez JA (2013) Action mechanisms of the
yeast Meyerozyma caribbica for the control of the phytopathogen
and crop management protocols, Biocultivos S.A. produces Colletotrichum gloeosporioides in mangoes. Biological Control, 65(3),
Fosfobiol based on Penicillium janthinellum, which acts as a 293-301. doi: 10.1016/j.biocontrol.2013.03.010
phosphorus solubiliser for cotton, rice, coffee, sugar cane, Bayman P, Otero JT (2006) Microbial endophytes of orchid roots, Microbial
root endophytes. Springer. Pp. 153-177.
corn, pastures and soybeans; Trifesol from Trichoderma viride, Behera BC, Sethi BK, Mishra RR, Dutta SK, Thatoi HN (2017) Microbial
which can be used as a biocontrol agent (mycoparasite), cellulases – diversity and biotechnology with reference to mangrove
plant growth promoter, and cellulose degrader; and a residue environment: a review. Journal of Genetic Engineering and Biotechnology,
15(1), 197-210. doi: 10.1016/j.jgeb.2016.12.001
treatment based on the Penicillium pinophillum and Pleurotus Bekirogullari M, Figueroa-Torres GM, Pittman JK, Theodoropoulos C
ostreatus consortium, as cellulose and lignin degraders (2020) Models of microalgal cultivation for added-value products
for the management of plant biomass residues. From the - a review. Biotechnology Advances, 44, 107609. doi: 10.1016/j.
biotechadv.2020.107 609
Natural Control company, the products based on native
Brahmachari G (2017) Discovery and development of neuroprotective
strains of fungi stand out: Anisagro (Metarhizium anisoplae), agents from natural products. Elsevier. 465p. doi: 10.1016/B978-0-12-
Vercani (Lecanicillium lecanii), Bassar (Beauveria bassiana), 809593-5.00001-X
Buchalo AS, Didukh MY, Mykchaylova OB, Lynovitska VM (2005) and semi-solid culture conditions on agro-industrial wastes. PLoS ONE,
Microstructures in medicinal mushroom cultures. International Journal of 8(9), e73721. doi: 10.1371/journal.pone.0073721
Medicinal Mushrooms, 7(3), 389-390. doi: 10.1615/IntJMedMushrooms. Granados GM, McTaggart AR, Rodas CA, Roux J, Wingfield MJ (2020)
v7.i3.380 Species of Cryphonectriaceae occupy an endophytic niche in the
Canellada F, Laca A, Laca A, Díaz M (2018) Environmental impact of cheese Melastomataceae and are putative latent pathogens of Eucalyptus.
production: a case study of a small-scale factory in southern Europe European Journal of Plant Pathology, 156(1), 273-283.
and global overview of carbon footprint. Science of the Total Environment, Heleno SA, Stojković D, Barros L, Glamočlija J, Soković M, Martins A,
635, 167-177. doi: 10.1016/j.scitotenv.2018.04.045 Ferreira IC (2013) Chemical composition and bioactivity of wild samples
Carrasco J, García‐Delgado C, Lavega R, Tello ML, De Toro M, Barba‐Vicente of Morchella esculenta from Portugal and Serbia. In: The 7th International
V, Preston GM (2020) Holistic assessment of the microbiome dynamics Workshop on Edible Mycorrhizal Mushrooms. Guatemala, La Antigua.
in the substrates used for commercial champignon (Agaricus bisporus) Available at: http://hdl.handle.net/10198/8690
cultivation. Microbial Biotechnology, 13(6), 1933-1947. Henao SZ, Vasquez M, Henao IC, Hoyos LFP, Torres JDS, Hoyos-Carvajal LM
Castillo-Saldarriaga C, Céspedes E, Gómez-Álvarez M (2018) Evaluation of (2019) Fungal endophytes in bananas cv Manzano affected by Fusarium.
the stability of granulated prototypes based on a Colombian yeast strain of African Journal of Agricultural Research, 14(7), 430-438.
Meyerozyma guilliermondii and oregano essential oil. Scientific Production Herrera T, Ulloa M (1998) El reino de los hongos: micología básica y aplicada.
Poster. Corporación Colombiana de Investigación Agropecuaria. Available Universidad Nacional Autónoma de México. Mexico, Mexico City. 552p.
at: https://bit.ly/3CYbRpn Hoffland E, Kuyper TW, Wallander H, Plassard C, Gorbushina AA,
Cepero de García MC, Restrepo S, Franco-Molano, A, Cárdenas M, Vargas N Haselwandter K, van Breemen N (2004) The role of fungi in weathering.
(2012) Biología de hongos. Bogota DC.: Ediciones Uniandes-Universidad Frontiers in Ecology and the Environment, 2(5), 258-264.
de los Andes. 497p. Hurtado Clopatosky S (2020) Aislamiento de endófitos en gulupa (Passiflora
Challa S, Dutta T, Neelapu NRR (2019) Fungal white biotechnology applications edulis Sims f.) y su potencial para promoción de crecimiento de la planta
for food security: opportunities and challenges. In: Yadav A, Singh S, y control del Fitopatógeno Fusarium oxysporum. Universidad Nacional de
Mishra S, Gupta A (eds). Recent Advancement in White Biotechnology Colombia. Colombia, Bogota DC. 168p.
Through Fungi. Springer. doi: 10.1007/978-3-030-14846-1_4 Hyde KD, Xu J, Rapior S, Jeewon R, Lumyong S, Niego AGT, Stadler M (2019)
Chaparro ML, Céspedes E, Cruz M, Castillo-Saldarriaga CR, Gómez-Álvarez The amazing potential of fungi: 50 ways we can exploit fungi industrially.
MI (2017) Fluidized bed drying of a granulated prototype based on a Fungal Diversity, 97(1), 1-136. doi: 10.1007/s13225-019-00430-9
potential probiotic yeast Meyerozyma guilliermondii: selection of process Jaramillo DA, Méndez MJ, Vargas G, Stashenko EE, Vasco-Palacios
parameters and drying protectant. Revista Mexicana de Ingeniería AM, Ceballos A, Caicedo NH (2020) Biocatalytic potential of native
Química, 16(2), 347-357. basidiomycetes from Colombia for flavour/aroma production. Molecules,
Chaparro DF, Rosas DC, Varela A (2009) Aislamiento y evaluación de la 25(18), 4344. doi: 10.3390/molecules25184344
actividad enzimática de hongos descomponedores de madera (Quindío, Khan FF, Ahmad K, Ahmed A, Haider S (2017) Applications of biotechnology
Colombia). Revista Iberoamericana de Micología, 26(4), 238-243. doi: in agriculture - review article. World Journal of Biology and Biotechnology,
10.1016/j.riam.2009.03.005 2(1), 139-142. doi: 10.33865/wjb.002.01.0013
Charria-Girón E, Espinosa MC, Zapata-Montoya A, Méndez MJ, Caicedo JP, Kitagaki H (2021) Medical application of substances derived from non-
Dávalos AF, Ferro BE, Vasco-Palacios AM, Caicedo NH (2021) Evaluation pathogenic fungi Aspergillus oryzae and A. luchuensis-containing Koji.
of the antibacterial activity of crude extracts obtained from cultivation Journal of Fungi, 7(4), 243.
of native endophytic fungi belonging to a tropical montane rainforest in Kogel KH, Franken P, Hückelhoven R (2006) Endophyte or parasite–what
Colombia. Frontiers in Microbiology, 2515. decides? Current Opinion in Plant Biology, 9(4), 358-363.
Christensen M, Bhattarai S, Devkota S, Larsen HO (2008) Collection and use Kües U, Badalyan SM (2017) Making use of genomic information to explore
of wild edible fungi in Nepal. Economic Botany, 62(1), 12-23. the biotechnological potential of medicinal mushrooms. In: Medicinal
Correa Soto CE, Gaviria Mendoza A (2010) Evaluación de la capacidad Plants and Fungi: Recent Advances in Research and Development.
antioxidante y actividad antitopoisomerasa I y/o II sobre cepas mutadas Singapore: Springer. Pp. 397-458. doi: 10.1007/978-981-10-5978-
de Saccharomyces cerevisiae de plantas de la ecorregión cafetera 0_13
colombiana. Undergraduation Thesis. Universidad Tecnologica de Kuldau GA, Yates IE (2000) Evidence for Fusarium endophytes in cultivated
Pereira. Colombia, Risaralda. Available at: http://repositorioslatinoameri and wild plants. In: Microbial Endophytes (CW Bacon, JF White, eds):
canos.uchile.cl/handle/2250/78852 85–120. Marcel Dekker, New York and Basel.
De Román M, Boa E (2004) Collection, marketing, and cultivation of edible Liabsuetrakul T, Choobun T, Peeyananjarassri K, Islam QM (2018)
fungi in Spain. Micologia Aplicada International, 16(2), 25-33. Prophylactic use of ergot alkaloids in the third stage of labour. Cochrane
Dreyfuss M, Petrini O (1984) Further investigations on the occurrence and Database of Systematic Reviews, 6(6), CD005456. doi: 10.1002/1465
distribution of endophytic fungi in tropical plants. Botanica Helvetica, 1858.CD005456.pub3
94(1), 33-40. Liao B, Huang H (2019) Structural characterisation of a novel polysaccharide
Flórez-Sampedro L, Zapata W, Orozco LP, Mejía AI, Arboleda C, Rugeles MT from Hericium erinaceus and its protective effects against H2O2-induced
(2016) In vitro anti-HIV-1 activity of the enzymatic extract enriched with injury in human gastric epithelium cells. Journal of Functional Foods, 56,
laccase produced by the fungi Ganoderma sp. and Lentinus sp. Vitae, 265-275. doi: 10.1016/j.jff.2019.03.028
23(2), 109-118. doi: 10.17533/udea.vitae.v23n2a03 Lozano Tovar MD, Tibasosa G, González CM, Alvarez KB, Hernandez MDPL,
Fonseca Blanco JDF, Hernandez MDPL, Galeano LSO, Nuñez JC, Tovar Villamizar FR (2020) Isolation and identification of microbial species
MDL (2020) Effect of addition of a specific mixture of yeast, lactic found in cocoa fermentation as microbial starter culture candidates
and acetic bacteria in the fermentation process to improve the quality for cocoa bean fermentation in Colombia. Pelita Perkebunan (a Coffee
and flavour of cocoa beans in Colombia. Pelita Perkebunan (a Coffee and Cocoa Research Journal), 36(3), 236-248. doi: 10.22302/iccri.jur.
and Cocoa Research Journal), 36(2), 154-172. doi: 10.22302/iccri.jur. pelitaperkebu nan.v36i3.443
pelitaperkebunan.v36i2. 438 Mejía-Otálvaro F, Merino-Restrepo A, Hormaza-Anaguano A (2021) Evaluation
Fonte SJ, Botero C, Quintero DC, Lavelle P, van Kessel C (2019) Earthworms of a Trametes pubescens laccase concentrated extract on allura red AC
regulate plant productivity and the efficacy of soil fertility amendments decolourisation without the addition of synthetic mediators. Journal
in acid soils of the Colombian Llanos. Soil Biology and Biochemistry, 129, of Environmental Management, 285, 112117. doi: 10.1016/j.jenv
136-143. doi: 10.1016/j.soilbio.2018.11.016 man.2021.112117
García-Estrada C, Martín JF, Cueto L, Barreiro C (2020) Omics approaches Melo JSR, Suarez FML (2016) Micobiota endófita del género Maxillaria Ruiz
applied to Penicillium chrysogenum and penicillin production: revealing & Pav. (Orchidaceae) en dos reservas naturales del departamento del
the secrets of improved productivity. Genes, 11(6), 712. Quindio. Revista de Investigaciones Universidad del Quindío, 28(1), 23-31.
Garzón Castaño SC, Hernández Londoño C (2009) Estudio comparativo Mendez MJ, Caicedo NH, Salamanca C (2018) Trametes elegans: a fungal
para la producción de etanol entre Saccharomyces cerevisiae silvestre, endophytic isolate from Otoba gracilipes as biocatalyst for natural
Saccharomyces cerevisiae ATCC 9763 y Candida utilis ATCC 9950. flavours production. New Biotech, 44, 1.
Undergraduation Thesis. Universidad Tecnologica de Pereira. Colombia, Meyer V, Basenko EY, Benz JP, Braus GH, Caddick MX, Csukai M, Wösten
Risaralda. Available at: https://repositorioslatinoamericanos.uchile.cl/ HA (2020). Growing a circular economy with fungal biotechnology: a
handle/2250/ 78502 white paper. Fungal Biology and Biotechnology, 7(1), 1-23. doi: 10.1186/
Gonzalez JC, Medina SC, Rodriguez A, Osma JF, Alméciga-Díaz CJ, Sánchez s40694-020-00095-z
OF (2013) Production of Trametes pubescens laccase under submerged Miles L, Lopera C, González S, de García MC, Franco A, Restrepo S (2012)
Exploring the biocontrol potential of fungal endophytes from an Andean applications. Melanin. Miroslav Blumenberg, IntechOpen. doi:
Colombian Paramo ecosystem, BioControl, 57(5), 697-710. 10.5772/67375
Money NP (2016) Fungi and biotechnology. In: The fungi. Waltham: Academic Ramírez JF, Medina S, Garcê N, Cifuentes T (2007) Effects of the
Press. 449p. doi: 10.1016/B978-0-12-382034-1.00012-8 supplementation with yeast (Saccharomyces cerevisiae) on milk yield,
Montoya Sánchez SH (2019) Evaluación in vitro de la capacidad antagónica and milk components of water buffalo cows from northeast of Colombia.
de hongos endófitos aislados de Moringa oleífera (l) en el control Italian Journal of Animal Science, 6(sup2), 502-503. doi: 10.4081/ijas.
de Pestalotia palmarum (Q). Universidad de Santander. Colombia, 2007.s2.5 02
Bucaramanga. 79p. Ramírez-Castrillón, M, Jaramillo-Garcia VP, Rosa PD, Landell MF, Vu D,
Moreno PL, Leal-Lara H, García-Pérez Á, Corona L, Romero-Pérez A, Fabricio MF, Valente P (2017) The oleaginous yeast Meyerozyma
Márquez-Mota CC (2019) PSXIV-14 evaluation of 21 fungal strains guilliermondii BI281A as a new potential biodiesel feedstock: selection
as pretreatment of corn stover: chemical composition and in vitro and lipid production optimisation. Frontiers in Microbiology, 8, 1776. doi:
digestibility. Journal of Animal Science, 97(Supplement_3), 444. doi: 10.3389/fmicb.2017.01776
10.1093/jas/ skz258.874 Ramírez-Castrillón M, Usman LM, Silva-Bedoya LM, Osorio-Cadavid E
Muñoz-Guerrero J, Guerra-Sierra BE, Alvarez JC (2021) Fungal endophytes (2019) Dominant yeasts associated to mango (Mangifera indica) and
of Tahiti Lime (Citrus citrus × latifolia) and their potential for control of rose apple (Syzygium malaccense) fruit pulps investigated by culture-
Colletotrichum acutatum JH Simmonds causing anthracnose. Frontiers based methods. Anais da Academia Brasileira de Ciências, 91 (04),
in Bioengineering and Biotechnology, 9, 255. e20190052. doi: 10.1590/0001-3765201920190052
Murillo W, Suárez H (2020) Evaluation of nutritional values of wild Schulz B, Boyle C (2005) The endophytic continuum. Mycological Research,
mushrooms and spent substrate of Lentinus crinitus (L.) Fr. Heliyon, 109(6), 661-686.
6(3), e03502. doi: 10.1016/j.heliyon.2020.e03502 Shurson GC (2018) Yeast and yeast derivatives in feed additives and
Nanda S, Rana R, Sarangi PK, Dalai AK, Kozinski JA (2018) A broad ingredients: sources, characteristics, animal responses, and
introduction to first-, second-, and third-generation biofuels. In: Recent quantification methods. Animal Feed Science and Technology, 235, 60-76.
advancements in biofuels and bioenergy utilisation. Singapore: Springer. Sierra Arenas YK, Díaz Castañeda CE, Hernández Mora JA (2015)
402p. doi: 10.1007/978-981-13-1307-3_1 Determinación del porcentaje de etanol producido por Saccharomyces
Nandy SK, Srivastava RK (2018) A review on sustainable yeast cerevisiae a partir de glicerina semipurificada. Semana de Divulgación
biotechnological processes and applications. Microbiological Research, Científica Institucional UDES, campus Cúcuta, Colombia. Available
207, 83-90. doi: 10.1016/j.micres.2017.11.013 at: http://service.udes.edu.co/semanadivulgacion/segundaSemana/
Navarro-Herrera YY, Ortíz-Moreno ML (2020) Yeast strains with memorias/ponencias/p12.pdf
antagonist activity against Colletotrichum gloeosporioides (Penz.) Singh B (2019) Production, characteristics, and biotechnological applications
Penz. & Sacc. and their phenotypic characterisation. Egyptian of microbial xylanases. Applied Microbiology and Biotechnology, 103(21),
Journal of Biological Pest Control, 30(1), 1-8. doi: 10.1186/s41938- 8763-8784. doi: 10.1007/s00253-019-10108-6
020-00231-2 Souza PMD, Bittencourt MLDA, Caprara CC, Freitas MD, Almeida RPCD,
Nevalainen H (2017) Editorial current advances in fungal biotechnology (part Silveira D, Magalhães PO (2015) A biotechnology perspective of fungal
I). Current Biotechnology, 6(2), 77. doi: 10.2174/22115501060217042 proteases. Brazilian Journal of Microbiology, 46, 337-346. doi: 10.1590/
4190046 S1517-838246220140359
Niño YM, Peña ER, Henao LG (2017) Aislamiento y producción de semilla Strobel G, Daisy B (2003) Bioprospecting for microbial endophytes and
de Auricularia fuscosuccinea (Mont.) Henn. y Crepidotus palmarum their natural products. Microbiology and Molecular Biology Reviews,
Sing. usados tradicionalmente en Pauna (Boyacá, Colombia). Revista 67(4), 491-502.
Colombiana de Ciencias Hortícolas, 11(1), 151-158. doi: 10.17584/ Tfelt-Hansen PC, Koehler PJ (2008) History of the use of ergotamine and
rcch. 2017v11i1.5616 dihydroergotamine in migraine from 1906 and onward. Cephalalgia,
Ordóñez NF, Tupac Otero J, Díez MC (2012) Hongos endófitos de orquídeas 28(8), 877-886. doi: 10.1111/j.1468-2982.2008.01578.x
y su efecto sobre el crecimiento en Vanilla planifolia Andrews. Acta United Nations (1992) The Convention on Biological Diversity. Quebec:
Agronómica, 61(3), 282-290. United Nations.
Ortiz-Monsalve S, Valente P, Poll E, Jaramillo-García V, Henriques JAP, Vargas-Isla R, Ishikawa NK, Py-Daniel V (2013) Contribuições etnomicológicas
Gutterres M (2019) Biodecolourization and biodetoxification of dye- dos povos indígenas da Amazônia. Biota Amazônia, 3(1), 58-65. doi:
containing wastewaters from leather dyeing by the native fungal strain 10.18561/2179-5746/biotaamazonia.v3n1p58-65
Trametes villosa SCS-10. Biochemical Engineering Journal, 141, 19-28. Velásquez-Riaño M, Carvajal-Arias CE, Rojas-Prieto NL, Ausecha-García
doi: 10.1016/j.bej.2018.10.002 SA, Vera-Díaz MÁ, Meneses-Sánchez JS, Villa-Restrepo AF (2018)
Osma J, Osma JF, Rodriguez Couto S, Toca Herrera JL (2007) Poly-R-478 Evaluation of a mixed simultaneous vinasse degradation treatment
and ABTS oxidation by the white-rot fungus Trametes pubescens on using Komagataeibacter kakiaceti GM5 and Trametes versicolor DSM
agar plates. Journal of Biotechnology, 131(1), S229. doi: 10.1016/j. 3086. Ecotoxicology and Environmental Safety, 164, 425-433. doi:
jbiotec. 2007.07.416 10.1016/j.ecoenv.2018.08.037
Oviedo Zumaqué L, Mantilla CL, Pantoja MM (2009) Levaduras autóctonas Vega FE, Posada F, Aime MC, Peterson SW, Rehner SA (2008) Fungal
con capacidad fermentativa en la producción de etanol a partir de pulpa endophytes in green coffee seeds. Mycosystema, 27(1), 75-84.
de excedentes de plátano Musa (AAB Simmonds) en el departamento de Vega FE, Posada F, Peterson SW, Gianfagna TJ, Chaves F (2006) Penicillium
Córdoba, Colombia. Revista Colombiana de Biotecnología, 11(1), 40-47. species endophytic in coffee plants and ochratoxin A production.
Pardo-Giménez A, Pardo JE, Zied DC (2017) Casing materials and Mycologia, 98(1), 31-42.
techniques in Agaricus bisporus cultivation. In: Cunha ZD, Pardo- Vega FE, Simpkins A, Aime MC, Posada F, Peterson SW, Rehner SA, Infante
Giménez A (eds). Edible and medicinal mushrooms: technology and F, Castillo A, Arnold AE (2010) Fungal endophyte diversity in coffee
applications. Pp. 385-413. plants from Colombia, Hawaii, Mexico and Puerto Rico. Fungal Ecology,
Parsa S, García-Lemos AM, Castillo K, Ortiz V, López-Lavalle LAB, Braun 3(3), 122-138.
J, Vega FE (2016) Fungal endophytes in germinated seeds of the Walteros Pinzón CD (2020) Caracterización de cepas de levadura colombiana
common bean, Phaseolus vulgaris. Fungal Biology, 120(5), 783-790. Saccharomyces cerevisiae para su potencial uso en la producción de
Pauro Roque JJ, Yucra MC, Aguilar RP, Canqui AM (2009) Estudios de cerveza “Colombian Ale”. Undergraduation thesis. Universidad de los
bioadsorción de plomo por Saccharomyces cereviceae en soluciones Andes. Colombia, Bogota DC. 56p.
acuosas. Revista Colombiana de biotecnología, 11(1), 33-39. Wösten HA (2019) Filamentous fungi for the production of enzymes,
Peña-Serna C, Castro-Gil C, Peláez-Jaramillo CA (2012) Evaluation of chemicals and materials. Current Opinion in Biotechnology, 59, 65-70.
ethanol production from two recombinant and a commercial strains doi: 10.1016/j.copbio.2019.02.010
of Saccharomyces cerevisiae (Fungi: Ascomycota) in sugar-cane Wu F, Yuan Y, Malysheva VF, Du P, Dai YC (2014) Species clarification of
molasses and rejected-banana juice from Uraba (Antioquia), Colombia. the most important and cultivated Auricularia mushroom “Heimuer”:
Actualidades Biológicas, 34(96), 21-31. evidence from morphological and molecular data. Phytotaxa, 186(5),
Pinzón-Osorio CA, Pinzón-Osorio J (2017) Primer registro de la familia 241-253. doi: 10.11646/phytotaxa.186.5.1
Morchellaceae (Ascomycota: Pezizales) para Colombia. Revista Wu G, Sun Y, Deng T, Song L, Li P, Zeng H, Tang X (2020) Identification
Peruana de Biología, 24(1), 05-110. doi: 10.15381/rpb.v24i1.12481 and functional characterisation of a novel immunomodulatory protein
Pombeiro-Sponchiado SR, Sousa GS, Andrade JC, Lisboa HF, Gonçalves RC from Morchella conica SH. Frontiers in Immunology, 11, 559770. doi:
(2017) Production of melanin pigment by fungi and its biotechnological 10.3389/fimmu.2020.559770
Sticta peltigerella
[Robert Lücking]
174 CATALOGUE OF FUNGI OF COLOMBIA
Chapter 14
Fungal Conservation in Colombia
Diego Simijaca 1,2,3*, Gregory Mueller 4,5 & Aída M. Vasco-Palacios2,5,6
1
Universidad Autónoma de Aguascalientes, Facultad de Ciencias Básicas, Centro de Ciencias Biológicas, Avenidad Universidad, # 940, Aguascalientes
20131, México.
2
Asociación Colombiana de Micología, ASCOLMIC.
3
Grupo Colombiano de Liquenología, GCOL.
4
Negaunee Institute for Plant Conservation Science and Action, Chicago Botanic Garden, Illinois, USA.
5
IUCN SSC Fungal Conservation Committee, FunCC.
6
Grupo de Microbiología Ambiental y Grupo BioMicro, Escuela de Microbiología, Universidad de Antioquia UdeA, Calle 70 No. 52–21, Medellín 050010,
Antioquia, Colombia.
ABSTRACT
There has been increasing activity in fungal conservation in Colombia over the past five years, leading to a higher profile for fungi
in conservation efforts. After reviewing the country’s fungal conservation status, further development and implementation of
measures to conserve fungi are still needed in Colombia. We attribute this to the fact that fungi are not explicitly included as an
element of the biological diversity in Colombia. Additionally, the generally low profile of fungi among conservation stakeholders
and decision-makers and the limited efforts by fungal scientists to put mycological knowledge into a conservation context has
hindered efforts. Colombia needs to focus efforts to advance the knowledge of the fungal diversity of its national territory,
including taxonomy, distribution, ecology, and threats, to enable fungi to be included within national conservation agendas.
The status of fungal knowledge in Colombia reflects the global situation, with only 7,241 of the estimated 300,000 species
documented and registered according to ColFungi. As a result, the extinction risk of South American fungal species is only
now beginning to be assessed. Colombian mycologists have been active in this field and are among the leaders in fungal
conservation efforts. Yet only 22 of the 7,241 species of fungi recorded from Colombia have had their global extinction risk
assessed and published by the IUCN Red List. This panorama highlights the urgency with which the fungi of Colombia should
be assessed in conservation efforts, making it necessary for national legislation to recognise fungi as pillars of biodiversity
and to promote strategies for their conservation.
RESUMEN
Flora, Fauna y una enorme bolsa de microorganismos (incluyendo los hongos) son los elementos que legalmente conforman la
biodiversidad biológica de Colombia. No obstante, la Funga dentro de esta colorida bolsa biodiversa parece ser un elemento
invisible o de claros matices, desestimado por los tomadores de decisiones y excluido de los instrumentos legales que
protegen las especies y los ecosistemas del país. En parte esta desatención se debe a su desconocimiento, teniendo en
cuenta que en este país “fungo diverso” se esperan cerca de 300.000 especies de hongos y se conocen apenas 7.273,
incluyendo 2.670 hongos liquenizados. Curiosamente, la Funga se ha estudiado destacando su potencial como insumo en la
industria alimentaria, farmacéutica y biotecnológica; incluso se ha puesto en evidencia el conocimiento tradicional y el arraigo
cultural hacia los hongos por parte de diferentes grupos étnicos colombianos, así como su potencial como indicadores de
calidad ambiental. Sin embargo, estos esfuerzos no han sido suficientes para hacer de los hongos elementos de alto perfil
en términos de la conservación y para dirigir la atención de las autoridades ambientales, educativas y políticas de Colombia.
Pocos instrumentos legales pueden relacionarse con la protección de la diversidad fúngica pese a que históricamente el
Código Nacional de Recursos Naturales Renovables y de Protección al Medio Ambiente obliga a Colombia a tomar cartas
en el asunto de la protección de la biodiversidad. Solo algunas resoluciones y acuerdos regionales incluyen a los líquenes
y restringen su uso como elementos decorativos en festividades o regulan su extracción en la licencia de proyectos de
infraestructura. En contraste, plantas y animales cuentan con listados de especies amenazadas visibilizados dentro de
resoluciones ministeriales y más de 40 normas que garantizan su protección. Así mismo, en los recursos de la IUCN se
han evaluado y publicado 26.000 especies plantas y 70.000 especies animales, pero solo 425 especies de hongos. Para
Colombia se han publicado 22 especies en la Lista roja de especies amenazadas a nivel global, cinco de estas como
amenazadas. En Suramérica, los micólogos colombianos han participado activamente en este trabajo y se encuentran entre
los líderes en estos esfuerzos. Al día de hoy, hay cerca de 141 especies de hongos no liquenizados se encuentran listas para
evaluar en la plataforma de la iniciativa de Lista roja global de hongos, y se espera que en el corto plazo estas sean evaluadas
por los especialistas y publicadas. Debido al especial interés en los líquenes, en 2019 se realizó la evaluación preliminar por
el grupo de liquenólogos colombianos de 151 especies, con 57% resultando en alguna categoría de amenaza (convenio No
19-098 adscrito entre el Instituto Humboldt, MADS e ICN). Estas evaluaciones son de suma importancia para visibilizar la
Funga en un país donde la deforestación, impulsada por la demanda de combustibles, redes de energía, minería y proyectos
de infraestructura amenaza los ecosistemas de Colombia. Este panorama resalta el color rojo en el que se debe subrayar la
funga de Colombia, haciendo necesario que la legislación nacional reconozca a los hongos como pilares de la biodiversidad
y promueva estrategias para la conservación de los hongos. Para esto se debe robustecer el conocimiento de la diversidad
fúngica en el territorio nacional, incluyendo campos como la taxonomía, la diversidad genética y de poblaciones, la evaluación
de los servicios ecosistémicos prestados por los hongos y las amenazas que actualmente impactan la diversidad fúngica. Este
fortalecimiento debe estar acompañado por la revisión sistemática y fortalecimiento de las colecciones fúngicas, fungarios
y ceparios existentes en el país, lo cual se logrará con la capacitación de investigadores y el reclutamiento de micólogos en
universidades e institutos de investigación. Con lo anterior se generaría un impacto positivo en la evaluación, categorización
y protección de la biodiversidad fúngica colombiana, que indudablemente impactará en la generación de documentos que
permitan a los tomadores de decisiones incluir a los hongos en las agendas nacionales de conservación.
regions such as the Caribbean coast, Orinoco and Pacific/ vascular plants and lichens from excessive exploitation for
Chocó, Insular areas, as well as the Cauca and Magdalena traditionally decorated Christmas cribs (CAR2 Agreement 022
River valleys are even more poorly known (Gaya et al., 2021). of 1993, CDMB3 Resolution 1986 of 1984, CORANTIOQUIA4
Some of those regions currently have the highest rates of Resolution 3183 of 2000, DAMA5 Resolution 1333 of
deforestation and ecosystems that are highly vulnerable 1997)6. Non-vascular plants and lichens have also been
to transformation and loss of ecosystem services (Etter et “occasionally” protected from logging and other projects
al., 2018; Clerici et al., 2020; IPBES, 2021). In addition to that impact ecosystems and biodiversity. In these cases,
diversity, data on fungal distribution, endemism, genetic projects need special permits that include management
diversity, communities, and ecosystem services are scarce, plans for Fauna, Flora, non-vascular plants, and lichens.
with the main threats and risks of extinction being unknown The list of threatened species to be protected was compiled
for most Colombian fungi. The explicit recognition of fungi as in the Resolution 1912 of 2014 (MADS)7 based on data
major components of biodiversity in the country's legislation published in red list books of groups of animals and plants
and policy is urgently needed. It should also include the (vascular and non-vascular: e.g., bryophytes; Linares & Uribe,
terms Fauna, Flora, and Funga when referring to biodiversity, 2002) and orchids (Calderón-Sáenz, 2007). Unfortunately,
as recently suggested by the IUCN (IUCN, 2021a). species of fungi, even lichenised fungi, were not included
in this resolution. A management plan for reducing the loss
CONSERVATION OF BIOLOGICAL DIVERSITY IN COLOMBIA: of biodiversity—sometimes including offsets or restoration
POLICY AND LEGISLATION of ecosystems and protection of certain species—also
Various national norms regulate critical aspects of biodiversity became mandatory following a mitigation hierarchy approach
management and stewarding environmental institutions. The and biodiversity offsetting principles (ten Kate et al., 2004;
National Code of Renewable and Non-renewable Natural MADS 2018). However, in 2019, decree 2106 (República
Resources and Protection of the Environment was created in de Colombia, 2019) dismissed the need for biological
1974 by the decree-law 2811 (República de Colombia, 1974). diversity impact statements for projects, resulting in
With the adoption of the 1991 Colombian Constitution, the adverse effects on the protection of natural ecosystems
country raised the management and protection of natural and the country’s biodiversity.
resources, the environment, and biodiversity (República
de Colombia, 1994). The National Biodiversity Policy that THE CONVENTION ON BIOLOGICAL DIVERSITY
has regulated Colombia since 1994 seeks to promote the In the past ten years, the development of the biodiversity
conservation, knowledge, and sustainable use of biodiversity research strategy for Colombia has followed the commitments
(MADS, 2012). However, this policy only recognises fauna, included in the Convention on Biological Diversity (CBD)
flora, and microorganisms as components of biological signed in Rio de Janeiro (Brazil) and ratified by Colombia
biodiversity. The fact that fungi were not considered an in 1994. The country has also adopted the Strategic Plan
element of the national biological diversity, at the same for Biological Diversity (PEDB 2011-2020) (CBD, 2021a),
level as Flora and Fauna, has left fungi outside the sphere including the Aichi Biodiversity Targets defined under the
of public policies. For 30 years, fungi have been relegated CBD (2011-2020) (CBD, 2021b). This plan provides an
to the background and not recognised as a component of overarching framework for biodiversity management and
Colombia’s biodiversity. This fact ignores the vital role they policy development to halt the loss of biodiversity. This
play in ecosystems and their close relationship with the process should begin with a national assessment of
country’s indigenous and rural cultures, and their potential biodiversity. However, Colombia did not include information
for the biotechnological development of the country. The on fungi in any of the first five Biodiversity National Reports
lack of explicit recognition under national law has resulted in (1998, 2005, 2008, 2010, 2014) (CBD, 1998 onwards;
fungi being unprotected in Colombia. Bello et al., 2014; Gómez et al., 2016). For the fourth report,
Local and regional policies have protected diversity, mycologists prepared a short text about fungal diversity
avoiding the over-harvesting of species and the trade of in Colombia and a case study on ectomycorrhizal fungi
endangered Fauna and Flora, and, exceptionally, also associated with Quercus humboldtii. Still, this information
including other organisms like “slimes”, “moulds”, and was not included in the final report due to “lack of space”
“parasites” (Resolution 0213 of 1977, INDERENA)1. At least (pers. observ.). The 2017 National Biodiversity Strategies
four regional resolutions were established to protect non- and Action Plan (NB) (CBD, 1998 onwards) were designed to
Procesos%20y%20procedimientos%20Vigente/Normatividad_Gnl/Resolucion%20213%20de%201977-Feb-01.pdf
Ministerio de Medio Ambiente y Desarrollo Sostenible (MADS)
7
identify gaps and weaknesses, as well as design actions to for each species, and the revision of habitat perturbation
overcome them. In this report, fungi were barely mentioned in collection sites to allocate threat categories. A total of
and always negatively, presenting only examples of plant and 841 species of lichenised fungi were assessed, resulting
animal pathogens. This is very troubling as these reports in 432 species listed as Critically Endangered, 180
are essential documents for decision-makers, but they as Endangered, 159 as Vulnerable, and 69 as Nearly
ignore fungal diversity and their importance for ecosystems Threatened. Nonetheless, a distributional analysis based
and humans. on the extent of occurrence (EOO) or area of occupancy
But things are changing, and information on fungi was (AOO) was not presented in this preliminary evaluation.
included for the first time in the Sixth Biodiversity National Also, some names used by the authors were not updated,
Report (2019) (CBD, 1998 onwards). This document and a few extralimital species were included, such as
included only general information taken from repositories of Chaenotheca furfuracea or Usnea plicata - both assessed
the Humboldt Institute. Although it did not include curated as Critically Endangered (CR), but recently excluded from
data from mycologists, it was the first time that fungi were the Colombian lichens checklist (Lücking et al., 2021b).
mentioned as part of the biological diversity of Colombia. In the same year, Garcia et al., (2007) reviewed the
In 2021, the Intergovernmental Science-Policy Platform on current knowledge of the uredinal-fungi (rust fungi) known
Biodiversity and Ecosystem Services report was prepared for from Colombia as the first step to create a Red List for
Colombia (IPBES 2021). It compiled relevant and published Colombian rust fungi. They observed that most collections
information about fungal diversity (genetic, species, and had been made on weedy plants, with few records from wild
ecosystem diversity levels) and presented attributes such as or threatened plants. They proposed some strategies to
the composition, structure, and function of this diversity. The document the hosts of endangered rusts within the National
report calls for scientific and monitoring programs to focus Program for the Study of the Country's Megadiversity
on the least-sampled groups of Colombia’s biodiversity, framework in the next 25 Years (Forero, 1999). The strategy
such as invertebrates, liverworts and hornworts, bacteria, included emphasising that collections from wild areas
archaea, and fungi (IPBES, 2021). should be prioritised in sites with high endemism rates
With the recent creation of a Ministry of Science, and where hosts are at risk of disappearing due to climate
Technology, and Innovation and the strengthening of research change (e.g., moors, coasts) as they are colonization fronts
capacities, the country’s high fungal diversity and the recent and centres of uredinal diversification. They emphasised
boom in information on and interest in the potential and the importance of working together with botanists who
importance of fungi has created a massive opportunity for know the hosts, and the same time, advising them to
the Colombian government to place this neglected group at collect botanical specimens showing disease symptoms,
the same level as Flora and Fauna. drying the material separately, and enriching the collection
of rust associated with native plants around the country.
RED LIST OF FUNGI Unfortunately, the red list book was never carried out, but
The International Union for Conservation of Nature (IUCN) the suggestions from 14 years ago remain valid.
Red List of threatened species assigns a category to each As mentioned above, lichens have a different
species according to its risk of extinction. The appropriate history because initially, they were studied by botanists
category is assigned for each species on the basis of specializing in bryophytes, making them more visible to the
assessments from each standard criteria that could consider research community (Aguirre & Rangel, 2017). In 2019, the
their vulnerability and threat characteristics (IUCN, 2021b). first national assessment of lichens species focusing on
Assessments are a powerful tool for supporting conservation endemic species was carried out, thanks to an initiative
policy, planning, and action. They help authorities to delineate led by the Instituto Alexander von Humboldt (Humboldt-
protected areas, guide allocation of funding, and influence MADS-ICN agreement No 19-098), in which the national
decision making. Recently a new standard tool was validated lichenologist community actively contributed. A total of
as a Red List of Threatened Species ally, the IUCN Green 151 species were preliminarily assessed, of which 57%
Status of Species, helping to evaluate species recovery and species were listed as Threatened: 56 species as Critically
conservation impact (Grace et al., 2021; IUCN, 2021b). Endangered (CR), 19 species as Endangered (EN), and ten
species as Vulnerable (VU) (Figure 1b, 3a). The resultant
RED LIST OF FUNGI IN COLOMBIA: LOCAL INITIATIVES list is an important input for biodiversity inventories that
National assessments arise from a predetermined list are required for infrastructure projects.
of species, and there is only one assessment for each
taxonomic group. In Colombia, national evaluations of GLOBAL RED LIST OF FUNGI
Fauna and Flora are published in the Red Book, but there Assessments of the risk to species made for the Global Red
is no published Red Book of Fungi to date. List are being carried out by taxonomic-specialist groups and
In 2007, Aguirre and Rangel carried out an exercise to aim to cover all the existing species in a group. For fungi,
categorise the threatened species of lichens and mosses there are five specialist groups: i) Chytrid, Zygomycete,
of Colombia, partially following the IUCN criteria. For this Downy Mildew, Slime Mould, ii) Cup-Fungus, Truffle and Ally,
analysis, they used herbarium collections, distribution data iii) Lichen, iv) Mushroom, Bracket and Puffball, and v) Rusts
A B
C D
FIGURE 1. A Colombian fungal species globally assessed and published in the IUCN Red List (data up to 2021). B Colombian fungal
species that have been globally proposed and published in The Global Fungal Red List Initiative (data up to July 2021). C Results of the
preliminary assessment of lichenised species, these evaluations were carried out by the Colombian lichenologist team (Humboldt-MADS-
ICN, agreement No 19-098). D Status in the evaluation process of all fungal species with distribution in Colombia that are currently on the
global fungal red list initiative platform.
and Smuts. A specific platform to assess fungal species interest of some mycologists—a dynamic process of work for
has been developed (A Global Fungal Red List, http://iucn. the conservation of fungi has started, encouraged by David
ekoo.se/en/iucn/welcome), and the final evaluations are Minter and Greg Mueller. Since 2011 various Colombian
published on the Global Red List (https://www.iucnredlist. mycologists have joined the Group for the Conservation of
org). Although fungi are one of the world’s most biodiverse Fungi in Latin America and the Caribbean, associated with
groups, they are the most under-represented multicellular the Latin American Mycology Association (ALM). Also, they
taxa on The IUCN Red List, with only 550 assessments of the were appointed to three of the five IUCN specialist groups:
over 150,000 described fungal species currently published Cup-Fungi, Truffles and Allies; Lichens; and Mushrooms,
or less than 0.3% (Gonçalves et al., 2021; IUCN 2022). Brackets and Puffballs. By being part of the specialist
In comparison, the 2021 edition of the Red List includes groups, the researchers could participate in fungal red-
54,127 and 79,858 species of plants and animals, listing workshops (2015, 2020). In addition, efforts have
respectively. In recent years—and thanks to the particular been made to increase the number of species proposed
FIGURE 2. A Gloiocephala quercetorum, saprotrophic fungi associated with Quercus humboldtii forests in a montane region in Colombia.
B Basidiomas of the ectomycorrhizal fungi Leccinum andinum, associated with Quercus species. C Basidiomas of the ectomycorrhizal fungi
Tylopilus obscurus associated with Quercus species. D Basidiomas of the endemic and ectomycorrhizal fungi Austroboletus amazonicus,
associated with Pseudomonotes tropenbosii (Dipterocarpaceae), Amazonian region. E Ascomas of Pseudolostoma volvatum associated with
P. tropenbosii (Dipterocarpaceae), Amazonian region. F Clavulina tepurumenga associated with Dicymbe uaipirensis (Dipterocarpaceae),
Amazonian region. G Lichen Sticta peltigerella. H Lichen Icmadophila aversa. (Photographs A-F by Aída Vasco; G by Robert Lücking; and H
by Bibiana Moncada.)
to be evaluated under the IUCN criteria. Two workshops on lands for cattle, land speculation, legal and illegal mining,
applying the IUCN criteria for fungi were recently offered urban expanse, and illicit crops (Figure 3b) (Clerici et al.,
to Latin American mycologists (2020–2021) to encourage 2021, Etter et al., 2008 Figure 4). Annual and perennial
the mycological community. The threat status of South non-timber crops have been identified as the major threat
American fungal species is only now beginning to be for fungal species. In Colombia, nearly four million hectares
assessed. Colombian mycologists have been active in this are occupied by different crops, with coffee, oil palms,
work and are among the leaders of these efforts. sugar cane, and cotton representing 48.7% of the total
Colombia is among the Latin American countries with the area (DANE, 2021). The threat known as biological resource
highest number of threatened living species on the IUCN Red use (logging and wood harvesting), or in other words,
List, grouping nearly 2.1% of total recorded species (IUCN, deforestation and habitat fragmentation, impacts nearly
2021b). A total of 721 Colombian plants are included in the 50–60% of all preliminarily assessed fungal species (Figure
2021 IUCN Red List. Amphibians with 289 listed species 3a). Deforestation is increasing in the country by about
follow in second place. Only 22 Colombian fungal species 800,000 hectares per year, mainly through converting native
have been globally assessed and published on the IUCN Red vegetation to pasture and extensive oil palm plantations
List (2021b). All but two of these belong to Agaricomycotina (Figure 3a, 4) (IDEAM, 2021; IPBES, 2021). Although natural
(mushrooms and relatives). The other two are ascomycetes forests in the Andean region are under anthropic pressures
belonging to Pezizomycotina, and two others, lichenised fungi. and the montane tropical forests currently occupy less
Austroboletus amazonicus is listed as Critically Endangered than 30% of their original extent (Forero-Medina & Joppa,
(Figure 2d). Four species were listed as Vulnerable (VU), 2010), the current nuclei of deforestation are located in
six as Nearly Threatened (NT), five as Least Concern (LC) the Orinoco basin, Amazonian region, Chocó biogeographic,
and two as Data Deficient (DD) (Figure 1). This effort has and Serranías in the Caribbean region (Etter et al., 2017;
already yielded positive and very encouraging results, with IPBES, 2021, Figure 4). Unfortunately, those correspond to
170 additional species proposed on the global fungal red list areas where there are the biggest gaps in the knowledge of
initiative site. Those species belonged to different groups that fungal diversity (Figure 3b, 4) (Chapter 3, 4).
include Macrofungi from Ascomycota and Basidiomycota (106 The conservation status of habitats and hosts is also
species), arbuscular mycorrhizae (15 species), myxomycetes essential for fungi. For example, Sticta peltigerela (VU,
(13 species), and lichens (54 species). Most unpublished B1ab (iii)) grows in non-polluted paramo and montane
species are in the preliminary step “proposed”, but others forest environments close to streams or floodplains with
are in various assessment levels (Figure 1b). Data for the clean waters. On the other hand, Hericium erinaceus (LC)
54 lichen species 48 endemic, will be published soon at the is probably a weak necrotrophic parasite growing on old
IUCN Red List of Threatened Species. deciduous and sclerophyllous trees, mainly old Quercus,
One of the most significant difficulties when evaluating in mature forests. Fomitiporia bambusarum (LC) is a
the species is the lack of information, especially on plant pathogen or wood saprotroph which has only been
population size, total distribution, and status. The reported growing on bamboos. Ectomycorrhizal fungi
categorisation of most fungal species has been assessed depend on their host, and this is the case for Austroboletus
amazonicus, a species that is restricted to small areas
on the basis of species distribution data, biological
associated with patches of Pseudomonotes tropenbosii
information, the threats posed to the host plants (e.g.:
(Dipterocarpaceae) in the Amazonia region (Vasco et al.,
pathogenic or symbiotrophs) and habitats (Corrales et al.,
2014, 2020). The ectomycorrhizal species associated with
2020; Vasco et al., 2020). For example, nearly all evaluated
Quercus in montane neotropical forests share the category
species are ectomycorrhizal fungi associated with Quercus
of vulnerable with their plant host, as almost 42% of the
humboldtii in tropical mountain forests (e.g., Vasco et al.
Q. humboldtii populations in Colombia are already lost
2020). These species have usually been recorded from just
(Cárdenas & Salinas, 2007). The remaining populations are
a few localities in the country Colombia, which means that
also highly fragmented, and the effect of this fragmentation
their total distribution is unknown. However, the geographic
on the diversity of the communities of ectomycorrhizal fungi
range was established using data on the distribution of
is unknown.
oak forests. This extrapolation is possible for fungi that
The introduction of exotic trees and their associated
establish this specific interaction type, but it is more difficult
fungi is one aspect that has received little attention
for other guilds, such as saprotrophic fungi or arbuscular
in tropical forests. Amanita muscaria is a mycobiont
mycorrhizal fungi.
associated with Pinus, that was introduced with the tree in
the nineties. This species is now competing and replacing
MAIN THREATS native ectomycorrhizal fungi on the roots of Quercus
The major causes of biodiversity loss in Colombia (Vargas et al., 2019). Some of these exotic species can be
are the fragmentation, degradation or loss of habitat, aggressive and more tolerant of environmental changes,
overexploitation, invasive species, pollution, and enabling them to out-compete native species.
climate changes (Clerici et al., 2020, 2021 Figure 3a,b). The impact of fertilizers and pesticides on the fungal
Conversions of primary forests in the country are primarily communities in natural systems in Colombia is unknown.
due to expansion of the agricultural frontier and grazing Colombia has the highest fertiliser consumption and the
FIGURE 3. A Major threats to fungi assessed for the IUCN Red List of Threatened Species. The data were downloaded from the IUCN Red
List of Threatened Species (2021-3); B Map of current threats affecting biological diversity in Colombia.
second-highest consumption of pesticides per hectare their primary condition may be the best near-term solution
in Latin America (www.foodandlandusecoalition.org). The to protect fungi. In Colombia, fungi have been indirectly
effect of anthropogenic nitrogen deposition that rises protected when habitats for animals and plants are
nitrogen levels, mainly from local and transboundary air conserved. The country has 56 protected areas (Hurtado-
pollution and fertilisers and fumigants, has been reported Guerra et al., 2013), with a total area representing more
widely in fungal species of temperate forests (Sapsford than 10% of the territory, even though this system does
et al., 2017; Lilleskov et al., 2019). In the tropics, studies not yet include all Colombian ecosystems. In addition,
show that nitrogen additions may reduce ectomycorrhizal deforestation in protected areas has increased during the
colonization of plant-host roots and may alter the fungal post-conflict period (Clerici et al., 2020).
composition (Corrales et al., 2017). Pesticides accumulate
in the soil over time, being transported and deposited by
WHERE SHOULD EFFORTS FOR FUNGAL CONSERVATION IN
air, and can thus contaminate non-agricultural areas. They
COLOMBIA BE CONCENTRATED?
may affect the nutritional quality of soils, disrupt enzymatic
activities, and alter soil fungal communities (Sapsford et A primary task should be to ensure that the Colombian
al., 2017). For example, insecticides and fungicides inhibit legislation recognises fungi on a par with animals and plants .
the colonisation of root plants by arbuscular mycorrhizas There is a greater awareness of the role and importance
and ectomycorrhizas (Laatikainen & Heinonen-Tanski, of fungi in ecosystems. Yet, their impact on life on this planet
2002; Karpouzas et al., 2014). This is an issue that must remains underestimated and unknown by the public, other
be taken into account for further studies and that impacts scientists and policy makers. The explicit recognition of fungi
the conservation of fungi. as major components of biodiversity in the country in terms
As we mentioned before, fungi play fundamental roles of legislation and policy is urgently needed, promoting the
in key ecosystem functions and services, so it is critical use of the terms fauna, flora and funga. Fungi need active
that they are included in conservation strategies. Given and specific inclusion in national surveys and inventories,
the lack of information, preserving tropical ecosystems in not just passive and implicit protection.
FIGURE 4. Distribution of the fungal species evaluated under the IUCN criteria (data up to 2021). The stars are published species and pink
circles are species that are under assessment. The red circles are the early deforestation warnings (2018).
Acknowledgements Corrales A, Sanchez Tello JD, Benjumea C, Nino Fernandez YM, Pena-
Canon R, Vasco-Palacios AM (2020). Lactifluus hallingii (amended
Special thanks are given to the Fungal Red List initiative for version of 2020 assessment). The IUCN RedList of Threatened
supplying the data; and to Instituto Alexander von Humboldt for Species 2020: e.T174799230A179540212
the data from the preliminary evaluation of prioritised lichens, DANE (2021) — Departamento Administrativo Nacional de Estadística
Agreement 19-098 Humboldt-MADS-ICN. We are thankful to – Encuesta Nacional Agropecuaria. Available at: https://www.dane.
gov.co/index.php/estadisticas-por-tema/agropecuario/ encuesta-
Yurani Caicedo Chalapud for helping with data processing, and the
nacional-agropecuaria-ena
reviewers Raquel Negrao and David Hammond for comments and Correa-Ochoa MA, Vélez-Monsalve LC, Saldarriaga-Molina JC, Jaramillo-
suggestions that improved the manuscript. Ciro MM (2020) Evaluation of the Index of Atmospheric Purity in an
American tropical valley through the sampling of corticolous lichens
in different phorophyte species. Ecological Indicators, 115, 106355.
References doi: 10.1016/j.ecolind.2020.106355
Dix NJ, Webster J (1995) Fungal ecology. Dordrecht: Springer.
Aguirre J, Rangel-Churio JO (2007) Amenazas a la conservación de las Donegan T, McMullan WM, Quevedo A, Salaman P (2013) Revision of
especies de musgos y líquenes en Colombia-Una aproximación the status of bird species occurring or reported in Colombia 2013.
inicial. Caldasia, 29(2), 235-262. Conservación Colombiana, 19, 3-10.
Avendaño JE, Bohórquez CI, Rosselli L, Arzuza-Buelvas D, Estela FA, Dugan FM (2011) Conspectus of world ethnomycology: fungi in ceremonies,
Cuervo AM, Stiles FG, Renjifo LM (2017) Checklist of the birds of crafts, diets, medicines, and myths. APS Press.
Colombia: a synthesis of the state of knowledge since Hilty and Etter A, McAlpine C, Possingham H (2008) Historical patterns and drivers
Brown (1986). Ornitologia Colombiana, 16, eA01–eA11. of landscape change in Colombia since 1500: a regionalized spatial
Bello JC, Baez M, Gómez MF, Orrego O, Nägele L (2014) Estado y approach. Annals of the Association of American Geographers, 98(1),
tendencias de la biodiversidad continental de Colombia. Bogota DC: 2-23.
Instituto Alexander von Humboldt. Etter A, Andrade A, Saavedra K, Amaya P, Arevalo P, Cortes J, Pacheco
Bernal R, Gradstein R, Celis M (2020) Catálogo de Plantas y C, Soler D (2017) Lista roja de ecosistemas de Colombia version
Líquenes de Colombia. Bogotá DC: Instituto de Ciencias 2.0. Pp.1-6.
Naturales, Universidad Nacional de Colombia. Available at: http:// Etter A, Andrade Á, Saavedra K, Cortés J (2018) Actualización de la Lista
catalogoplantasdecolombia. unal.edu.co Roja de los Ecosistemas Terrestres de Colombia. In: Moreno LA,
Betancur J, Sarmiento H, Toro-González L, Valencia J (2015) Plan para el Rueda C, Andrade GI (eds). Biodiversidad 2017: Estado y tendencias
estudio y la conservación de las orquídeas en Colombia. Bogotá DC: de la biodiversidad continental de Colombia. Bogotá DC: Instituto de
Ministerio de Ambiente y Desarrollo Sostenible. Universidad Nacional Investigación de Recursos Biológicos Alexander von Humboldt. 19p.
de Colombia. 336p. Fajardo J, Lessmann J, Bonaccorso E, Devenish C, Muñoz J (2014)
Cárdenas D, Salinas N (2007) Libro rojo de plantas de Colombia. Especies Combined use of systematic conservation planning, species
maderables amenazadas I Parte. Bogotá DC: Instituto Amazónico de distribution modelling, and connectivity analysis reveals severe
Investigaciones Científicas SINCHI. Ministerio de Ambiente, Vivienda conservation gaps in a megadiverse country (Peru). PLoS ONE,
y Desarrollo Territorial. 10(3), e0122159. doi: 10.1371/journal.pone.0122159
Calderón-Sáenz E (2007) Libro rojo de plantas de Colombia, volumen 6: Forero E (1999) La sistemática en Colombia para el siglo XXI. Revista
Orquídeas, primera parte. Bogota DC: Serie Libros Rojos de Especies de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales,
Amenazadas de Colombia. Instituto Alexander von Humboldt y 23(86), 129-138.
Ministerio de Ambiente, Vivienda y Desarrollo Territorial. 828p. Forero-Medina G, Joppa L (2010) Representation of global and national
CEIBA (2019) Especies de Líquenes Priorizadas en Colombia. Available at: conservation priorities by Colombia’s protected area network. PLoS
http://i2d.humboldt.org.co/ceiba/resource.do?r=rrbb_liquenes_2019 ONE, 5(10), e13210. doi: 10.1371/journal.pone.0013210
Chiriví J, Danies G, Sierra R, Schauer N, Trenkamp S, Restrepo S, Frąc M, Hannula SE, Bełka M, Jędryczka M (2018) Fungal biodiversity
Sanjuan T (2017) Metabolomic profile and nucleoside composition and their role in soil health. Frontiers in Microbiology, 9, 707. doi:
10.3389/fmicb.2018.00707
of Cordyceps nidus sp. nov. (Cordycipitaceae): a new source of active
Gadd G, Watkinson SC, Dyer PS (2007) Fungi in the Environment, vol. 25.
compounds. PLoS ONE, 12(6), e0179428. doi: 10.1371/journal.
Cambridge: Cambridge University Press.
pone.0179428
Gaya E, Vasco-Palacios AM, Vargas-Estupiñán N, Lücking R, Carretero
Clerici N, Armenteras D, Kareiva P, Botero R, Ramírez-Delgado JP, Forero-
J, Sanjuan T, Moncada B, Allkin B, Bolaños Rojas AC, Castellanos-
Medina G, Ochoa J, Pedraza C, Schneider L, Lora C, Gómez C, Linares
Castro C, Coca LF, Corrales A, Cossu T, Davis L, dSouza J, Dufat A,
M, Hirashiki C, Biggs D (2020) Deforestation in Colombian protected
Franco-Molano AE, García F, Gómez-Montoya N, González-Cuellar FE,
areas increased during post-conflict periods. Scientific Reports,
Hammond D, Herrera A, Jaramillo-Ciro MM, Lasso Benavides C, Mira
10(1), 1-10. doi: 10.1038/s41598-020-61861-y
MP, Morley J, Motato-Vásquez V, Niño-Fernández Y, Ortiz-Moreno ML,
Clerici N, López S, Molina LF, Pinilla MC, Ríos H, Sánchez A, Villegas C,
Peña-Cañón ER, Ramírez-Castrillón M, Rojas T, Ruff J, Simijaca D,
Villegas JC. (2021). Capítulo 4. Motores directos de transformación Sipman HJM, Soto-Medina E, Torres G, Torres-Andrade PA, Ulian T,
y pérdida de biodiversidad. En: IPBES Evaluación Nacional de White K, Diazgranados M (2021) ColFungi Colombian resources for
Biodiversidad y Servicios Ecosistémicos. fungi made accessible. Richmond: Royal Botanic Gardens, Kew. 36p.
CBD (1998 onwards) — Convention on Biological Diversity National García GD, Buriticá P, Henao MLM (2007) Elementos para la elaboración
Reports and NBSAPs (Reports at 1998, 2005, 2008, 2010, 2014). del libro rojo de hongos royas de Colombia. Revista de la Academia
Available at: https://www.cbd.int/reports/search/?country=co Colombiana de Ciencias Exactas Físicas y Naturales, 31(121), 449-468.
CBD (2021a) — Convention on Biological Diversity National Biodiversity Gómez MF, Moreno LA, Andrade GI, Rueda C (2016) Biodiversidad 2015.
Strategies and Action Plans (NBSAPS). Available at: https://www.cbd. Estado y tendencias de la biodiversidad continental de Colombia.
int/nbsap/about/latest/ Bogota DC: Instituto Alexander von Humboldt.
CBD (2021b) — Convention on Biological Diversity Strategic Plan Gonçalves SC, Haelewaters D, Furci G, Mueller GM (2021) Include all
for Biodiversity 2011-2020, including Aichi Biodiversity Targets fungi in biodiversity goals. Science, 373(6553), 403. doi: 10.1126/
(PEDB 2011-2020). Available at: https://www.cbd.int/kb/record/ science.abk1312
decision/12268 González FE, Lasso-Benavides CM, Adrada-Gómez BY, Sanabria-Diago
Corrales A, Turner BL, Tedersoo L, Anslan S, Dalling JW (2017). Nitrogen OL, Vasco-Palacios AM (2021) Estudio etnomicológico con tres
addition alters ectomycorrhizal fungal communities and soil enzyme comunidades rurales ubicadas en la zona andina del departamento
activities in a tropical montane forest. Fungal Ecology, 27, 14-23 del Cauca, Colombia. Boletín de Antropología, 36(62), 147-164.
Grace MK, Akçakaya HR, Bennett EL, Brooks TM, Heath A, Hedges S, TW, Meyer W, Nilsson HR, Öpik M, Robert V, Stadler M, Thines M,
Vasco-Palacios AM, Young S (2021) Testing a global standard for Vu D, Yurkov AM, Zhang N, Schoch CL (2021a) Fungal taxonomy and
quantifying species recovery and assessing conservation impact. sequence-based nomenclature. Nature Microbiology, 6, 540-548.
Conservation Biology, early view. doi: 10.1111/cobi.13756 doi: 10.1038/s41564-021-00888-x
Hurtado Guerra A, Santamaria Gomez M, Matallana Tobon CL (2013). Lücking R, Moncada B, Soto-Medina E, Simijaca D, Sipman HJ (2021b)
Plan de Investigación y Monitoreo del Sistema Nacional de Áreas Nomenclatural and taxonomic update to the Catálogo de Líquenes de
Protegidas (Sinap): avances construidos desde la Mesa de Colombia. Revista de la Academia Colombiana de Ciencias Exactas,
Investigación y Monitoreo entre 2009 y 2012. Bogota, D.C.: Instituto Físicas y Naturales, 45(174), 147-190.
de Investigación de Recursos Biológicos Alexander von Humboldt y MADS (2012) — Ministerio de Ambiente y Desarrollo Sostenible Política
Parques Nacionales Naturales de Colombia. 200 pp. Nacional de Gestión Integral de la Biodiversidad y sus Servicios
IDEAM (2021) Resultados de Monitoreo de Deforestación en 2019. Ecosistémicos PNGIBSE. Available at: https://www.minambiente.gov.
Available at: http://www.ideam.gov.co documents/10182/105413996/ co/ index.php/bosques-biodiversidad-y-servicios-ecosistematicos/
presentacion balancedeforestacion2019/7c9323fc-d0a1-4c95-b1a1- politica-nac ional-de-biodiversidad
1892b162c067 MADS (2018) — Ministerio de Ambiente y Desarrollo Sostenible
INDERENA (1977) — Instituto Nacional de los Recursos Naturales Resolución 256, por la cual se adopta la actualización del Manual de
Resolución 0213 de 1977. Available at: https://cvc.gov.co/sites/ Compensaciones Ambientales del Componente Biótico. Bogota DC.:
default/files/ Sistema_Gestion_de_Calidad/Procesos%20y%20 Diario Oficial 50.525.
procedimientos%20Vigente/Normatividad_Gnl/Resolucion%20 Méndez MJ, Caicedo NH, Salamanca C (2018) Trametes elegans: a fungal
213%20de%201977-Feb-01.pdf endophytic isolate from Otoba gracilipes as biocatalyst for natural
Instituto de Investigación de Recursos Biológicos Alexander von flavours production. New Biotechnology, 44, S75. doi: 10.1016/j.
Humboldt (2017) Boletín de prensa Instituto Humboldt - Biodiversidad nbt.2018.05.892
colombiana: números para tener en cuenta. Available at: http:// Mueller GM, Foster M, Bills G (2004) Biodiversity of Fungi. Inventory and
w w w.humboldt.org.co/es/boletines-y- comunicados/item/1087- Monitoring Methods. Burlington: Academic Press (Elsevier).
biodiversidad-colombiana-numero-tener-en-cuenta Peña-Cañón ER, Enao-Mejía LG (2014) Conocimiento y uso tradicional
IPBES (2021) Evaluación Nacional de Biodiversidad y Servicios de hongos silvestres de las comunidades campesinas asociadas
Ecosistémicos. Resumen para Tomadores de Decisión. Available a bosques de roble (Quercus humboldtii) en la zona de influencia
at: http://www.humboldt.org.co/images/pdf/RTD%20Final%20 de la Laguna de Fúquene, Andes Nororientales. Etnobiología, 12(3),
v290521%20Lanzamiento.pdf 28-40.
IUCN (2021a) IUCN SSC acceptance of Fauna Flora Funga. Available at: Pérez-Moreno J, Guerin-Laguette A, Arzú RF, Yu FQ (2020) Mushrooms,
https://www.iucn.org/commissions/species-survivalcommission/ Humans and Nature in a Changing World: Perspectives from Ecological,
about/ssc -committees/fungalconservation-committee Agricultural and Social Sciences. Springer Nature.
IUCN (2022). The IUCN Red List of Threatened Species. Version 2021-3. Ramírez-Morán NA, León-Gómez M, Lücking R (2016) Use of lichen
https://www.iucnredlist.org. ISSN 2307-8235. Search on 2021-09-
biotypes as bioindicators of perturbation in fragments of high
09
Andean Forest (“Encenillo” Biological Reserve, Colombia). Caldasia,
IUCN (2021b) The IUCN Redlist of Threatened Species (2021) Summary
38(1), 31-52.
Statistics. Available at https://www.iucnredlist.org/resources/
República de Colombia (1974) Decreto 2811 del 18 de diciembre de
summary-statistics
1974. Código Nacional de Recursos Naturales Renovables y de
Jaramillo DA, Méndez MJ, Vargas G, Stashenko EE, Vasco-Palacios
Protección al Medio Ambiente.
AM, Ceballos A, Caicedo NH (2020) Biocatalytic potential of native
República de Colombia (1994) Ley 165 de 1994 Por medio de la cual
basidiomycetes from Colombia for flavour/aroma production.
se aprueba el "Convenio sobre la Diversidad Biológica", hecho en
Molecules, 25(18), 4344. doi: 10.3390/molecules25184344
Río de Janeiro el 5 de junio de 1992. Available at: https://test-
Karpouzas DG, Papadopoulou E, Ipsilantis I, Friedel I, Petric I, Udikovic-
www. minambiente.gov.co/wp-content/uploads/2020/06/Ley-165-
Kolic N, Djuric S, Kandeler E, Menkissoglu-Spiroudib F, Martin-Laurent
F (2014) Effects of nicosulfuron on the abundance and diversity of de-1994.pdf
arbuscular mycorrhizal fungi used as indicators of pesticide soil República de Colombia (2019) Decreto 2106 de 2019 por el cual
microbial toxicity. Ecological Indicators, 39, 44-53. se dictan normas para simplificar, suprimir y reformar trámites,
Koleff P, Urquiza-Haas T (2011) Planeación para la conservación de la procesos y procedimientos innecesarios existentes en la
biodiversidad terrestre en México: retos en un país megadiverso. administración pública. Available at: https://dapre.presidencia.gov.
México DF: Comisión Nacional para el Conocimiento y Uso de la co/normativa/normativa/ DECRETO%202106%20DEL%2022%20
Biodiversidad, Comisión Nacional de Áreas Naturales Protegidas. DE%20NOVIEMBRE%20DE%202019.pdf
Laatikainen T, Heinonen-Tanski H (2002) Mycorrhizal growth in pure Rodríguez OE, Andrade WA, Díaz FE, Moncada B (2015) Actividad
cultures in the presence of pesticides. Microbiological Research, antimicrobiana de líquenes de la cuenca alta del río Bogotá. Nova,
157(2), 127-137. 13(23), 65-72.
Lilleskov EA, Kuyper TW, Bidartondo MI, Hobbie EA (2019) Atmospheric Samper K (1998) Biodiversity research in Colombia: what we know and
nitrogen deposition impacts on the structure and function of forest what we need to know. In: Seminar proceedings: Research in tropical
mycorrhizal communities: a review. Environmental Pollution, 246, rainforests: its challenges for the future, Wageningen (Netherlands),
148-162. doi: 10.1016/j.envpol.2018.11.074 25-26 Nov 1997. Tropenbos Foundation.
Linares EL, Uribe-M J (2002) Libro rojo de Briofitas de Colombia. Libros Sapsford SJ, Paap T, Hardy GESJ, Burgess TI (2017) The ‘chicken or the
rojos de especies amenazadas de Colombia. Bogota DC: Instituto de egg’: which comes first, forest tree decline or loss of mycorrhizae?
Ciencias Naturales-Universidad Nacional de Colombia. Ministerio del Plant Ecology, 218(9), 1093-1106. doi: 10.1007/s11258-017-0754-6
Medio Ambiente. ten Kate K, Bishop J, Bayon R (2004) Biodiversity offsets: views,
López-Legarda XL, Echavarría CA, Sánchez FS (2015) Producción de experience, and the business case. London, UK, Gland, Switzerland
polisacáridos a partir de Ganoderma sp., aislado en la región andina. and Cambridge, UK: IUCN and Insight Investment.
Revista Colombiana de Biotecnología, 17(2), 44-54. Vargas N, Gonçalves SC, Franco-Molano AE, Restrepo S, Pringle A (2019)
Lücking R, Hodkinson BP, Leavitt SD (2017) The 2016 classification In Colombia the Eurasian fungus Amanita muscaria is expanding its
of lichenised fungi in the Ascomycota and Basidiomycota – range into native, tropical Quercus humboldtii forests. Mycologia,
approaching one thousand genera. The Bryologist, 119, 361-416. doi: 111(5), 758-771. doi: 10.1080/00275514.2019.1636608
10.1639/0007-2745-119.4.361 Vasco-Palacios AM, Suaza SC, Castanõ-Betancur M, Franco-Molano
Lücking R, Aime MC, Robberts B, Miller AN, Aoki T, Ariyawansa HA, AE (2008) Conocimiento etnoecológico de los hongos entre los
Cardinali G, Crous PW, Druzhinina IS, Geiser DM, Hawksworth DL, indígenas Uitoto, Muinane y Andoke de la Amazonía Colombiana.
Hyde KD, Irinyi L, Jeewon R, Johnston PR, Kirk PM, Malosso E, May Acta Amazonica, 38, 17-30.
ABSTRACT
Colombia ranks among the 17 countries considered megadiverse, hosting almost 10% of the planet’s biodiversity and only
being surpassed by Brazil, a country seven times larger than Colombia. Unfortunately, fungi are mostly left out of these counts.
One of the issues faced when attempting to report the fungal diversity in Colombia is the inability to account for the fungal
specimens scattered in national and international collections, which are often not properly curated or digitised. In this chapter,
we describe the first attempt to compile the available information on fungal collections from Colombia as a contribution
towards a more realistic estimation of their diversity and attempt to establish a baseline to improve mycological knowledge in
the country. Databases of national and international biological collections and personal databases from several mycologists
were consulted, using the keywords “Colombia”, “fungi”, “hongos”, “lichens”, and “liquenes”. Uncatalogued Colombian
collections were also examined when possible, and experts and collection managers were also consulted. All metadata
associated with the voucher specimens (e.g., locality, collection date, ecology and classification) were extracted and analysed
using descriptive statistics. We found, unsurprisingly, that species of fungi that are considered economically important or
with potential biotechnological applications, or otherwise conspicuous or enigmatic, were overall the best represented within
national and international collections. We highlight the need to increase sampling efforts in the country, as well as the need
for regularisation and proper digitisation of specimens from biological collections so that they can be appropriately accounted
for in future fungal research. This chapter is expected to be an invitation for mycologists to increase their efforts to improve
the knowledge of Colombian funga through proper curation and documentation of their collections, and urges the institutions
that promote biological research to support the growth and maintenance of Colombian fungal collections.
RESUMEN
Colombia está entre los 17 países considerados megadiversos, con casi el 10% de la biodiversidad del planeta, sólo superado
por Brasil, un país siete veces mayor que Colombia. Desafortunadamente, la mayoría de los hongos quedan fuera de estos
recuentos. Uno de los problemas al intentar informar sobre la diversidad de hongos en el país es la incapacidad de dar cuenta
de los especímenes de hongos dispersos en colecciones nacionales e internacionales que a menudo no están debidamente
curadas o documentadas. En este capítulo realizamos el primer intento de recopilar la información disponible sobre colecciones
de hongos en Colombia para contribuir a una estimación más realista de su diversidad y establecer una base para mejorar
el conocimiento micológico en el país. Para ello, se consultaron bases de datos de colecciones biológicas nacionales e
internacionales y bases de datos personales de varios micólogos, utilizando los modificadores de palabras clave “Colombia”
y “fungi” o “hongos”. También se examinaron colecciones colombianas no catalogadas y se consultaron especialistas y
directores de colecciones. Todos los metadatos asociados a los especímenes avalados (por ejemplo, localidad, fecha de
recolección, ecología, clasificación, etc.) fueron extraídos y analizados usando estadísticas descriptivas. Encontramos, como
era de esperar, que las especies de hongos considerados económicamente importantes o con potenciales aplicaciones
biotecnológicas, o alternativamente enigmáticas o conspicuas, fueron en general las mejor representadas dentro de las
colecciones nacionales e internacionales. Destacamos aquí la necesidad de incrementar los esfuerzos de muestreo en el
país, así como la regularización y adecuada digitalización de los especímenes de colecciones biológicas para que puedan ser
debidamente contabilizados en futuras investigaciones fúngicas. Se espera que este capítulo sea una invitación para que
los micólogos incrementen sus esfuerzos para mejorar el conocimiento sobre la funga colombiana, mediante la curación y
documentación apropiada de sus colecciones, así como instar a las instituciones y entidades de fomento a la investigación
biológica a apoyar el crecimiento y mantenimiento de las colecciones de la funga colombiana.
A
Aections
Major group
MajorA group A
Major collections
group
Major collections
Major
group
collections
group collections
collections Fungal
Fungal collections in Fungal
collections
Colombia collections
Fungal
Fungal collections
in collections
Colombia
in Colombia
Fungal
in Colombia
collections
in Colombiain Colombia Specimens
Specimens per fungariumSpecimens
per fungarium
per fungari
Specimens perSpecimens
Speci
fungarium per f
microfungi AM microfungi AM microfungi AM
yeasts 1% microfungi AM
yeasts yeasts CBUMAG:LIC HEUS CBUMAG:LIC CBUMAG:LIC HEUS HEUSCBUMAG:LIC
s 1%AM 1% LLANOS CBUMAG:LIC
microfungi 0% 0% 1% yeasts 0% 1% LLANOS 1%
0% CBUMAG:LIC 1% 0%LLANOS 0% 1%HEUS
1% yeasts 0% 1%
MEDEL 1%Otros 1% HEUS1% 1%
MEDEL LLANOS
Otros MEDEL Otros
0%
JBB JBBLLANOS JBB 0% JBB
0%
filamentous fungifilamentous fungi filamentous fungi TOLI 0% 2% 1% TOLI 0% JBB
MEDEL 2% 1%
TOLI
Otros 0% 2% MEDE
AFungal collections
plant pathogens plant pathogens plant pathogens 1% 1% 1% TOLI 1%
Major group collections Fungal collections in Colombia
omopathogenic entomopathogenic
filamentous entomopathogenic
fungi entomopathogenic JBB 1% 1% 0%
and yeasts
filamentous fungi plantand yeasts
pathogens and yeasts
entomopathogenic 1% ANDES ANDES TOLI 0% 1%
2% ANDES
in Colombia Specimens per fungarium
4% 4% 4% ANDES 1%
0%
ant pathogens 3% 3% 0% and yeasts
entomopathogenic
4%
0% 3% 0% CAUP 1% CAUP 1%
CAUP
ANDES
C
hens, macrofungi and macrofungi
lichens, yeasts 3% macrofungi
lichens,
0%
COAH 2% COAH ANDES 2%COAH 2%
1% 1% COAH CAUP 2%
4%
3% 0% 1% CAUP 2%
lichens, macrofungi 2% COAH 2% 1%
2%
ns, macrofungi 2% 2% microfungi AM 2% COAH 2% 2% 1% 2%
2% yeasts
sts)
nt pathogens
2% (rusts) plant pathogens (rusts) 1% CBUMAG:LIC 2% HT HT HT
HTHEUS
A
plant pathogens (rusts) 0%
Major group collections Fungal collections in Colombia Specimens per fungar
)and macrofungi and macrofungi and macrofungi 1% LLANOS 2%0% 2% 2% HT 2%
6% 6%
filamentous fungi 1%
UPTC-H MEDEL Otros UPTC-H HT UPTC-H 2% UPTC-H
6% plant pathogens entomopathogenic JBB 2% UPTC-H
and yeasts 2% TOLI 0% 2% 2%
UPTC-H HUEB 2% 2%
4% 1% HUEB 1% HUEB 2% HUEB
microfungi AM 3% 0%
2% UDBC
3% 21% 3% HUEB UDBC 21%3%UDBC 2
yeasts lichens, macrofungi ANDES CAUP 3% HUEB CBUMAG:LIC HEUS
1% 3% UDBC
0% 2% COAH 2% 1% 3% 1% LLANOS 0% 21%
2% 1% MEDEL Otros
plant pathogens (rusts) HPUJ HPUJ HPUJ
JBB HPUJ
filamentous fungi TOLI
HPUJ 0% 2% 5%
plant pathogens entomopathogenic and macrofungi HT 5% 5%
HPUJ 5%
1%
and yeasts 5%1%
4%
3% 0% 6% 2% 5% ANDES CAUP
macrofungilichens, macrofungi
macrofungi lichensmacrofungilichens lichens
UPTC-H
MMUNM COAH FAUC2% MMUNM 1%
MMUNM FAUC MMUNM FA
lichens macrofungi
18%
macrofungi
66% 2%18% 66% 18% 66% lichens 66% 2%
HUEB 7% 2% 15%
7% 7% MMUNM FAUC 15% 7% 1
18% lichens 66% UDBC 21% MMUNM 7%
ant pathogens
18% (rusts) 3% HT 15%
66% 7%
and macrofungi 2%
6% CUVC UPTC-H CUVC CUVC
HPUJ 9% 9% CUVC
9% 2% CUVC
5% HUA HUEB HUA 9% HUA
9%
3% COL COL UDBC
macrofungi COL 12% 12% 12% HUA
lichens
FAUC 12% 12% COL
18% MMUNM 12% COL 12%
66%
15% HPUJ 12%
7% 12%
5%
macrofungi CUVC F
18%
lichens
66%
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PercentageHUA Percentage
of
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TULV, 5Percentage 5 collections
UNO,of major group
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COL 12%
11 major group collections
TULV, CM-EM- CM-EM- CM-EM-
CCMhf-UV USAB-CCMhf-UV
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12% EA-EDCMUN CMUN CMUN CCMhf-UV USA
ercentage of major group 5 5collections
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TULV, 2% UDEA 2% BIO MHNMC 2% 1% ColMicroFede-FNC
MHNMC
CM-EM- 1%0% USAB- 2%
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JBB, 11 JBB,
CUVC, UNO, 4CUVC, 5 JBB, 11 CUVC, 5 MHNMC 1% 0%
EA-ED 1%
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JBB, HT, 15 HT, 1 1% CMIUQ CMUN
1%1%0%2% CCMhf-UV
MHNMC 1% 0% 1% UDEA 0% 9% BIO 1% CMIUQ
0% 0% 0% 0%
HT, 1
JBB, 11 ANDES,
CUVC,
UPTC-H, 1 HT, 1 1 9
UPTC-H, UPTC-H, 1 1% UNCSAB 2% 0% MHNMC1%
CMUCMUNCSAB1%
UDEA CMIUQ 0% 1% BIOCMIUQCMUCM
UNCSAB ColMicroFede-FNC
1% CMUCM
1% CMIUQ UNCSAB0% CMUCM 0
0% HUA
NDES, 9 UPTC-H, 1 CUVC,95 ANDES, 9 ANDES, CMPUJ CMPUJ CMPUJ 1% CMIUQ 0% 0% CMPUJ 0%
HT, 1 ANDES, 9 UPTC-H, 1 0% 0% 0%UNCSAB 0% 0%1%
5 IBUN, 15 9 IBUN, 15
ANDES, UPTC-H, 1 IBUN, 15 2% 1% 2% 1% 2%
UNCSAB 1% CMPUJ1% CMUCM
1% 2% 0% COL 0% 12%
IBUN, 15 CMPUJ 0%2% 0%
1% IBUN 12%
IBUN,
CUVC, 1515 CUVC, 15
CUVC, 15
CUVC, 15
MPA-CENIPALMA
IBUN
Percentage of specimens per fungarium
2% IBUN
MPA-CENIPALMA MPA-CENIPALMA
2% 2% IBUNMPA-CENIPALMA
IBUN
pe specimens
2040 2040 2040 2040 12040Total number of25type specimens
Percentage of specimens per culture collection Percentage of s 23%
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FIGURE 2. History and composition of fungal collections in Colombia summarised. A Centre: intensity map showing the Colombian Departments
with the highest concentration of collections. Top left: pie chart depicting the proportion of major life forms found in collections. Bottom
left: pie chart depicting the total number of type specimens per fungarium. Top right: pie chart depicting the proportion of specimens per
fungarium. Bottom right: pie chart depicting the proportion of specimens present in culture collections in Colombia. B Plot depicting the
timeline of the foundation of officially registered fungal collections in Colombia and the date of the first fungal specimen being registered
for each collection.
average of two biological collections were registered per for the project “The Mycological Flora of Colombia”, carried
year in the RNC, with the years 2001 and 2019 holding out by the New York Botanical Garden and the Instituto de
the highest number of registered collections, with 12 and Ciencias Naturales (ICN; Universidad Nacional de Colombia;
8 registrations, respectively. The RNC (2021) reported four 1978-1983), and over 9,000 lichenised fungi gathered as
major registration gaps during 2004–2007, 2009, 2013– part of the binational project ECOANDES, with partners
2014, and 2016. Of the 66 registered fungal collections, from The Netherlands and Colombia (1980–1983); and 5)
only 11 have their databases fully to partially available for the Museo Micológico Universidad Nacional Sede Medellín
download on the Information System on Biodiversity of (MMUNM), with 8,301 specimens, 60% of which represent
Colombia website (SiB, 2021). The remaining collections do phytopathogens, including the Buriticá collection with 3,000
not have their digitised databases available for download on rusts associated with native plant species of Colombia and
the SiB or GBIF websites. duplicates of the phytopathogenic fungi collected by Carlos
Chardón and Rafael Toro in the 1920s, as well as 160 type
Fungal collections of Colombia in numbers specimens. Other important collections are microscope slide
The 234 biological collections recognised by the Registro collections, such as the one hosted by Hongos-Univalle (UV-
MICO) with 2,514 slides. Additionally, the International Center
Nacional de Colecciones (RNC) include over 14 million
for Tropical Agriculture (CIAT) has held for years a collection
records comprising all biological groups. A total of ≈181,000
of 466 arbuscular mycorrhizal fungi, with 12 species
fungi are listed (http://rnc.humboldt.org.co/admin/index.
described from Colombia (Peña-Venegas & Vasco-Palacios,
php/reporte/reporte), comprising dry specimens and strain/
2019), a collection not yet registered with the RNC (García
culture collections. However, this number contains various
et al., 2000).
duplicate entries and some incorrect numbers. After
Colombia has 18 fungal culture collections (Figure
correcting and removing duplicate entries, we reduced the
2D), the largest one being the Colección Nacional de
total to 90,000 specimens, representing only half of the
Fitopatógenos (CNF), which holds 5,295 specimens, followed
original count (see Supplementary Table T1). This illustrates
by the Microorganismos Corporación para Investigaciones
the difficulties in arriving at a clear figure for the number of
Biológicas (MicroCIB), with 2,443 yeasts and filamentous
fungal specimens in the country. How many of these are
fungi, and the Banco de Germoplasma de Microorganismos –
catalogued, that is, digitised, also remains uncertain. For
AGROSAVIA (BGM-A), which hosts 548 accessions including
example, GBIF only contains a little over 15,000 records arbuscular mycorrhizal fungi, yeast, and filamentous fungi,
from Colombian fungaria, most from COL (> 9,000), HPUJ for biological control. The remaining 15 culture collections
(almost 4,000), Unillanos (nearly 600), and ANDES (300). comprise between 500 and a single specimen (Figure 2).
On the other hand, the UDBC collection is fully digitised, but When considering type specimens, we found 565
its data are not yet publicly available. This database includes nomenclatural types housed in Colombian collections
more than 25,000 records, representing at least 10,000 out of at least 989 types that are based on Colombian
more than the number currently available in GBIF. fungal specimens. The fungal collections with the highest
After a thorough review of the data on Colombia’s percentage of nomenclatural types are the Museo
fungaria and ceparia (culture collections), including Micológico (MMUNM), with 160, Herbario Forestal Gilberto
unpublished data provided by curators, we arrived at a Emilio Mahecha Vega Sección No Vasculares (UDBC), with
total of 138,108 fungal accessions deposited in the 66 147, the Herbario Universidad de Antioquia (HUA), with
Colombian collections across the country. This number 91, the Herbario Nacional Colombiano (COL), with 81, the
represents almost 50,000 more specimens than indicated Herbario de la Universidad de Caldas (FAUC), with 30,
by the corrected data from the RNC (Table 1, Figure 2A). and the Universidad Nacional de Colombia Sede Medellín
This difference is partly because 11 of those collections (IBUM), with 15 (Table 1, Figure 2). From the remaining
do not have their specimens registered in the RNC. Out fungal collections in Colombia, only ten hold type specimens
of these 138,108 accessions, 124,930 correspond to dry within their collections.
specimens, 10,414 are strain/culture collections, and 2,764 The Colombian department with the largest number
are microscope slides archived in two collections (Table 1; of fungal collections is Bogotá (Distrito Capital), with 17
Figure 2A). When considering all fungi, including lichens, collections (31%), followed by Antioquia, with nine (16%),
Colombia’s largest collections are: 1) the Herbario Forestal Valle del Cauca, with six (11%), and Caldas and Magdalena,
Gilberto Emilio Mahecha Vega Sección No Vasculares of the with three (5%) each (Figure 2A). Eleven departments
Universidad Distrital Francisco José de Caldas (UDBC), the (Boyacá, Cauca, La Guajira, Huila, Meta, Nariño, Norte de
most important collection of lichenised fungi, with 25,670 Santander, Quindío, Santander, Sucre, and Tolima) each hold
specimens; 2) the herbarium at the Universidad de Caldas a single collection, whereas the 15 remaining departments
(FAUC) with 18,000 specimens of lichenised fungi; 3) the have no fungal collections within their limits (Figure 2A).
fungarium of the Universidad de Antioquia (HUA), with When it comes to the main life forms of fungi represented
the largest collection of macrofungi (13,000 specimens) in Colombian collections, lichens are predominant (66%),
and 2,500 specimens of lichenised fungi; 4) the Herbario followed by macrofungi (16%), yeasts and filamentous
Nacional Colombiano (COL), with over 13,000 specimens, microfungi, such as moulds (13%), and phytopathogenic
including approximately 4,000 macrofungi mostly collected fungi (12%; Figure 2A).
TABLE 1. Colombian biological collections registered in the National Registry of Biological collections (RNC) that contain fungi. Acronym
of herbaria/fungaria, Institution, city, Department, major life forms, number of specimens, number of identified species, curator of the
collection, and origin of the data are indicated (GBIF, 2021). Data not available in GBIF were obtained directly from specialists.
Total Proportion
Type Data
Acronym Institution Department Major groups number identified Fungal curator
specimens origin
specimens species
Strain/culture collections
Arbuscular
25 67% 0
mycorrhiza
Carolina
Yeasts 240 100% 0 Carolina
González
BGM-A AGROSAVIA Cundinamarca González
Filamentous fungi 281 20% 0 Almario,
Almario
RNS
Nematophagous
2 100% 0
fungi
Universidad del Valle del Filamentous Germán A.
CCMhf-UV 46 100% 0 RNC
Valle Cauca fungi, yeasts Bolívar
Carmenza
Federación Gongora
Nacional de Filamentous Botero,
CEN Caldas 222 70% 0 RNC
Cafeteros - fungi, yeasts Luis Miguel
CENICAFÉ Constantino
Chuaire
Aída Vasco
(environmental
CM-EM- Universidad de Filamentous Aída
Antioquia 63 44% 0 fungi), Diana
UDEA Antioquia fungi, yeasts Vasco
González Gil
(clinical fungi)
Universidad del Filamentous
CMIUQ Quindío 40 60% 0 — RNC
Quindío fungi, yeasts
Pontificia Angela María
Filamentous
CMPUJ Universidad Bogotá 177 82% 0 Alvarado RNC
fungi, yeasts
Javeriana Fernández
Universidad
Filamentous
CMUCM Católica de Caldas 15 87% 0 — RNC
fungi, yeasts
Manizales
Universidad de Filamentous
CMUN Nariño 100 100% 0 — RNC
Nariño fungi, yeast
CNF AGROSAVIA Cundinamarca Plant pathogens 5,295 51% 0 — RNC
Federación Diannefair
ColMicro-
Nacional de Santander Filamentous fungi 17 0% 0 Duarte RNC
Fede-FNC
Cacaoteros Hernandez
Universidad
EA-ED Antioquia Filamentous fungi 166 10.5% 0 — RNC
EAFIT
Daniel Uribe
Universidad Velez, Fabio
Filamentous
IBUN Nacional de Bogotá 208 5% 15 Ancizar RNC
fungi, yeasts
Colombia Aristizabal
Gutierrez
TABLE 1. (continued)
Total Proportion
Type Data
Acronym Institution Department Major groups number identified Fungal curator
specimens origin
specimens species
Instituto de
Investigaciones
MHNMC Marinas y Magdalena Filamentous fungi 120 7.5% 0 — RNC
Costeras
Invemar
Universidad
Filamentous
MHN-UPN Pedagógica Bogotá 277 0% 0 Hugo Mauricio RNC
fungi, yeasts
Nacional
Corporación
para Marcela RNC,
Filamentous
MicroCIB Investigaciones Antioquia 2,443 88% 0 Gaviria David
fungi, yeasts
Biológicas Camino Parra
Medellín
Leidy Johanna
Corporación 73 44% 0 Contreras
Centro de Entomopathogenic Arias
MPA- RNC
Investigación Bogotá fungi,
CENIPALMA Yuri Adriana
en Palma de filamentous fungi
Aceite 142 32% 0 Mestizo
Garzón
Universidad
Magally
Nacional de Filamentous
UNCSAB Antioquia 110 85% 0 Romero RNC
Colombia, sede fungi, yeasts
Tabares
Medellin
Luis Alejandro
Universidad de Filamentous
USAB-BIO Bogotá 33 79% 0 Acosta RNC
la Sabana fungi, yeasts
Gonzalez
Valle del Arbuscular
NR CIAT 466 ND 0 — RNC
Cauca mycorrhiza
Fungaria
Fundación Luis
Luis Gerardo
AFP Universitaria de Cauca Lichens 470 ND 0 Gerardo
Chilito
Popayán Chilito
Silvia Restrepo
Universidad de (Levaduras
ANDES Bogotá Macrofungi 2,339 46.2% 1 RNC
los Andes y hongos
microscópicos)
Macrofungi 31 ND 0 María
Fernanda
Lozano y
Fernando Maria
Universidad de
BOG Bogotá Sarmiento Fernando
la Salle Lichens 245 ND 0 Parra, Lozano
Conservador:
José Warles
Díaz
Universidad del
CAUP Cauca Lichens 1,558 6% 0 — RNC
Cauca
TABLE 1. (continued)
Total Proportion
Type Data
Acronym Institution Department Major groups number identified Fungal curator
specimens origin
specimens species
Macrofungi 70 ND 0 Kevin Ramírez
Roncallo,
María A.
Negritto, Jorge
Luna Fontalvo,
Universidad del Kevin
CBUMAG Magdalena Alejandra
Magdalena Lichens 1,300 0% 0 Ramírez
Barrios and
Cristina
Abaunza,
Alberto Rafael
Páez Redondo
Colección de
Ludy
Plantas Vivas Juan Diego
Yaneth
CDMB del Jardín Santander Lichens 69 ND 0 Ramírez
Archila
Botánico Eloy Román
Durán
Valenzuela
Curador
Instituto Wilson Ricardo
COAH Bogotá Lichens 2,362 ND 0 Dairon
SINCHI Álvaro Alba
Cárdenas
Jaime Aguirre Lauren
Lichens 10,901 ND 39
(Líquenes) Raz
Data
Universidad from
COL Nacional de Bogotá other
Colombia sources
Macrofungi 4,000 ND 42 —
ca.
Dumont
et al.
(1978)
Ana
Edier Soto,
Universidad del Valle del Lichens, Cristina
CUVC 9,500 75% 15 Ana Cristina
Valle Cauca macrofungi Bolaños
Bolaños
and RNC
Herbario Lichens 18,000 ND 30 Luis
Luis Fernando
FAUC Universidad de Caldas Fernando
Macrofungi 500 ND 0 Coca
Caldas Coca
Universidad de Norte de
HECASA Lichens 250 4% 0 — RNC
Pamplona Santander
Lichens 513 78.5% 0 — RNC,
Jorge
Universidad de Jorge David
HEUS Sucre David
Sucre Macrofungi 14 100% 0 Mercado Mercado
Gomez Gomez
Facultad de
Agronomía,
HFAB Universidad Bogotá Macrofungi 32 0% 0 — RNC
Nacional de
Colombia
TABLE 1. (continued)
Total Proportion
Type Data
Acronym Institution Department Major groups number identified Fungal curator
specimens origin
specimens species
Universidad de Macrofungi 40 0% 0
HG Guajira — RNC
la Guajira Lichens 25 0% 0
Fundación Manuel David Jairo
HJJT Bogotá Lichens 40 50% 0
Trópico Alto Cortés Pardo Pinto
Pontificia Macrofungi 400 22% 0 Nestor
HPUJ Universidad Bogotá Miguel León-G Julio
Javeriana Lichens 5,800 12% 0 García
Macrofungi 1 0% 1 Favio
Manuel David
Herbario Ávila y
HT Bogotá Cortés y
Tropical Lichens 2,700 90% 0 Astrid
Daniela Jara
Caro
Macrofungi Ana Esperanza
(Ascomycota, Franco, Felipe
Esperanza
Basidiomycota, 12,443 46% 91 Cardona,
Franco
Myxomycetes/ Natalya Gómez
Universidad de Mycetozoa) Montoya
HUA Antioquia
Antioquia Felipe
Margarita
Cardona,
Jaramillo Ciro,
Lichens 2,500 ND 0 Margarita
Giovanny
Jaramillo
Pérez
Ciro
Universidad
HUCO Católica de Antioquia Lichens 71 63% 0 — RNC
Oriente
Lichens 1,900 ND 0
Universidad del Macrofungi Hector Orlando
HUEB Bogotá RNC
Bosque (Ascomycota, 1,922 6% 115 Lancheros
Basidiomycota)
Lichens 13 33% 0 Wilmar
Universidad Valle del Oswaldo
ICESI —
ICESI Cauca Macrofungi 7 0% 0 Diaz
Vasco
Jorge Londoño,
Jardín Botánico
Margarita
Joaquín Antonio Norberto
JAUM Antioquia Lichens 124 ND 0 Jaramillo y
Uribe de López
Juan Carlos
Medellín
Benavides
Jardín Botánico Angélica
JBB Bogotá Lichens 1,430 40% 11 —
de Bogotá Aponte
Jardín Botánico
JBJCM-CV Bogotá Lichens 12 25% 0 — RNC
de Bogotá
Martha Lucía
Macrofungi 509 38% 0
Universidad de Ortiz Luz Stella
LLANOS Meta
los Llanos Mónica Suárez
Lichens 724 10.6% 0
Merchán
TABLE 1. (continued)
Total Proportion
Type Data
Acronym Institution Department Major groups number identified Fungal curator
specimens origin
specimens species
Universidad
MEDEL Nacional de Antioquia Lichens 500 40% 0 — RNC
Colombia
Universidad Plant pathogens Mauricio
Mauricio
MMUNM Nacional Sede, Antioquia (rusts), 8,301 83% 160 Salazar
Salazar Yepes
Medellin macrofungi Yepes
Aida
Universidad de Aida Elena
PSO Nariño Lichens 380 ND 0 Elena
Nariño Baca
Baca
Hilda
Universidad
SURCO Huila Macrofungi 56 ND 0 — Dueñas
Surcolombiana
(Directora)
Universidad del Héctor Héctor
TOLI Tolima Lichens 1,119 ND 0
Tolima Esquivel Esquivel
Instituto para Edier Soto,
la Investigación Martha Ruiz,
y Preservación Edgar Linares,
Alejandro
del Patrimonio Valle del Ana Cristina
TULV Lichens 487 ND 5 Castaño
Cultural y Cauca Bolaños,
Naranjo
Natural del Katherine
Valle del Cauca Osorio, Aura
- Inciva María Rivera
Macrofungi 421 0% 0 Bibiana
Universidad
Moncada,
Distrital Bibiana
UDBC Bogotá Alejandra
Francisco José Lichens 27,320 41% 147 Moncada
Suárez, Diego
de Caldas
Simijaca
Maria
Universidad del
UNO Atlántico Lichens 361 ND 4 Pierine España Cristina
Norte
Martinez
Diego Andrés
Universidad Moreno,
Maria
Pedagógica y Lina Marcelo
UPTC-H Boyacá Lichens 3,054 3.2% 1 Eugenia
Tecnológica de Lozano y
Morales
Colombia UPTC Maria Eugenia
Morales
Universidad del
UTMC Magdalena Lichens 7 57.2% 0 — RNC
Magdalena
Microscope slide collections
Hector Orlando
Universidad del Microfungi
HUEB Bogotá 250 0% 0 Lancheros RNC
Bosque (Zygomycota)
Redondo
María Inés
Álvarez, Luz
Universidad del Valle del Filamentous
UV-MICO 2,514 42.8% 0 Dary Caicedo RNC
Valle Cauca fungi, yeasts
(Micología
Clínica)
FIGURE 3. Colombian fungi in international biological collections. A Intensity map showing the global distribution of fungal species. Pie
charts display the proportion of species belonging to different life forms and the proportion of species per collection. B Intensity map
showing the global distribution of fungal specimens. Inset plot shows the total number of fungi collections and specimens per continent.
Pie charts display the proportion of major life forms.
Fungi of Colombia in international biological collections 2). The herbaria or fungaria with the largest percentage
As mentioned above, Colombian specimens were largely of fungal specimens collected in Colombia are B (38%),
deposited in international collections for a long time. NY (23%), F (5%), and BR, CUP, and S (about 4% each),
Currently, 19,444 specimens and about 2,918 species with the remaining collections accounting for 1% or less of
of fungi collected from Colombia are deposited in 72 Colombian specimens (Table 2, Figure 3). When looking at
international biological collections (GBIF, 2021; Westerdijk species counts, Europe concentrates the highest diversity
Fungal Diversity Centre, 2021). From this total, 37 collections (1,927; 56%), followed by North America (1,354; 39%),
represent fungaria, with 16,988 specimens, and one with the rest of the continents falling below 3%. Country-
ceparium (culture collection), comprising 263 specimens wise, the USA is the country with the highest number of
(Table 2). No records of Colombian samples were found at species counts (1,347; 30%) followed closely by Germany
the Komarov Botanical Institute Culture Collection (LE-BIN) (1,305; 29%), and with much more reduced numbers the
or the Culture Collections Information Worldwide (CCINFO), Netherlands (433; 10%), Belgium (321; 7%), Sweden (291;
part of the World Data Center for Microorganisms (WDCM). 6%); Norway (223; 5%), and the remaining countries with
It is important to note that these figures only include the 3% or less (Figure 3).
available information in databases and are missing data When looking at main life forms represented in
from some crucial collections, such as BAF, which contains international collections, Berlin (B) was the collection with the
some of Singer’s collections. We only obtained secondary largest percentage of lichenised fungi from Colombia (32%),
references on type specimens and the phytopathogenic whereas the New York Botanical Garden (NY) has a varied
collections of Mayor that are in NEU (see Chapter 2). collection with non-lichenised Ascomycota representatives,
When looking at the global distribution of these a substantial number of ectomycorrhizal fungi from oak
international collections, Europe is the continent with forests collected in the 1980s by R. Halling, G. Mueller and
the largest number of fungal specimens originating from A. Franco-Molano, and the Dumont collections made during
Colombia housed outside the country (51%), followed by the “The Mycological Flora of Colombia” project. The most
North America (47%), South America (1%), and Oceania important plant pathogenic collection was found in BPI (15%).
(0.3%) (Figure 3). The USA is the country with the largest When assessing the main life forms globally, lichenised fungi
number of fungal specimens from Colombia (9,079; 42%), dominated the collections (9,641 specimens; 58%), followed
followed by Germany (6,867; 37%), Belgium (674; 3%), by non-lichenised Ascomycota (3,221; 19%), Basidiomycota
Sweden and the Netherlands (662; 3% each), Norway (403; (2,663; 16%), and rusts and smut fungi (478; 3%), with the
2%), and some others with less than 2% (Figure 3, Table remaining forms covering less than 2% (Figure 3).
TABLE 2. International biological collections with fungi of Colombia. Acronym of herbaria/fungaria, institution, country, major fungi groups
found in the collections, number of specimens, number of determined species, percentage of determined species, and number of type
collections are indicated (GBIF, 2021).
Total Proportion
Type
Acronym Institution Country Major groups number identified
specimens
specimens species
Strain/culture collections
Filamentous fungi
The CBS-KNAW culture The
CBS (macro- and microfungi), 264* 99.2% 62
collection Netherlands
yeasts
Fungaria
ASU The Arizona State University USA Lichens 50 98% 1
Lichens, lichenicolous
B Botanischer Garten Berlin Germany 6,534 82.4% 36
fungi
Universidad de Buenos Aires Macrofungi
BAFC Argentina 11 100% 11
Herbario (Basidiomycota)
BCN University of Barcelona Spain Lichens 3 100% 0
BG University of Bergen Norway Lichens 18 94.4% 0
BISH Bishop Museum USA Microfungi (Ascomycota) 1 100% 0
BM(NHMUK) The Natural History Museum UK Lichens 114 95.6% 49
TABLE 2. (continued)
Total Proportion
Type
Acronym Institution Country Major groups number identified
specimens
specimens species
Plant pathogenic,
U.S. National Fungus
BPI (USDA) USA macrofungi, 3,105 87.4% 91
Collections, USDA-ARS
Myxomycetes/Mycetozoa
Lichens, macrofungi
BR Meise Botanic Garden Belgium 670 86.1% 44
(Basidiomycota)
CANB Australian National Herbarium Australia Lichens 33 45.5% 7
CHRB Rutgers University USA Plant pathogens (rusts) 2 100% 0
CMMF Jardin botanique de Montréal Canada Microfungi (Ascomycota) 3 33.3% 0
CNAL Canadian Museum of Nature Canada Lichens 3 100% 0
Cornell Plant Pathology
CUP USA Plant pathogens 657* 8.2% 70
Herbarium
DAR Orange Agricultural Institute Australia Plant pathogens (rusts) 3 33.3% 0
Lichens, macrofungi
DUKE Duke University USA 90 76.7% 0
(Basidiomycota)
Royal Botanic Garden Arbuscular mycorrhyzal
E UK 73 94.8% 47
Edinburgh fungi (BRIL), lichens
ECON Harvard University USA Lichens 1 0% 0
Field Museum of Natural Macrofungi
F USA 913 53% 188
History (Basidiomycota), lichens
plant pathogens (rusts),
FH Harvard University USA 81 92.6% 37
lichens
Universidade Federal de Macrofungi
FLOR Brazil 3 66.7% 0
Santa Catarina (Basidiomycota)
Universidad Nacional
FM-UNAM Autónoma de México (UNAM) Mexico Human pathogenic fungi 14 100% 0
Facultad de Medicina
Senckenberg Gesellschaft für
Naturforschung: Senckenberg
FR Germany Lichens 41 95.1% 0
Forschungsinstitut und
Naturmuseum
Conservatoire et Jardin
G botaniques de la Ville de Switzerland Lichens 98 94.9% 98
Genève
Macrofungi
GB University of Gothenburg Sweden 34 94.1% 0
(Basidiomycota), lichens
GENT Ghent University Belgium Lichens 4 25% 0
Lichens, plant
GZU Karl-Franzens-Universität Graz Austria pathogens 12 100% 0
(Sordariomycetes)
H University of Helsinki Finland Lichens 6 100% 0
Tasmanian Museum and Art
HO Australia Lichens 1 100% 0
Gallery
Universidade Federal do Rio Lichens, macrofungi
ICN Brazil 26 100% 1
Grande do Sul (Basidiomycota)
Plant pathogens
IMI CABI Bioscience UK Centre UK 154 63.6% 0
(Sordariomycetes)
TABLE 2. (continued)
Total Proportion
Type
Acronym Institution Country Major groups number identified
specimens
specimens species
Instituto Nacional de Macrofungi
INPA Brazil 8 87.5% 1
Pesquisas da Amazônia (Basidiomycota)
Macrofungi
K Royal Botanic Gardens, Kew UK 98 94.9% 16
(Basidiomycota)
L Naturalis Netherlands Lichens 261 87.7% 13
LD Lund University Sweden Lichens 30 100% 7
Macrofungi
LIL-Fungi Fundación Miguel Lillo Argentina 2 100% 0
(Basidiomycota)
Macrofungi
LSUM Louisiana State University USA 87 54% 3
(Basidiomycota), lichens
Staatliche
M Naturwissenschaftliche Germany Lichens 291 92.8% 258
Sammlungen Bayerns (SNSB)
Lichens, plant
MA Real Jardín Botánico Spain 29 96.6% 0
pathogens (rusts)
Facultad de Farmacia,
MAF Universidad Complutense, Spain Lichens 2 50% 0
Madrid
MBM Museu Botânico Municipal Brazil Macrofungi (Ascomycota) 1 0% 1
Royal Botanic Gardens
MEL Australia Lichens 15 33.3% 1
Victoria
Universidad Nacional
MEXU Mexico Lichens 15 100% 0
Autónoma de México
Macrofungi
MICH University of Michigan USA 34 97.1% 3
(Basidiomycota)
Macrofungi
MIL Milwaukee Public Museum USA 3 100% 0
(Basidiomycota)
Lichens, plant
MIN University of Minnesota USA 52 92.3% 1
pathogens (rusts)
MN Liceo Classico “Virgilio” Italy Lichens 2 100% 0
MO Missouri Botanical Garden USA Lichens 1 0% 0
MSC Michigan State University USA Lichens 11 81.8% 0
Royal Botanic Gardens &
NSW Australia Lichens 1 0% 0
Domain Trust
Discomycetes,
The New York Botanical
NY USA macrofungi 4,023 69% 132
Garden
(Basidiomycota)
Macrofungi
O University of Oslo Norway 383 86.2% 3
(Basidiomycota), lichens
Muséum National d’Histoire
PC (MNHN) France Lichens 222 84.2% 12
Naturelle
Manaaki Whenua - Landcare
PDD New Zealand Plant pathogens (rusts) 102 97.1% 25
Research
Swedish Museum of Natural Plant pathogens (rusts),
S Sweden 570 87.5% 236
History lichens
TABLE 2. (continued)
Total Proportion
Type
Acronym Institution Country Major groups number identified
specimens
specimens species
Macrofungi
SP-FUNGI Instituto de Botânica IBT Brazil 63 39.7% 0
(Basidiomycota)
Institute of Agricultural and
Environmental Sciences of Macrofungi
TAAM Estonia 16 87.5% 0
the Estonian University of Life (Basidiomycota)
Sciences
Lichens 2 100% 0
TENN University of Tennessee USA Macrofungi
(Basidiomycota), 36 63.9% 1
Sordariomycetes
National Museum of Nature
TNS Japan Lichens 54 98.1% 3
and Science
Norwegian University of
TRH Norway Lichens 2 100% 0
Science and Technology
Macrofungi
TUF (UTE) University of Tartu Estonia 97 25.8% 0
(Basidiomycota), lichens
Myxomycetes/
UARK University of Arkansas USA 56 100% 0
Mycetozoa
Universidad Nacional Federico
UFV Peru Macrofungi (Ascomycota) 1 100% 0
Villarreal
Lichens, macrofungi
UPS Museum of Evolution Sweden 28 89.3% 6
(Basidiomycota)
Universidade Federal de Macrofungi
URM Brazil 40 92.5% 2
Pernambuco (Basidiomycota)
US Smithsonian Institution USA Lichens 497 97.2% 14
Macrofungi
UTC Utah State University USA 34 94.1% 0
(Basidiomycota)
Naturhistorisches Museum
W Austria Lichens 7 71.4% 0
Wien
University of Wisconsin
WIS USA Lichens 93 97.8% 3
Herbarium
the number of accessioned Colombian specimens in a global also be greater than that for other groups of fungi because
database such as GBIF amounts to only ca. 36,000, a they are included in the list of veto species that must be
figure that falls behind those for much smaller, temperate assessed in all environmental impact studies for projects
countries such as Germany (320,000), Japan (200,000), that may affect Colombian ecosystems and biodiversity
France (125,000), the UK (80,000), or Italy (54,000). (Decree 0213 of 1977, INDERENA) (Chapter 14). Finally,
The reasons for the underestimated numbers of fungi other groups that have received disproportionate attention
of Colombia are various (see Chapter 3). On the one hand, are yeasts (Silva-Bedoya et al., 2014; Morales-López et
fungi have not been as extensively studied in Colombia al., 2017, see Chapter 7), rusts (Yepes and González,
as they have been in other countries. This scenario is 2019, see Chapter 9), and smuts (Piepenbring, 2002, see
partly due to Colombia having a relatively small group Chapter 9), due to their importance in biotechnology and
of specialised mycologists, especially when compared agriculture.
with other research areas. On the other hand, many of
the aforementioned Colombian collections (e.g., COL, CONCLUSIONS
MMUNM) and some international collections (e.g., BAF,
Biological collections play a fundamental role in
NEU) are yet to be digitised. These collections may contain
documenting our planet’s biodiversity across space and
unexpected diversity, including species new to science,
time (Pyke & Ehrlich, 2010; Daru et al., 2019; Rønsted et
potentially endemic to Colombia, or species associated
al., 2020). They can be used to reconstruct past and predict
with threatened habitats that deserve special conservation
future biota, for example, due to climate changes. They
status. Uncatalogued collections may also hold lost type
provide the foundation for describing and understanding
specimens. Colombian collections may also add value due
Colombia’s fungal diversity. Their diversity provides a basis
to their broader temporal, geographical and taxonomic
for finding solutions on food security, human health, and
coverage when compared to collections housed at
environmental conservation. Considering that Colombia
international institutions. They may hold a higher proportion
has turned its gaze to the bioeconomy to sustain economic
of specimens collected in ecosystems that are difficult to
development, it must support biological collections as a
access. In addition, these collections have a fundamental
source of information for such development. Therefore,
role in training new generations of Colombian mycologists
it is important to nourish these collections continuously
and in being centres of reference for the development
with further specimens and to provide them with much-
of ecological and environmental research, amongst
needed specialised curators and additional curatorial
other disciplines of mycology in Colombia. Nevertheless,
resources. Combining a well-planned collection digitisation
unrecognised species and otherwise critical collections may
program with the unification of databases, using broadly
also be found among international collections. Increased
accepted taxonomy, will greatly increase the accessibility
cross-institutional communication and collaboration are
of this natural capital. It will allow for comparative studies,
essential to facilitate access to this information and to
metadata analysis, mapping, and modelling species
foster future fungal research, ultimately facilitating more
distributions for biodiversity management, identifying
accurate fungal diversity estimates for Colombia.
knowledge gaps, or proposing solutions to decision-makers
Another issue made obvious from our compilation of
in determining the best compromises between human and
the available fungaria and culture collections within and
environmental needs.
outside Colombia is that Colombia has not been uniformly
surveyed. Gaya et al. (2021) already mentioned that some
economically important areas, such as the ‘Eje Cafetero’ Acknowledgements
or the Andean region, have been more intensively collected
than others. The areas particularly affected by the civil The authors thank Robert Lücking and Viviana Motato for comments
and suggestions that helped improve this manuscript, and the
conflict are the most unexplored to date (e.g., Amazonia and
members of the Colombian Association of Mycology (ASCOLMIC),
Orinoquía). This highlights the need to increase sampling
who provided information relevant for the development of this
efforts in underexplored regions across the Colombian chapter. The authors also thank the following curators who kindly
territory. The recently launched Catalogue of Fungi of provided updates or new data on collections not included in public
Colombia (ColFungi, 2021) represents a good opportunity databases: Angélica Aponte, Ludy Yaneth Archila Durán, Favio
to overcome some of these issues and to further promote Ávila, Aida Elena Baca, Ana Cristina Bolaños, Dairon Cárdenas,
efforts to increase the quantity, quality, and accessibility of Felipe Cardona, Astrid Caro, Alejandro Castaño Naranjo, Luis
fungal collections in Colombia. Gerardo Chilito, Luis Fernando Coca, Wilmar Oswaldo Diaz Vasco,
The fungal group that dominates the hitherto gathered Hilda Dueñas, Héctor Esquivel, Maria Fernando Lozano, Esperanza
and catalogued collections are lichenised fungi. This is an Franco, Nestor Julio García, Carolina González Almario, Margarita
example of research and collection bias and not a reflection Jaramillo Ciro, Norberto López, Maria Cristina Martinez, Maria
of a higher diversity of this functional group in Colombia, Eugenia Morales, Jairo Pinto, Kevin Ramírez, Lauren Raz, Mauricio
Salazar Yepes, and Luz Stella Suárez.
given that many of the studies on Colombian fungi to
date have focused on lichens (Lücking et al., 2021). The
knowledge on and number of collections of lichens might
Thamnolia vermicularis
[Bibiana Moncada]
208 CATALOGUE OF FUNGI OF COLOMBIA
Chapter 16
Annotated Checklist of Fungi of Colombia
Tiziana Antonella Cossu1, Robert Lücking 2,3,4, Natalia Vargas-Estupiñán5,6, Julia Carretero1, Aída M. Vasco-Palacios3,7, Bibiana Moncada2,3,4,8,
Paul Kirk1, Rafael Felipe de Almeida1, Ester Gaya1, Luis Fernando Coca4,9, Joaquim de Souza1, David Díaz-Escandón4,10,11, Ana Esperanza
Franco-Molano12, Nataly Gómez-Montoya12, Fabiola Eugenia González-Cuellar3,13,14, Margarita María Jaramillo-Ciro3,4,15, Viviana Motato-
Vásquez3,16,17, Yeina Milena Niño Fernandez18, Martha Lucia Ortiz-Moreno3,19, Ehidy Rocío Peña-Cañón3,18, Meike Piepenbring20, Mauricio
Ramírez-Castrillón3,21, Mauricio Salazar-Yepes22, Tatiana Sanjuan3,23, Diego Simijaca4,8,24, Edier Soto-Medina4,10 & Mauricio Diazgranados1*
1
Royal Botanic Gardens, Kew, UK.
2
Botanischer Garten und Botanisches Museum, Freie Universität Berlin, Germany.
3
Asociación Colombiana de Micología (ASCOLMIC).
4
Grupo Colombiano de Liquenología (GCOL).
5
Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Bogotá, Colombia.
6
Laboratorio de Micología y Fitopatología, Universidad de los Andes, Bogotá, Colombia.
7
Grupo BioMicro y de Microbiología Ambiental, Colección de Microorganismos, Escuela de Microbiología, Universidad de Antioquia UdeA, Medellín,
Colombia.
8
Licenciatura en Biología, Universidad Distrital Francisco José de Caldas, Bogotá, Colombia.
9
Herbario Universidad de Caldas - FAUC, Manizales, Colombia.
10
Grupo de Ecología y Diversidad Vegetal, Departamento de Biología, Universidad del Valle, Cali, Colombia.
11
Department of Biological Sciences CW405, University of Alberta, Edmonton, Alberta, Canada.
12
Laboratorio de Taxonomía y Ecología de Hongos (TEHO), Instituto de Biología, Facultad de Ciencias Exactas y Naturales, Universidad de Antioquia,
Medellín, Colombia.
13
Departamento de Medicina Interna, Universidad del Cauca, Popayán, Colombia.
14
Semillero de Investigación en Etnobiología (SIE), Universidad del Cauca, Cauca, Colombia.
15
Facultad de Ingeniería, Universidad de San Buenaventura, Medellín, Colombia.
16
Grupo de Investigación en Biología de Plantas y Microorganismos (BPM), Departamento de Biología, Facultad de Ciencias Naturales y Exactas,
Universidad del Valle, Cali, Colombia.
17
Calima, Fundación para la Investigación de la Biodiversidad y Conservación en el Trópico, Cali, Colombia.
18
Línea de Usos y Saberes de Hongos en Andes Nororientales, Grupo de Investigación Biología para la Conservación, Universidad Pedagógica y Tecnológica
de Colombia- Tunja, Colombia.
19
Grupo de Sustentabilidad Ambiental (SUSA), Departamento de Biología y Química, Facultad de Ciencias Básicas e Ingenierías, Universidad de los
Llanos, Villavicencio, Colombia.
20
Mycology Working Group, Goethe University, Frankfurt am Main, Germany.
21
Grupo de Investigación iCUBO, Departamento de Ingeniería Bioquímica, Facultad de Ingeniería, Universidad Icesi, Cali, Colombia.
22
Universidad Nacional de Colombia, sede Medellín, Museo Micológico-MMUNM, Medellín, Colombia.
23
Grupo Micólogos Colombia.
24
Departamento de Biología, Universidad Autónoma de Aguascalientes, Aguascalientes, México.
ABSTRACT
A comprehensive checklist of the fungi known for 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
checklist includes supraspecific taxonomic ranks, accepted species and authors, synonyms and authors, common names,
species origin (when available), geographic information (regions, departments, elevation range), trophic mode, host/
associated species and family (if pertinent), conservation status at global and national levels (assessments of the accepted
name), and level 1 of the category of use. The taxonomic coverage and statistics are provided in Chapter 3. Analysis of
specific groups of fungi can be found in Chapters 4–9. Notes on the geographic distribution are provided in Chapter 10.
Analyses on the conservation status are included in Chapter 14. At the end of the catalogue, indexes of synonyms, families
and genera are also provided. For full details on each species, readers should scan the QR codes or click in the species
hyperlink to visit the species profiles in ColFungi. These include morphological descriptions, geographic distribution and
maps, synonymy, notes on uses, links to fungarium specimens, field images of fungi, illustrations, economic botany items,
bibliography and additional sources.
RESUMEN
En este capítulo se presenta una lista de verificación completa de los hongos 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, sinónimos y autores, nombres comunes, origen de
las especies (si estaba disponible), información geográfica (regiones, departamentos, rango de elevación), modo trófico,
especies y familias hospedantes o en asociación (si aplica), estado de conservación global y nacional (evaluaciones de
riesgo de extinción para nombres aceptados), y nivel 1 de la categoría de uso. La cobertura taxonómica y las estadísticas de
diversidad son presentadas en el Capítulo 3. Análisis de grupos específicos de hongos pueden encontrarse en los capítulos
4–9. En el Capítulo 10 se presenta una descripción general de la distribución geográfica. Los análisis sobre el estado de
conservación de los hongos se incluyen en el Capítulo 14. Al final del catálogo, también se proporcionan índices de sinónimos,
familias y géneros. Para obtener detalles completos sobre la especie, los lectores pueden escanear los códigos QR y visitar
los perfiles de especies en ColFungi que incluyen descripciones morfológicas, distribución geográfica y mapas, sinonimia,
notas sobre usos, enlaces a especímenes de Fungario, imágenes de hongos vivos, ilustraciones, elementos de botánica
económica, bibliografía y fuentes adicionales.
groups in question. Potential homonyms were checked provided also information on additional global assessment
by filtering identical names with different authorship. already revised but not yet published in the IUCN Red List
Correct orthography and author citations were checked by portal. Common names and languages, when available from
automated comparison with a download of all fungal names the original sources, were added to the species. QR codes
from Index Fungorum (2021). For most of these tasks, linking the ColFungi species profiles were created for each
automated formulas in a Microsoft Excel master file were species from its Permanent URLs using the R package
used. From the master file all listed and checked species QRcode version 0.1.3 (Teh, 2015).
records, a master file of currently accepted scientific names
was extracted. 4. Use data and their classification
The subsequent reconciliation of the list of currently Metadata on the use of each species were collected from
accepted scientific names against Index Fungorum was numerous and different scientific resources, details of which
done primarily using Microsoft Access. The full list of are available on bibliography section of each species in
names extracted from the taxonomic backbones mentioned ColFungi.
above was matched with our list linking genus and specific The categorisation of fungal uses followed a revised
epithets. Unmatched names were checked to find the simplified version of the Level 1 states proposed by Cook
underlying reasons, such as potential misspellings in the (1995) in the Economic Botany Data Collections Standard, as
sources, invalidly published names, or names not yet defined in the WCUP (Diazgranados et al., 2020), adapted
registered in Index Fungorum, and errors were corrected for fungi:
in the list, with accompanying updates in Index Fungorum
where applicable. For unmatched names not readily • Animal Food (AF): for vertebrate animals only;
reconciled with the taxonomic backbones, taxonomists
were consulted for advice. Index Fungorum numbers were • Environmental Uses (EU): examples include biofertilizers,
assigned to all names. biocontrol agents, pesticide detoxifiers, wastewater
The reconciled master list of accepted species was purifiers, indicators of the presence of metals, indicators
again carefully analysed by taxonomists experts on specific of the presence of pollution;
fungal groups of Colombia, who provided further corrections • Fuels (FU): e.g., used in production of oil as raw material
and adjustments, including additional information, such for biodiesels, ethanol production, etc.;
as on anamorph names and names with unresolved
• Gene Sources (GS): fungi with beneficial genetic traits e.g.
nomenclatural status.
used in recombinant technology;
3. Annotation of species-related metadata • Human Food (HF): food, including beverages, for humans
Metadata related to the accepted species in the checklist only;
were retrieved from more than 750 scientific resources such • Invertebrate Food (IF): only fungi eaten by invertebrates
as scientific journal publications, books, technical reports, useful to humans, such as silkworms, lac insects and
theses, and databases. The full list of sources related to edible grubs, are covered here;
each species is available in the bibliography of ColFungi.
• Materials (MA): pigments, enzymes with industrial use,
Indications of origin (i.e., endemic, native) were obtained
used in cosmetics;
from the Catalogue of Plants and Lichens of Colombia
(Sipman & Aguirre-C., 2016, 2019) for lichens, and through • Medicines (ME): both human and veterinary;
a literature review for other fungal groups. We also combined • Poisons (PO): fungi that are poisonous to vertebrates
the geographic information in the CPLC with information and non-vertebrates (including non-vertebrate animals,
coming from other scientific sources to determine the plants, bacteria and fungi), both accidentally and
presence of a given species in regions and departments. purposefully, e.g., for hunting and fishing;
Information on trophic mode/guild was extracted mainly
from FUNGuild (Nguyen et al., 2016) and complemented by • Social Uses (SU): fungi used for social purposes,
other scientific resources as scientific journal publications, which are not definable as food or medicines, for
books, technical reports, theses. Host/Associated species instance, masticatories, smoking materials, narcotics,
and families (when pertinent) were extracted from various hallucinogens and psychoactive drugs, contraceptives
scientific sources and compiled with the help of mycologists and abortifacients, and fungi with ritual or religious
with expertise, such as in smuts and rusts. Fungal habit significance (as also proposed by Gruca et al. 2014).
was compiled from different scientific sources. The global
conservation status was retrieved from the IUCN Red List 5. Database normalisation
portal (IUCN, 2020) and matched to the CFC using Microsoft The data were structured, normalised, and formatted
Access, considering only the assessments related to the according to the standards agreed for this catalogue and
accepted name. National assessments were based on data accounting for space limitations. Please see the list of
from the Grupo Colombiano de Liquenología (GCOL), who abbreviations used below.
(14), Diaporthales (32), Diversisporales (51), Dothideales (46), Blastocladiaceae (2), Bolbitiaceae (10), Boletaceae
(9), Echinosteliida (1), Entomophthorales (6), Entylomatales (43), Boletinellaceae (5), Botryobasidiaceae (6), Botry-
(9), Eremithallales (6), Erythrobasidiales (1), Eurotiales osphaeriaceae (16), Brigantiaeaceae (1), Bulleribasidi-
(136), Exobasidiales (2), Filobasidiales (9), Geastrales (8), aceae (2), Cainiaceae (1), Calcarisporiaceae (1), Calici-
Geoglossales (4), Georgefischeriales (4), Gloeophyllales aceae (75), Callistosporiaceae (3), Calostomataceae (1),
(3), Glomerales (45), Glomerellales (36), Gomphales (19), Candelariaceae (8), Capnodiaceae (6), Castanediellaceae
Graphidales (79), Gyalectales (124), Helicobasidiales (1), (2), Catillariaceae (3), Celotheliaceae (3), Cenangiaceae
Helotiales (187), Herpomycetales (1), Hymenochaetales (4), Ceratiomyxidae (2), Ceratobasidiaceae (3), Cerato-
(109), Hypocreales (405), Incertae sedis (163), cystidaceae (7), Cerrenaceae (3), Chaconiaceae (7),
Laboulbeniales (16), Lauriomycetales (1), Lecanorales Chaetomiaceae (17), Chaetosphaeriaceae (5), Chaeto-
(824), Lecideales (12), Leotiales (10), Lepidostromatales thyriaceae (2), Chlorociboriaceae (2), Choanephoraceae
(3), Leprocaulales (1), Liceida (7), Lichenoconiales (2), (1), Chromocyphellaceae (4), Chrysotrichaceae (5), Chytri-
Lichenostigmatales (4), Lichinales (1), Lineolatales (1), diaceae (1), Cladoniaceae (77), Cladosporiaceae (16),
Magnaporthales (3), Malasseziales (3), Meliolales (27), Claroideoglomeraceae (5), Clastodermatidae (1), Clavari-
Microascales (25), Microbotryales (6), Microthyriales (12), aceae (5), Clavicipitaceae (27), Coccocarpiaceae (13),
Monoblastiales (13), Mortierellales (26), Mucorales (40), Coccodiniaceae (1), Coccotremataceae (1), Cochlearomy-
Muyocopronales (1), Mycocaliciales (5), Mycosphaerellales cetaceae (2), Coelomomycetaceae (1), Coenogoniaceae
(1), Myriangiales (6), Mytilinidiales (2), Nephridiophagales (47), Coleosporiaceae (7), Collemataceae (56), Coni-
(1), Odontotrematales (1), Olpidiales (1), Onygenales (21), ochaetaceae (6), Coniocybaceae (7), Coniophoraceae (1),
Ophiostomatales (4), Orbiliales (11), Ostropales (506), Coniothyriaceae (2), Cordanaceae (1), Cordieritidaceae
Paraglomerales (3), Parasympodiellales (1), Patellariales (8), Cordycipitaceae (38), Corticiaceae (12), Cortinari-
(2), Peltigerales (256), Peronosporales (23), Pertusariales aceae (21), Coryneliaceae (2), Corynesporascaceae (3),
(58), Pezizales (33), Phacidiales (3), Phaeomoniellales Crepidotaceae (14), Cribrariidae (5), Cronartiaceae (4),
(3), Phallales (12), Phomatosporales (1), Phyllachorales Cryphonectriaceae (7), Cryptobasidiaceae (1), Cryptococ-
(84), Physarida (40), Plasmodiophorida (2), Platygloeales caceae (9), Cucurbitariaceae (2), Cuniculitremaceae (2),
(4), Pleosporales (150), Pneumocystales (1), Polyporales Cunninghamellaceae (8), Cylindriaceae (1), Cyphellaceae
(258), Protostelida (2), Pucciniales (554), Pyrenulales (2), Cyphellophoraceae (1), Cyphobasidiaceae (1), Cysto-
(75), Rhipidiales (1), Rhizocarpales (11), Rhizophlyctidales basidiaceae (3), Cystocoleaceae (1), Cystostereaceae (1),
(1), Rhizophydiales (1), Rhytismatales (14), Russulales Dacampiaceae (2), Dacrymycetaceae (14), Dactylo-
(90), Saccharomycetales (105), Saccharomycodaceae sporaceae (1), Davidiellaceae (1), Debaryomycetaceae
(1), Schaereriales (1), Schizosaccharomycetales (1), (3), Dianematidae (2), Diaporthaceae (16), Diatrypaceae
Sebacinales (3), Septobasidiales (2), Sordariales (36), (5), Didymellaceae (24), Didymiidae (16), Didymospha-
Sporidiobolales (9), Stemonitida (31), Stereopsidales eriaceae (6), Diplocystidiaceae (2), Dipodascaceae (6),
(2), Strigulales (23), Symbiotaphrinales (1), Taphrinales Discinellaceae (4), Diversisporaceae (7), Dothideaceae
(1), Teloschistales (51), Thelebolales (7), Thelenellales (2), Dothidotthiaceae (1), Drepanopezizaceae (4), Eballis-
(6), Thelephorales (17), Tilletiales (6), Togniniales (2), traceae (2), Echinostelidae (1), Ectolechiaceae (80),
Trechisporales (20), Tremellales (40), Triblidiales (1), Elaphomycetaceae (2), Elsinoaceae (6), Endomelanconi-
Trichiida (22), Trichosporonales (16), Trypetheliales (102), opsidaceae (2), Englerulaceae (2), Entolomataceae (27),
Tubeufiales (5), Umbelopsidales (7), Umbilicariales (11), Entomophthoraceae (4), Entylomataceae (9), Eocronarti-
Urocystidales (6), Ustilaginales (54), Valsariales (1), aceae (2), Epibryaceae (2), Erysiphaceae (12), Erythroba-
Venturiales (6), Verrucariales (15), Vezdaeales (1), Xylariales sidiaceae (1), Etheirophoraceae (1), Euantennariaceae
(126), Xylobotryales (1). (1), Exobasidiaceae (1), Fibroporiaceae (1), Filobasidi-
aceae (4), Fomitopsidaceae (9), Fuscideaceae (3), Gano-
Families: Abrothallaceae (8), Acarosporaceae (6), Acaulo- dermataceae (1), Geastraceae (8), Gelatinodiscaceae (4),
sporaceae (21), Acrospermaceae (1), Agaricaceae (43), Geoglossaceae (4), Georgefischeriaceae (1), Gigaspo-
Ajellomycetaceae (3), Albatrellaceae (1), Albuginaceae (7), raceae (23), Gloeophyllaceae (3), Glomeraceae (40),
Amanitaceae (27), Amaurochaetidae (6), Ambisporaceae Glomerellaceae (29), Glomosporiaceae (3), Gomphaceae
(4), Amniculicolaceae (4), Amphisphaeriaceae (9), Amylo- (19), Gomphillaceae (79), Graphidaceae (486), Graphos-
corticiaceae (3), Ancylistaceae (2), Anthracoideaceae tromataceae (13), Grifolaceae (1), Gymnosporangiaceae
(24), Aphelariaceae (1), Apiosporaceae (3), Apoharknessi- (1), Gyroporaceae (2), Haematommataceae (10), Halo-
aceae (1), Arachnopezizaceae (2), Archaeosporaceae (3), sphaeriaceae (7), Hamatocanthoscyphaceae (1), Hansfor-
Arthoniaceae (87), Arthopyreniaceae (1), Arthrodermata- diaceae (1), Harknessiaceae (1), Helicobasidiaceae (1),
ceae (12), Arthrorhaphidaceae (2), Ascobolaceae (5), Helicogoniaceae (1), Helotiaceae (17), Helvellaceae (3),
Aspergillaceae (132), Asterinaceae (22), Asterotexaceae Hericiaceae (3), Herpomycetaceae (1), Herpotrichiellaceae
(1), Astrosphaeriellaceae (2), Atheliaceae (9), Auriculari- (23), Hyaloriaceae (2), Hyaloscyphaceae (5), Hydnaceae
aceae (18), Auriscalpiaceae (4), Backusellaceae (1), Baeo- (40), Hydnangiaceae (10), Hydnodontaceae (20),
mycetaceae (7), Bankeraceae (5), Bartaliniaceae (1), Hygrophoraceae (105), Hymenochaetaceae (65), Hyme-
Basidiobolaceae (1), Beltraniaceae (1), Bionectriaceae nogastraceae (56), Hyphodermataceae (14), Hypoc-
reaceae (78), Hypoxylaceae (36), Icmadophilaceae (15), aceae (2), Pyronemataceae (10), Pythiaceae (7), Radulo-
Incertae sedis (462), Incrustoporiaceae (3), Induratiaceae mycetaceae (3), Ramalinaceae (129), Ramboldiaceae (3),
(1), Inocybaceae (9), Irpicaceae (10), Laboulbeniaceae Raveneliaceae (18), Reticulariidae (4), Rhizocarpaceae
(16), Lachnaceae (50), Laetiporaceae (3), Lamproderma- (11), Rhizophlyctidaceae (1), Rhizopodaceae (3), Rhizopo-
tidae (5), Lasiosphaeriaceae (5), Lauriomycetaceae (1), gonaceae (1), Rhynchogastremaceae (6), Rhytismataceae
Lecanographaceae (10), Lecanoraceae (49), Lecideaceae (14), Rickenellaceae (8), Robillardaceae (1), Roccellaceae
(12), Leotiaceae (4), Lepidostromataceae (3), Leprocaul- (49), Roccellographaceae (2), Russulaceae (46), Rutstro-
aceae (1), Leptodontidiaceae (1), Leptosphaeriaceae (4), emiaceae (11), Saccharomycetaceae (35), Saccharomyco-
Letrouitiaceae (4), Liceidae (3), Lichenoconiaceae (2), daceae (5), Saccotheciaceae (7), Saksenaeaceae (6),
Lichtheimiaceae (2), Lineolataceae (1), Lophiostomata- Salispinaceae (1), Sarcoscyphaceae (6), Sarcosomata-
ceae (1), Lycoperdaceae (18), Lyophyllaceae (5), Lyrom- ceae (2), Schaereriaceae (1), Schizoparmaceae (1),
mataceae (1), Macrocystidiaceae (1), Macrovalsariaceae Schizophyllaceae (4), Schizoporaceae (26), Schizosaccha-
(1), Magnaporthaceae (1), Malasseziaceae (3), Malmide- romycetaceae (1), Sclerodermataceae (8), Sclerotiniaceae
aceae (23), Marasmiaceae (100), Marthamycetaceae (3), (13), Sebacinaceae (3), Septobasidiaceae (2), Skierkaceae
Massarinaceae (3), Megalosporaceae (7), Megasporaceae (1), Solenopeziaceae (2), Sordariaceae (5), Sparassi-
(1), Melampsoraceae (3), Melanommataceae (3), daceae (1), Sphaerophoraceae (3), Sphaer-ophragmiaceae
Melaspileaceae (6), Meliolaceae (27), Meripilaceae (6), (2), Sphinctrinaceae (1), Spirographaceae (3), Sporidiobol-
Meruliaceae (12), Metschnikowiaceae (5), Microascaceae aceae (9), Sporormiaceae (2), Stachybotryaceae (4), Stec-
(10), Microbotryaceae (4), Micropeltidaceae (1), Micro- cherinaceae (17), Stemonitidae (4), Stephanosporaceae
sphaeropsidaceae (1), Microtheliopsidaceae (1), Micro- (2), Stereaceae (20), Stereocaulaceae (30), Stereopsi-
thyriaceae (11), Mikronegeriaceae (3), Milesinaceae (5), daceae (2), Stictidaceae (17), Strigulaceae (23), Stropha-
Mniaeciaceae (1), Mollisiaceae (9), Monascaceae (2), riaceae (26), Suillaceae (2), Symbiotaphrinaceae (1),
Monoblastiaceae (13), Morchellaceae (2), Mortierellaceae Syncephalastraceae (3), Synchytriaceae (4), Taphrinaceae
(26), Mrakiaceae (1), Mucoraceae (15), Muyocopronaceae (1), Telimenaceae (3), Teloschistaceae (39), Tephrome-
(1), Mycenaceae (70), Mycocaliciaceae (4), Mycoporaceae lataceae (3), Teratosphaeriaceae (7), Tetraplospha-
(4), Mycosphaerellaceae (136), Mycosyringaceae (1), eriaceae (1), Thelenellaceae (6), Thelephoraceae (12),
Mytilinidiaceae (2), Myxotrichaceae (7), Nectriaceae Tilletiaceae (6), Tilletiariaceae (1), Togniniaceae (2), Toru-
(144), Neolauriomycetaceae (1), Niaceae (8), Niessliaceae laceae (2), Tranzscheliaceae (2), Trapeliaceae (10), Trem-
(4), Nitschkiaceae (3), Ochrolechiaceae (4), Odontotrem- atosphaeriaceae (1), Tremellaceae (16), Triblidiaceae (1),
ataceae (1), Oliveoniaceae (1), Olpidiaceae (1), Omphalo- Trichiidae (20), Tricholomataceae (8), Trichomeriaceae (3),
taceae (67), Onygenaceae (4), Opegraphaceae (28), Ophi- Trichomonascaceae (9), Trichosporonaceae (16), Tricladi-
ocordycipitaceae (42), Ophioparmaceae (2), Ophiostoma- aceae (1), Trimorphomycetaceae (2), Trypetheliaceae
taceae (4), Orbiliaceae (11), Oxyporaceae (1), Panaceae (100), Tubariaceae (1), Tuberaceae (1), Tubeufiaceae (5),
(10), Pannariaceae (36), Paraglomeraceae (3), Parasym- Tulasnellaceae (1), Tympanidaceae (1), Typhulaceae (1),
podiellaceae (1), Parmeliaceae (411), Parodiellaceae (2), Umbelopsidaceae (7), Umbilicariaceae (6), Uncolaceae
Parodiopsidaceae (4), Patellariaceae (2), Paxillaceae (2), (3), Urocystidaceae (2), Uropyxidaceae (12), Ustilagi-
Peltigeraceae (151), Peltulaceae (1), Peniophoraceae (14), naceae (30), Ustilentylomataceae (2), Valsaceae (3),
Perisporiopsidaceae (2), Peronosporaceae (16), Pertusari- Valsariaceae (1), Varicellariaceae (2), Venturiaceae (6),
aceae (35), Pestalotiopsidaceae (19), Pezizaceae (4), Verrucariaceae (15), Vezdaeaceae (1), Vibrisseaceae (1),
Pezizellaceae (9), Phacidiaceae (2), Phaeococcomyceta- Vizellaceae (1), Wickerhamomycetaceae (4), Xenasmata-
ceae (4), Phaeodimeriellaceae (1), Phaeosphaeriaceae ceae (6), Xylariaceae (66), Xylobotryaceae (1).
(10), Phakopsoraceae (43), Phallaceae (12), Phanero-
chaetaceae (16), Phleogenaceae (4), Phlyctidaceae (4),
Phomatosporaceae (1), Phragmidiaceae (11), Phragmoxe- References
nidiaceae (1), Phyllachoraceae (81), Phyllostictaceae (1), Bernal, R., Gradstein, S.R. and Celis, M. (eds.) (2016a). Catálogo de Plantas
Phyllotopsidaceae (2), Physalacriaceae (26), Physaridae y Líquenes de Colombia. Vol. I: Líquenes a Lythraceae. Bogotá: Instituto
(40), Physciaceae (88), Pilobolaceae (1), Piskurozymaceae de Ciencias Naturales, Universidad Nacional de Colombia, 993 pp.
Bernal, R., Gradstein, S.R. and Celis, M. (eds.) (2016b). Catálogo de
(5), Planistromellaceae (2), Plasmodiophoridae (2), Platy- Plantas y Líquenes de Colombia. Vol. II: Magnoliaceae a Zygophyllaceae
gloeaceae (2), Plectosphaerellaceae (7), Pleosporaceae – Especies Introducidas y Cultivadas. Bogotá, Instituto de Ciencias
(64), Pleurotaceae (27), Ploettnerulaceae (2), Pluteaceae Naturales, Universidad Nacional de Colombia, 1,556 pp.
(15), Pneumocystaceae (1), Podoscyphaceae (12), Bernal, R., Gradstein, S. & Celis, M. (eds.) (2019). Catálogo de
Plantas y Líquenes de Colombia. Bogotá: Instituto de Ciencias
Podosporaceae (2), Polycoccaceae (2), Polyporaceae Naturales, Universidad Nacional de Colombia. Available at http://
(121), Polystomellaceae (1), Porinaceae (72), Porothe- catalogoplantasdecolombia.unal.edu.co.
leaceae (1), Protoscyphaceae (1), Psathyrellaceae (22), ColFungi (2021). Useful Fungi of Colombia. Facilitated by the Royal Botanic
Pseudeurotiaceae (7), Pseudoperisporiaceae (3), Pseu- Gardens, Kew. Published on the Internet; https://colfungi.org/
Cook, F. (1995). Economic Botany Data Collection Standards, Prepared
dopyrenochaetaceae (1), Psilolechiaceae (1), Psoraceae for the International Working Group on Taxonomic Database for Plant
(1), Pterulaceae (5), Pucciniaceae (316), Pucciniastraceae Science (TDWG). Royal Botanic Gardens, Kew.
(9), Pucciniosiraceae (13), Pyrenulaceae (75), Pyriculari- Diazgranados, M., Allkin, B., Black, N., Cámara-Leret, R., Canteiro, C.,
Carretero, J., Eastwood, R., Hargreaves, S., Hudson, A., Milliken, W.,
Nesbitt, M., Ondo, I., Patmore, K., Pironon, S., Turner, R. & Ulian,
T. (2020). World Checklist of Useful Plant Species. In: Royal Botanic
Gardens, K. (ed.). Knowledge Network for Biocomplexity. Available
online: https://knb.ecoinformatics.org/view/doi:10.5063/F1CV4G34.
GBIF (2021). GBIF: Global Biodiversity Information Facility. Facilitated by
the Global Biodiversity Information Facility. Published on the Internet;
https://www.gbif.org/.
Gruca, M., Cámara-Leret, R., Macía, M.J., & Balslev, H. (2014). New
categories for traditional medicine in the Economic Botany Data
Collection Standard. Journal of Ethnopharmacology, 155(2), 1388–
1392. https://doi.org/10.1016/j.jep.2014.06.047
Index Fungorum (2021). Index Fungorum. Facilitated by the Index Fungorum
Partnership. Published on the Internet; http://www.indexfungorum.
org/.
IUCN (2020). The IUCN Red List of Threatened Species. Version 2021-1.
Published on the Internet; http://www.iucnredlist.org/. Downloaded 24
August 2021.
Nguyen, N.H., Song, Z., Bates, S.T., Branco, S., Tedersoo, L., Menke, J.,
Schilling, J.S. and Kennedy, P.G., (2016). FUNGuild: an open annotation
tool for parsing fungal community datasets by ecological guild. Fungal
Ecology, 20, 241–248.
Piepenbring, M. (2002). Annotated checklist and key for smut fungi in
Colombia [Lista anotada y clave para los ustilaginales de Colombia].
Caldasia, 24(1), 103-119.
SIB Colombia (2021). Sistema de Información sobre Biodiversidad de
Colombia. Published on the Internet; https://datos.biodiversidad.co/.
Sipman, H. J. M. & Aguirre-C., J. (2016) Líquenes. In: Bernal, R., Gradstein,
S. R. & Celis, M. Catálogo de Plantas y Líquenes de Colomia. Volumen
1. Universidad Nacional de Colombia, Facultad de Ciencias, Instituto
de Ciencias Naturales, Bogotá: 159-281.
Species Fungorum (2021). Species Fungorum. Facilitated by the Index
Fungorum Partnership. Published on the Internet; http://www.
speciesfungorum.org/.
Teh, V. (2015). QRcode Generator for R (0.1.3) [Online]. https://doi.
org/10.5281/zenodo.5040088. [Accessed].
Tropicos (2021). Tropicos.org. Facilitated by the Missouri Botanical Garden.
Published on the Internet; https://www.tropicos.org/.
Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L.,
Herendeen, P.S., Knapp, S., Kusber, W.-H., Li, D.-Z., Marhold, K., May,
T.W., McNeill, J., Monro, A.M., Prado, J., Price, M.J. & Smith, G.F.
(eds.) (2018). International Code of Nomenclature for Algae, Fungi, and
Plants (Shenzhen Code). Regnum Vegetabile 159. Glashütten: Koeltz
Botanical Books. DOI https://doi.org/10.12705/Code.2018
Fungi, Blastocladiomycota, Incertae sedis, Fungi, Blastocladiomycota, Incertae sedis, Fungi, Blastocladiomycota, Incertae sedis,
Blastocladiomycetes, Incertae sedis, Blastocladiomycetes, Incertae sedis, Blastocladiomycetes, Incertae sedis,
Blastocladiales, Blastocladiaceae Blastocladiales, Blastocladiaceae Blastocladiales, Coelomomycetaceae
1. Blastocladiella colombiensis Karling IF 2. Clavochytridium colombiense Doweld IF 3. Coelomomyces reticulatus Couch & A.J.
No: 103090 Trophic mode/Guild: No: 550342 Walker IF No: 114691 Trophic
saprotroph/undefined saprotroph mode/Guild: pathotroph/animal pathogen
Fungi, Chytridiomycota, Chytridiomycotina, Fungi, Chytridiomycota, Chytridiomycotina, Fungi, Chytridiomycota, Incertae sedis,
Chytridiomycetes, Chytridiomycetidae, Rhizophydiomycetes, Incertae sedis, Incertae sedis, Incertae sedis,
Rhizophlyctidales, Rhizophlyctidaceae Rhizophydiales, Incertae sedis Nephridiophagales, Incertae sedis
10. Karlingia lobata Karling 11. Batrachochytrium dendrobatidis 12. Nephridiophaga lucihormetica R.
IF No: 287362 Longcore, Pessier & D.K. Nichols IF No: Radek, Wellmanns & A. Wolf
450228 Trophic mode/Guild: pathotroph/ IF No: 543141
animal pathogen
Fungi, Ascomycota, Pezizomycotina, Fungi, Ascomycota, Pezizomycotina, Fungi, Ascomycota, Pezizomycotina, Sordariomycetes,
Sordariomycetes, Xylariomycetidae, Sordariomycetes, Xylariomycetidae, Xylariomycetidae, Xylariales, Xylariaceae
Xylariales, Induratiaceae Xylariales, Robillardaceae 4620. Dematophora acutispora (Theiss.) C. Lamb., Wittstein & M.
4618. Induratia suturae (Kudalkar, Strobel 4619. Robillarda sessilis (Sacc.) Sacc. IF Stadler IF No: 827530 Trophic mode/Guild: saprotroph
& Riy.–Ul–Hass.) Samarak., Thongbai, K.D. No: 225336 /undefined saprotroph
Hyde & M. Stadler IF No: 833463 Trophic
mode/Guild: saprotroph/ undefined
saprotroph
Fungi, Basidiomycota, Agaricomycotina, Fungi, Basidiomycota, Incertae sedis, Fungi, Basidiomycota, Incertae sedis,
Agaricomycetes, Incertae sedis, Incertae Incertae sedis, Incertae sedis, Incertae Incertae sedis, Incertae sedis, Incertae
sedis, Incertae sedis sedis, Incertae sedis sedis, Incertae sedis
7078. Odonticium helgae Hjortstam & 7079. Brevicellopsis allantospora 7080. Radulodontia pyriformis Hjortstam
Ryvarden IF No: 103621 Trophic mode (Hjortstam & Ryvarden) Hjortstam & & Ryvarden IF No: 507302 Trophic mode/
/Guild: saprotroph/undefined saprotroph Ryvarden IF No: 507301 Trophic mode Guild: saprotroph/undefined saprotroph
Habitat: Saprotroph Distribution: Europe, /Guild: saprotroph/ undefined saprotroph
South America Elev.: 500 m Dept.: MAG Dept.: MAG
Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae
sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis,
Incertae sedis Incertae sedis Incertae sedis
7081. Asterinites colombiensis Doub. & D. 7082. Asterinites tellezii Doub. & D. Pons 7083. Dicellaesporites magnus Doub. & D.
Pons ex Kalgutkar & Janson. IF No: ex Kalgutkar & Janson. IF No: 483253 Pons IF No: 485252 Trophic mode/Guild:
483252 Trophic mode/Guild: fossil Fungi Trophic mode/Guild: fossil Fungi fossil Fungi
Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae
sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis,
Incertae sedis Incertae sedis Incertae sedis
7084. Diporisporites elongatus Hammen 7085. Diporisporites minutiporatus 7086. Diporisporites minutus Hammen IF
IF No: 330110 Trophic mode/Guild: fossil Hammen IF No: 330111 Trophic mode/ No: 330112 Trophic mode/Guild: fossil
Fungi Guild: fossil Fungi Fungi
Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae
sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis,
Incertae sedis Incertae sedis Incertae sedis
7087. Inapertisporites minutus Hammen 7088. Inapertisporites typicus Hammen IF 7089. Inapertisporites variabilis Hammen
IF No: 332523 Trophic mode/Guild: fossil No: 332524 Trophic mode/Guild: fossil IF No: 332525 Trophic mode/Guild:
Fungi Fungi fossilFungi/
Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae
sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis,
Incertae sedis Incertae sedis Incertae sedis
7090. Incertisporites polygranulatus 7091. Lacrimasporonites arcuatus Doub. 7092. Lacrimasporonites ovalis Doub. & D.
Hammen IF No: 332526 Trophic mode/ & D. Pons IF No: 485259 Trophic mode/ Pons IF No: 485260 Trophic mode/Guild:
Guild: fossil Fungi Guild: fossil Fungi fossil Fungi
Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae
sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis,
Incertae sedis Incertae sedis Incertae sedis
7093. Mediaverrunites fournieri Elsik & 7094. Mediaverrunites magnus Elsik & 7095. Monoporisporites annulatus
Jarzen IF No: 568010 Trophic mode/Guild: Jarzen IF No: 568012 Trophic mode/Guild: Hammen IF No: 334446 Trophic mode/
fossil Fungi fossil Fungi Guild: fossil Fungi
Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae
sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis,
Incertae sedis Incertae sedis Incertae sedis
7096. Monoporisporites buerglii Hammen 7097. Monoporisporites grandis Hammen 7098. Monoporisporites minutus Hammen
IF No: 334447 Trophic mode/Guild: fossil IF No: 334449 Trophic mode/Guild: fossil IF No: 334450 Trophic mode/Guild: fossil
Fungi Fungi Fungi
Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae
sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis,
Incertae sedis Incertae sedis Incertae sedis
7099. Multicellaesporites cerrejonensis 7100. Pluricellaesporites erdtmanii 7101. Pluricellaesporites filiformis
Doub. & D. Pons Hammen IF No: 337279 Trophic mode/ Hammen IF No: 337280 Trophic mode/
IF No: 485250 Guild: fossil Fungi Guild: fossil Fungi
Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae
sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis,
Incertae sedis Incertae sedis Incertae sedis
7102. Pluricellaesporites krauselii 7103. Pluricellaesporites minutigranulatus 7104. Pluricellaesporites typicus Hammen
Hammen IF No: 337282 Trophic mode/ Hammen IF No: 337283 Trophic mode/ IF No: 337287 Trophic mode/Guild: fossil
Guild: fossil Fungi Guild: fossil Fungi Fungi
Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae
sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis,
Incertae sedis Incertae sedis Incertae sedis
7105. Polyadosporites firbasii Hammen IF 7106. Polyadosporites garciabarrigae 7107. Polyadosporites suescae Hammen
No: 337454 Trophic mode/Guild: fossil Hammen IF No: 337455 Trophic mode/ IF No: 337456 Trophic mode/Guild: fossil
Fungi Guild: fossil Fungi Fungi
Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae
sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis,
Incertae sedis Incertae sedis Incertae sedis
7108. Polyporisporites elongatus Hammen 7109. Spirotremesporites duenasii Elsik IF 7110. Spirotremesporites ecuatorialis
IF No: 337477 Trophic mode/Guild: fossil No: 130384 Trophic mode/Guild: fossil Dueñas IF No: 112635 Trophic mode/
Fungi Fungi Guild: fossil Fungi
Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae
sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis,
Incertae sedis Incertae sedis Incertae sedis
7111. Spirotremesporites multiplex 7112. Spirotremesporites simplex Dueñas 7113. Striadyadosporites constrictus
Dueñas IF No: 112636 Trophic mode/ IF No: 112637 Trophic mode/Guild: fossil Dueñas IF No: 112047 Trophic mode/
Guild: fossil Fungi Fungi Guild: fossil Fungi
Fungi, Fossil Fungi, Incertae sedis, Incertae Fungi, Fossil Fungi, Incertae sedis, Incertae Protozoa, Amoebozoa, Mycetozoa,
sedis, Incertae sedis, Incertae sedis, sedis, Incertae sedis, Incertae sedis, Myxogastrea, Columellinia,
Incertae sedis Incertae sedis Clastodermatida, Clastodermatidae
7114. Striadyadosporites elongatus 7115. Triporisporites minutus Hammen IF 7116. Clastoderma debaryanum A. Blytt
Dueñas IF No: 112651 Trophic mode/ No: 340461 Trophic mode/Guild: fossil IF No: 248890
Guild: fossil Fungi Fungi
Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis,
Peronosporea, Albuginidae, Albuginales, Peronosporea, Albuginidae, Albuginales, Peronosporea, Albuginidae, Albuginales,
Albuginaceae Albuginaceae Albuginaceae
7231. Albugo candida (Pers. ex J.F. Gmel.) 7232. Albugo chardonii W. Weston IF No: 7233. Albugo ipomoeae–panduratae
Roussel IF No: 122113 Trophic mode/ 534448 Trophic mode/Guild: pathotroph (Schwein.) Swingle IF No: 120694 Trophic
Guild: pathotroph/plant pathogen /plant pathogen mode/ Guild: pathotroph/plant pathogen
Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis,
Peronosporea, Albuginidae, Albuginales, Peronosporea, Albuginidae, Albuginales, Peronosporea, Albuginidae, Albuginales,
Albuginaceae Albuginaceae Albuginaceae
7234. Pustula tragopogonis (Pers.) Thines 7235. Wilsoniana bliti (Biv.) Thines 7236. Wilsoniana platensis (Speg.) Thines
IF No: 346454 IF No: 345130 IF No: 345146
Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis,
Peronosporea, Albuginidae, Albuginales, Peronosporea, Peronosporidae, Peronosporea, Peronosporidae,
Albuginaceae Peronosporales, Peronosporaceae Peronosporales, Peronosporaceae
7237. Wilsoniana portulacae (DC.) Thines 7238. Halophytophthora vesicula 7239. Peronosclerospora sorghi (W.
IF No: 345084 (Anastasiou & Churchl.) H.H. Ho & S.C. Weston & Uppal) C.G. Shaw
Jong IF No: 126021 IF No: 319377
Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis,
Peronosporea, Peronosporidae, Peronosporea, Peronosporidae, Peronosporea, Peronosporidae,
Peronosporales, Peronosporaceae Peronosporales, Peronosporaceae Peronosporales, Peronosporaceae
7240. Peronospora dianthicola R. Heim 7241. Peronospora farinosa (Fr.) Fr. 7242. Peronospora sparsa Berk.
IF No: 508269 IF No: 232886 IF No: 143418
Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis,
Peronosporea, Peronosporidae, Peronosporea, Peronosporidae, Peronosporea, Peronosporidae,
Peronosporales, Peronosporaceae Peronosporales, Peronosporaceae Peronosporales, Peronosporaceae
7243. Phytophthora betacei M.F. Mideros, 7244. Phytophthora cactorum (Lebert & 7245. Phytophthora cinnamomi Rands
L.E. Lagos & S. Restrepo IF No: 815748 Cohn) J. Schröt. IF No: 199322 IF No: 260884
Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis,
Peronosporea, Peronosporidae, Peronosporea, Peronosporidae, Peronosporea, Peronosporidae,
Peronosporales, Peronosporaceae Peronosporales, Peronosporaceae Peronosporales, Peronosporaceae
7246. Phytophthora citricola Sawada 7247. Phytophthora drechsleri Tucker 7248. Phytophthora infestans (Mont.) de
IF No: 260970 IF No: 251892 Bary IF No: 232148
Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis,
Peronosporea, Peronosporidae, Peronosporea, Peronosporidae, Peronosporea, Peronosporidae,
Peronosporales, Peronosporaceae Peronosporales, Peronosporaceae Peronosporales, Peronosporaceae
7249. Phytophthora nicotianae Breda de 7250. Phytophthora palmivora (E.J. Butler) 7251. Phytophthora sojae Kaufm. & Gerd.
Haan IF No: 194443 E.J. Butler IF No: 194605 IF No: 303624
Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis,
Peronosporea, Peronosporidae, Peronosporea, Peronosporidae, Peronosporea, Peronosporidae,
Peronosporales, Peronosporaceae Peronosporales, Peronosporaceae Peronosporales, Pythiaceae
7252. Plasmopara borreriae (Lagerh.) 7253. Plasmopara viticola (Berk. & M.A. 7254. Globisporangium splendens (Hans
Constant. IF No: 354627 Curtis) Berl. & De Toni IF No: 208592 Braun) Uzuhashi, Tojo & Kakish.
IF No: 517635
Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis,
Peronosporea, Peronosporidae, Peronosporea, Peronosporidae, Peronosporea, Peronosporidae,
Peronosporales, Pythiaceae Peronosporales, Pythiaceae Peronosporales, Pythiaceae
7255. Globisporangium sylvaticum (W.A. 7256. Lagenidium giganteum Couch ex 7257. Pythium arrhenomanes Drechsler IF
Campb. & F.F. Hendrix) Uzuhashi, Tojo & Redhead No: 162965 Trophic mode/Guild:
Kakish. IF No: 517636 IF No: 551798 pathotroph/plant pathogen
Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis, Chromista, Oomycota, Incertae sedis,
Peronosporea, Peronosporidae, Peronosporea, Peronosporidae, Peronosporea, Peronosporidae,
Peronosporales, Pythiaceae Peronosporales, Pythiaceae Peronosporales, Pythiaceae
7258. Pythium graminicola Subraman. IF 7259. Pythium insidiosum De Cock, L. 7260. Pythium myriotylum Drechsler IF No:
No: 119045 Trophic mode/Guild: Mend., A.A. Padhye, Ajello & Kaufman IF 271965 Trophic mode/Guild: pathotroph/
pathotroph/plant pathogen No: 130421 Trophic mode/Guild: plant pathogen
pathotroph/plant pathogen
Chromista, Oomycota, Incertae sedis,
Peronosporea, Peronosporidae,
Rhipidiales, Salispinaceae
7261. Salispina spinosa (Fell & Master)
Marano, A.L. Jesus & Pires–Zottar.
IF No: 551604
Trametes polyzona
CATALOGUE OF FUNGI OF COLOMBIA [Ana Cristina Bolaños] 425
Cortinarius violaceus
[Natalia Vargas-Estupiñan]
ASCOMYCOTA
Laminae
Selected species of Colombian fungi
Agonimia foliacea Robert Lücking Akanthomyces tuberculatus Tatiana Sanjuan Allographa angustata Robert Lücking
Allophoron farinosum Robert Lücking Alternaria alternata Álvaro Rúa Annulohypoxylon thouarsianum Nataly Gómez-
Montoya
Ascopolyporus polychrous Aida Marcela Vasco-Palacios Aspergillus section Fumigati Álvaro Rúa Aspergillus section Nigri Álvaro Rúa
Aspergillus section Nigri Álvaro Rúa Aspergillus section Terrei Álvaro Rúa Aspergillus section Terrei Álvaro Rúa
Auriculora byssomorpha Robert Lücking Baeomyces rufus Robert Lücking Bathelium madreporiforme Robert Lücking
Beauveria locustiphila Tatiana Sanjuan Blackwellomyces sp. Tatiana Sanjuan Brigantiaea leucoxantha Robert Lücking
Bunodophoron melanocarpum Robert Lücking Byssoloma subdiscordans Robert Lücking Caloplaca brebissonii Robert Lücking
Chlorociboria aeruginosa Robert Lücking Cladia aggregata Robert Lücking Cladonia confusa Robert Lücking
Cladonia grayi Robert Lücking Cladonia secundana Robert Lücking Coccocarpia pellita Robert Lücking
Crocodia aurata Robert Lücking Cookeina tricholoma Viviana Motato-Vásquez Cresponea melanocheiloides Robert Lücking
Cryptothecia striata Robert Lücking Curvularia sp. Álvaro Rúa Dibaeis columbiana Robert Lücking
Diorygma poitaei Robert Lücking Dyplolabia afzelii Robert Lücking Enchylium conglomeratum Robert Lücking
Entonaema cinnabarinum Nataly Gómez-Montoya Erioderma divisum Robert Lücking Exophiala jeanselmei Álvaro Rúa
Fusarium oxysporum Álvaro Rúa Fusarium oxysporum Álvaro Rúa Glyphis scyphulifera Robert Lücking
Graphis dolichographa Robert Lücking Gyalidea hyalinescens Robert Lücking Helminthocarpon leprevostii Robert Lücking
Herpothallon rubrocinctum Robert Lücking Heterodermia comosa Robert Lücking Hypoxylon rubiginosum Nataly Gómez-Montoya
Hortaea werneckii Álvaro Rúa Icmadophila aversa Robert Lücking Leotia lubrica Aida Marcela Vasco-Palacios
Leptogidium dendriscum Robert Lücking Lobariella sipmanii Robert Lücking Lopezaria versicolor Robert Lücking
Morchella sp. Tatiana Sanjuan Neoscytalidium dimidiatum Álvaro Rúa Neoscytalidium dimidiatum Álvaro Rúa
Ophiocordyceps australis Tatiana Sanjuan Ophiocordyceps fulgoromorphila Kent Brothers Oropogon loxensis Robert Lücking
Paecilomyces variotti Álvaro Rúa Pannaria rubiginosa Robert Lücking Parainoa subconcolor Robert Lücking
Peltigera vainioi Robert Lücking Penicillium sp. Álvaro Rúa Penicillium sp. Álvaro Rúa
Phyllopsora breviuscula Robert Lücking Phillipsia domingensis Ana Esperanza Franco-Molano Phyllopsora pyxinoides Robert Lücking
Placopsis rhodocarpa Álvaro Rúa Placopsis roivainenii Álvaro Rúa Polycauliona candelaria Robert Lücking
Porpidia macrocarpa Álvaro Rúa Pseudochapsa subdactylifera Álvaro Rúa Pseudocyphellaria sandwicensis Robert Lücking
Psoroma hypnorum Robert Lücking Punctelia stictica Robert Lücking Pyrenula astroidea Robert Lücking
Pyrenula ochraceoflava Robert Lücking Pyxine coccifera Robert Lücking Ramalina celastri Robert Lücking
Ramboldia russula Robert Lücking Rhabdodiscus anamorphoides Robert Lücking Rhizina undulata Nataly Gómez-Montoya
Rhizocarpon geographicum Robert Lücking Rusavskia elegans Robert Lücking Sarcographa cinchonarum Robert Lücking
Stereocaulon novogranatense Robert Lücking Stereocaulon ramulosum Robert Lücking Sticta dilatata Robert Lücking
Stigmatochroma metaleptoides Robert Lücking Strigula smaragdula Robert Lücking Tephromela atra Robert Lücking
Usnea baileyi Robert Lücking Xanthoparmelia taractica Robert Lücking Xanthoparmelia taractica Nataly Gómez-Montoya
Xylaria curta Nataly Gómez-Montoya Xylaria multiplex Robert Lücking Xylaria polymorpha Nataly Gómez-Montoya
Amanita citrina Natalia Vargas Estupiñan Amanita colombiana Natalia Vargas Estupiñan
Amanita flavoconia Nataly Gómez-Montoya Amanita flavoconia Natalia Vargas Estupiñan Amanita rubescens Aida Marcela Vasco-Palacios
Amanita virosa Natalia Vargas Estupiñan Amanita xerocybe Aida Marcela Vasco-Palacios
Aseroe rubra Nataly Gómez-Montoya Auricularia delicata Robert Lücking Auricularia fuscosuccinea Ana Esperanza Franco-Molano
Auricularia mesenterica Ana Cristina Bolaños Auricularia mesenterica Aida Marcela Vasco-Palacios Auricularia polytricha Ana Cristina Bolaños
Austropuccinia psidii Mauricio Salazar Blumenavia angolensis Nataly Gómez-Montoya Calvatia cyathiformis Nataly Gómez-Montoya
Calostoma cinnabarinum Natalia Vargas Estupiñan Calostoma cinnabarinum Bibiana Moncada Cantharellus coccolobae Aida Marcela Vasco-Palacios
Campanella caesia Robert Lücking Cerioporus flavus Nataly Gómez-Montoya Chlorophyllum molybdites Ana Esperanza Franco-Molano
Clathrus archeri Ana Esperanza Franco-Molano Clavariachaete rubiginosa Nataly Gómez-Montoya Clavulina kunmudlutsa Aida Marcela Vasco-Palacios
Collybia plectophylla Ana Cristina Bolaños Collybia plectophylla Aida Marcela Vasco-Palacios
Clavulinopsis fusiformis Nataly Gómez-Montoya Coltricia perennis Gineth Adriana Calderón Coprinellus disseminatus Robert Lücking
Cortinarius violaceus Natalia Vargas Estupiñan Cortinarius violaceus Aida Marcela Vasco-Palacios Cortinarius iodes Aida Marcela Vasco-Palacios
Cotylidia aurantica Aida Marcela Vasco-Palacios Craterellus boyacensis Tatiana Sanjuan Craterellus fallax Natalia Vargas Estupiñan
Crepidotus appalachianensis Nataly Gómez-Montoya Crossopsora byrsonimae Meike Piepenbring Craterellus fallax Aida Marcela Vasco-Palacios
Cyanoboletus pulverulentus Nataly Gómez-Montoya Cyathus striatus Aida Marcela Vasco-Palacios Cymatoderma caperatum Nataly Gómez-Montoya
Cyptotrama asprata Nataly Gómez-Montoya Cystidiodontia laminifera Viviana Motato-Vásquez Dacrymyces lacrymalis Nataly Gómez-Montoya
Dacryopinax spathularia Kent Brothers Dictyonema sericeum Robert Lücking Earliella scabrosa Nataly Gómez-Montoya
Entoloma incanum Nataly Gómez-Montoya Favolus brasiliensis Aida Marcela Vasco-Palacios Filobletus gracilis Nataly Gómez-Montoya
Flavodon flavus Aida Marcela Vasco-Palacios Flavodon flavus Aida Marcela Vasco-Palacios Fulvifomes inermis Nataly Gómez-Montoya
Fuscocerrena portoricensis Viviana Motato-Vásquez Fuscoporia senex Nataly Gómez-Montoya Fuscoporia wahlbergii Nataly Gómez-Montoya
Ganoderma australe Ana Cristina Bolaños Ganoderma australe Robert Lücking Fuscoporia wahlbergii Nataly Gómez-Montoya
Ganoderma multiplicatum Ana Cristina Bolaños Geastrum fimbriatum Nataly Gómez-Montoya Geastrum pectinatum Aida Marcela Vasco-Palacios
Geastrum saccatum Nataly Gómez-Montoya Geastrum triplex Nataly Gómez-Montoya Gloeocantharellus uitotanus Aida Marcela Vasco-Palacios
Gloiocephala quercetorum Natalia Vargas Estupiñan Hericium erinaceus Ana Esperanza Franco-Molano Hemileia vastatrix Mauricio Salazar
Hexagonia tenuis Ana Cristina Bolaños Hohenbuehelia nigra Nataly Gómez-Montoya Hydnodon thelephorus Aida Marcela Vasco-Palacios
Hydnum repandum Aida Marcela Vasco-Palacios Hydnum repandum Nataly Gómez-Montoya Laccaria gomezii Nataly Gómez-Montoya
Laccaria laccata Nataly Gómez-Montoya Lactarius gerardii Robert Lücking Lactarius gerardii Aida Marcela Vasco-Palacios
Lactarius chrysorrheus Natalia Vargas Estupiñan Lactarius indigo Aida Marcela Vasco-Palacios Lactarius indigo Aida Marcela Vasco-Palacios
Lactifluus hallingii Robert Lücking Lactifluus hallingii Natalia Vargas Estupiñan Laetiporus sulphureus Ana Esperanza Franco-Molano
Leccinum andinum Aida Marcela Vasco-Palacios Lentinus tricholoma Nataly Gómez-Montoya Lentinus crinitus Nataly Gómez-Montoya
Lepidostroma calocerum Robert Lücking Lepista nuda Aida Marcela Vasco-Palacios Leucoagaricus rubrotinctus Natalia Vargas Estupiñan
Leucocoprinus fragilissimus Natalia Vargas Estupiñan Leucopaxillus gracillimus Aida Marcela Vasco-Palacios
Lycoperdon sp. Aida Marcela Vasco-Palacios Lycoperdon sp. Natalia Vargas Estupiñan
Macrolepiota colombiana Ana Esperanza Franco-Molano Maramius cladophyllus Nataly Gómez-Montoya Maramius rhabarbinus Nataly Gómez-Montoya
Mutinus caninus Ana Cristina Bolaños Mycena alcalina Natalia Vargas Estupiñan Mycena holoporphyra Ana Esperanza Franco-Molano
Mycena pura Natalia Vargas Estupiñan Mycena pura Natalia Vargas Estupiñan Oudemansiella Ana Cristina Bolaños Panaeolus Ana Esperanza
canarii semiovatus Franco-Molano
Panaeolus Ana Esperanza Phaeoclavulina Aida Marcela Phaeomarasmius Natalia Vargas Phallus indusiatus Aida Marcela
sphinctrinus Franco-Molano zippelii Vasco-Palacios sp. Estupiñan Vasco-Palacios
Phanerochaete australis Viviana Motato-Vásquez Phellinus setulosus Nataly Gómez-Montoya Phlebia chrysocreas Viviana Motato-Vásquez
Pisolithus arhizus Nataly Gómez-Montoya Phlebiopsis crassa Aida Marcela Vasco-Palacios Phylloporus fibulatus Nataly Gómez-Montoya
Pleurotus djamor Ana Cristina Bolaños Pleurotus djamor Nataly Gómez-Montoya Pleurotus ostreatoroseus Ana Cristina Bolaños
Pluteus nanus Nataly Gómez-Montoya Podaxis pistilaris Aida Marcela Vasco-Palacios Polyporus leprieurii Nataly Gómez-Montoya
Polyporus udus Kent Brothers Pseudohydnum gelatinosum Aida Marcela Vasco-Palacios Psilocybe cubensis Natalia Vargas Estupiñan
Psilocybe cubensis Natalia Vargas Estupiñan Pycnoporus sanguineus Natalia Vargas Estupiñan Ramaria gracilis Nataly Gómez-Montoya
Rhizochaete radicata Viviana Motato-Vásquez Rhodophana nitellina Nataly Gómez-Montoya Ramaria stricta Bibiana Moncada
Rigidoporus microporus Viviana Motato-Vásquez Russula emetica Nataly Gómez-Montoya Russula emetica Nataly Gómez-Montoya
Scleroderma areolatum Nataly Gómez-Montoya Schizophyllum commune Natalia Vargas Estupiñan Schizophyllum commune Robert Lücking
Scleroderma flavidum Robert Lücking Stalgilomyces sp. Tatiana Sanjuan Stereopsis radicans Nataly Gómez-Montoya
Stereum complicatum Viviana Motato-Vásquez Stereum ostrea Robert Lücking Strobilomyces confusus Natalia Vargas Estupiñan
Thelephora cf. palmata Aida Marcela Vasco-Palacios Trametes elegans Natalia Vargas Estupiñan
Tetrapyrgos alba Nataly Gómez-Montoya Trametes pavonia Aida Marcela Vasco-Palacios Trametes polyzona Ana Cristina Bolaños
Trametes versicolor Robert Lücking Trametes villosa Natalia Vargas Estupiñan Trametes villosa Nataly Gómez-Montoya
Tremella fuciformis Ana Cristina Bolaños Tremella mesenterica Aida Marcela Vasco-Palacios Tricholoma sp. Natalia Vargas Estupiñan
Trogia papyracea Aida Marcela Vasco-Palacios Trogia papyracea Natalia Vargas Estupiñan Truncospora ochroleuca Nataly Gómez-Montoya
Xerocomellus chrysenteron Natalia Vargas Estupiñan Ustilago maydis Meike Piepenbring Xylodon radula Viviana Motato-Vásquez
Aseroe rubr
[Nataly Gómez-Montoya]
458 CATALOGUE OF FUNGI OF COLOMBIA
Index of Families
Bellicidia 281 Bunodophoron 121, 130, 131, Catenulopsora 353 Cionothrix 364
Belonium 322 284, 428 Catillaria 263 Circinella 108, 218
Belonopsis 321 Byssocaulon 228 Catillochroma 282 Cladia 263, 429
Beltrania 343 Byssolecania 260 Celothelium 248 Cladonia 36, 82, 263–265,
Beltraniella 344 Byssoloma 261, 428 Cephalotrichiella 340 429
Berkeleyomyces 339 Byssomerulius 408 Ceraceomyces 397 Cladophialophora 247
Biatora 281 Byssonectria 325 Ceratiomyxa 423 Cladosporium 143, 157, 170,
Biatoropsis 372 Byssosphaeria 238 Ceratobasidium 143, 156 232
Bibbya 281 Ceratocarpia 246 Clandestinotrema 305
Binderoboletus 61 C Ceratocystis 339, 340 Clarohilum 233
Bionectria 73, 328 Cabalodontia 415 Cercidospora 246 Claroideoglomus 220
Bioscypha 323 Cacumisporium 349 Cercospora 37, 170, 232, 233 Clastoderma 421
Biostictis 315 Cadophora 322 Cercosporella 233 Clathrus 402, 443
Bipolaris 239 Caeoma 366 Cericium 376 Clavaria 68, 375
Biscogniauxia 345 Calathella 383 Cerion 324 Clavariachaete 443
Bisporella 69, 71, 319 Calcarisporium 329 Cerioporus 48, 411, 443 Clavariopsis 240
Bjerkandera 410 Calenia 296, 297 Ceriporia 408, 409 Clavatospora 340
Blackwellomyces 330, 428 Caleniopsis 297 Ceriporiopsis 409 Claviceps 166, 329
Blastenia 294 Calicium 256 Cerotelium 353 Clavispora 222
Blastobotrys 224 Calidion 365 Cerrena 407 Clavochytridium 216
Blastocladiella 216 Callistosporium 375 Cetraria 268, 269 Clavulina 61, 129, 180, 403,
Blastospora 352 Calocera 370 Cetraspora 219, 220 443
Blastosporella 383 Calocybe 383 Chaconia 352 Clavulinopsis 375, 443
Blumenavia 402, 442 Calogaya 294 Chadefaudia 340 Clithris 318
Blumeria 319 Calonectria 333 Chaenotheca 178, 231 Clitocybe 395
Boeremia 237 Calopadia 261 Chaenothecopsis 255 Clitocybula 395
Bogbodia 394 Caloplaca 294, 295, 428 Chaetocalathus 383 Clitopilus 376
Bogoriella 245 Calostilbe 333 Chaetomidium 343 Clonostachys 170, 328
Bolbitius 375 Calostoma 48, 399, 442 Chaetopsina 333 Clypeolum 350
Boletellus 398 Calothyrium 242 Chaetoseptoria 349 Clypeopyrenis 248
Boletinellus 399, 442 Calvatia 383, 442 Chaetosphaeria 341 Coccidioides 255
Boletus 48, 127, 398 Calvitimela 285 Chaetothyrium 247 Coccocarpia 82, 286, 429
Boreostereum 404 Calycellina 322 Chaetotyphula 393 Coccodiella 341
Botryobasidium 402 Calycina 71, 322 Chalara 322 Coccomyces 324
Botryodiplodia 326 Calyptella 395 Chalciporus 398 Coccostroma 341
Botryodontia 404 Camarotella 341 Chapsa 304, 305 Cochlearomyces 323
Botryosphaeria 170, 241 Camillea 68, 345, 346 Chardonia 350 Cochliobolus 239
Botryostroma 241 Campanella 16, 48, 383, 443 Chardoniella 112, 114, 116, Codinaea 341
Botryotinia 322 Campanophyllum 376 364 Coelomomyces 108, 216
Botryotrichum 143, 343 Camposporium 349 Cheilymenia 325 Coenogonium 298–300
Botrytis 157, 159, 170, 322 Camptomeris 349 Chiodecton 229 Coleosporium 352
Bovista 382, 383 Campylocarpon 333 Chlorencoelia 318 Collaria 422
Brasilicia 260 Campylospora 349 Chlorociboria 48, 149, 318, Collariella 343
Bresadolia 121, 126, 411 Canasta 365 429 Collema 286
Brevicellicium 419 Candelabrochaete 415 Chlorophyllum 373, 443 Colletotrichum 143, 170, 327,
Brevicellopsis 420 Candelaria 231 Choanephora 217 328
Brigantiaea 294, 428 Candida 48, 68, 92, 96, 98, Chromocyphella 375 Collybia 48, 395, 443
Brownliella 294 99, 100, 107, 224, 225 Chroodiscus 305 Colpoma 324
Brunneochlamydosporium 328 Canoparmelia 268 Chrysocelis 116, 352 Coltricia 404, 443
Brunneocorticium 383 Cantharellus 61, 157, 402, Chrysocyclus 354 Coltriciella 129, 404
Bryochiton 246 403, 442 Chrysofolia 326 Comatricha 421
Bryorella 246 Capnodium 232 Chrysoporthe 170, 326 Conferticium 418
Bryoria 268 Capronia 247 Chrysopsora 354 Coniarthonia 227
Bryostigma 230 Carbacanthographis 304 Chrysosporium 255 Conidiobolus 108, 216
Buellia 47, 256 Caribaeomyces 242 Chrysothrix 228 Conidiosporomyces 368
Buelliella 246 Castanediella 345 Chytridium 216 Coniella 326
Bulbothrix 268 Catacauma 341 Cinnamomeonectria 333 Coniocarpon 227
Bullatina 296 Catenulocercospora 232 Cintractia 117, 368, 369 Coniochaeta 325, 326
Favolus 158, 168, 169, 411, Geastrum 48, 401, 447 H Hydnopolyporus 48, 124, 409,
446 Gelatinomyces 323 Haddowia 413 448
Fellhanera 262 Geminispora 342 Haematomma 265, 266 Hydnoporia 405
Fellomyces 371 Genea 325 Haiyanga 340 Hydnotrya 72
Fibrillithecis 306 Geoglossum 72, 255 Halophytophthora 423 Hydnum 403, 449
Fibrodontia 406 Geomyces 324 Hamatocanthoscypha 319 Hydropisphaera 329
Fibroporia 408 Gerronema 396 Hannaella 371 Hydropus 387
Filobasidium 370, 371 Gerwasia 354 Hanseniaspora 98, 223 Hygroaster 379
Filoboletus 387, 446 Gibbera 241 Hansfordia 346 Hygrocybe 379, 380, 449
Filosporella 323 Gibberella 170 Hansfordiella 242 Hygrophorus 380
Fimetariella 344 Gibellula 331 Harknessia 326 Hymenochaete 124, 405
Fissurina 306 Gigaspora 220 Hegermila 320 Hymenopsis 350
Fistulina 396 Gilmaniella 350 Heiomasia 310 Hymenoscyphus 319, 320
Fistulinella 398, 446 Glabrocyphella 396 Helicobasidium 352 Hymenotorrendiella 320
Flabellimycena 387 Glaucotrema 306 Helicogloea 351 Hyperdermium 331
Flabelloporina 300 Gliocephalotrichum 336 Helicogonium 323 Hyperphyscia 258, 259
Flabellospora 350 Gliocladiopsis 336 Helicomyces 240 Hyphoderma 408
Flagelloscypha 388 Gliomastix 339 Heliscella 350 Hyphodermella 410
Flagellospora 323 Gliophorus 379 Heliscus 336 Hyphodontia 406, 407
Flakea 250 Globisporangium 424 Helminthocarpon 230, 431 Hypholoma 394
Flammulina 157, 391 Globonectria 328 Helminthosporiella 238 Hyphopichia 225
Flaviporus 124, 415, 446 Gloeocantharellus 128, 401, Helminthosporium 238 Hyphopolynema 350
Flavobathelium 243 447 Helotium 319 Hypocenomyce 318
Flavodon 409, 446 Gloeodontia 418 Helvella 72, 325 Hypochnicium 415
Flavoparmelia 269 Gloeomucro 403 Hemileia 366, 448 Hypocrea 331
Flavopunctelia 269 Gloeophyllum 404 Hemimycena 387 Hypocrella 329
Fomes 411, 412 Gloeoporus 409 Hemitrichia 422 Hypocreopsis 331
Fomitella 412 Gloeotinia 319 Hennenia 116, 366 Hypogymnia 269
Fomitiporella 405 Gloiocephala 61, 180, 391, Hericium 48, 61, 157, 169, Hypomontagnella 346
Fomitiporia 61, 181, 405 448 181, 416, 448 Hypomyces 331
Fomitopsis 408 Glomus 39, 108, 141, 171, Herpomyces 255 Hypotrachyna 269–272
Fonsecaea 247 221 Herpothallon 227, 431 Hypotrachynicola 349
Foraminispora 412 Glyphis 306, 430 Herpotrichia 238 Hypoxylon 346, 347, 431
Fouragea 228 Golovinomyces 319 Heterocephalacria 371 Hypsizygus 383
Fuligo 421 Gomphillus 297 Heterochaete 400, 401
Fulvia 233 Gongronella 217 Heterodermia 258, 431 I
Fulvifomes 405, 446 Goplana 352 Heteroradulum 401 Icmadophila 180, 316, 431
Fulvophyton 230 Grammothele 413 Hevansia 331 Ijuhya 329
Funalia 412 Graphis 82, 307–309, 431 Hexagonia 413, 448 Illosporium 339
Funneliformis 220 Grifola 408 Hirsutella 337 Ilyonectria 339
Furcasterigmium 328 Grovesia 323 Histoplasma 254 Imshaugia 272
Furtadoa 412 Guepinia 401 Hjortstamia 410 Inapertisporites 420
Fusarium 39, 68, 143, 169, Guepiniopsis 370 Hobsonia 351 Incrupila 320
170, 171, 334–336, 430 Gyalectidium 297 Hohenbuehelia 391, 392, 448 Incrustocalyptella 376
Fuscidea 318 Gyalidea 297, 431 Holocotylon 373 Induratia 347
Fuscocerrena 412, 447 Gyalideopsis 297, 298 Homophron 393 Inflatostereum 415
Fuscoporia 405, 447 Gyalolechia 295 Homostegia 247 Inocephalus 377
Fuscutata heterogama 220 Gymnographopsis 310 Hortaea 431 Inocybe 382
Fusicolla 336 Gymnopilus 381 Humicola 344 Inonotus 405
Gymnopus 389, 448 Humphreya 413 Inosperma 382
G Gymnosporangium 352 Huneckia 295 Intralichen 350
Gaeumannomyces 341 Gymnostellatospora 324 Hyalocamposporium350 Iodophanus 325
Galerina 380 Gyoerffyella 319 Hyalopsora 363 Ionomidotis 318
Gallowayella 295 Gyrodontium 399 Hyaloria 371 Irenina 341
Galzinia 404 Gyronactis 229 Hyaloscypha 320 Irenopsis 341
Ganoderma 124, 157, 159, Gyroporus 399 Hydnangium 377 Irpex 409
169, 412, 447 Gyrothrix 350 Hydnellum 419 Isaria 331
Gassicurtia 257 Gyrotrema 310 Hydnodon 448 Issatchenkia 222
Morchella 72, 168, 169, 325, Neofusicoccum 241 Oudemansiella 157, 158, 391, Penicilliopsis 68, 251
433 Neometulocladosporiella 322 452 Penicillium 39, 107, 143, 166,
Morganella 383 Neonectria 73, 337 Ovatospora 344 170, 171, 251–253, 434
Mortierella 69, 108, 216, 217 Neonothopanus 390 Ovicuculispora 329 Penidiella 236
Mucidula 391 Neopestalotiopsis 345 Oxyporus 406 Peniophora 416
Mucor 108, 157, 217, 218 Neophysopella 366 Peniophorella 406
Multicellaesporites 420 Neoprotoparmelia 121, 130, P Penzigomyces 350
Multiclavula 403 131, 273 Pachykytospora 413 Perenniporia 413
Munkiodothis 342 Neoscytalidium 110, 241, 433 Paecilomyces 143, 251 Perichaena 422
Musicillium 328 Neottiosporina 350 Paecilomyces 434 Periconia 240
Musidium 328 Nephridiophaga 216 Palifer 407 Peristomialis 329
Mutatoderma 408 Nephroma 290 Pallidocercospora 234 Peroneutypa 345
Mutinus 402, 451 Nephromopsis 273 Pallidogramme 312 Peronosclerospora 423
Mycena 60, 387, 388, 451, Neurospora 344 Panaeolina 397 Peronospora 423
452 Niesslia 337 Panaeolus 48, 58, 397, 452 Perrotia 321
Mycetinis 390 Nigelia 330 Pandora 216 Pertusaria 317
Mycoacia 409 Nigrohydnum 416 Panellus 388 Pestalotia 345
Mycocalicium 255 Nigromacula 350 Pannaria 14, 288, 434 Pestalotiopsis 345
Mycochlamys 350 Nigroporus 415 Pantospora 234 Petriella 340
Mycofalcella 323 Nigrospora 143, 349 Panus 164, 168, 169, 409, Peziza 68, 325
Mycoporum 238 Nigrovothelium 245, 433 410 Pezizella 322
Mycosphaerella 233 Niorma 295 Papiliotrema 371 Phacidium 324
Mycosyrinx 118, 370 Niptera 321 Paracercosporidium 96, 234 Phacopsis 276
Mycovellosiella 234 Nitschkia 327 Paracoccidioides 254 Phaeoacremonium 327
Myelochroa 273 Nochascypha 396 Paraconiothyrium 237, 238 Phaeoclavulina 401, 452
Myeloconis 300 Normandina 250, 433 Paracremonium 337 Phaeocollybia 381
Myriochapsa 310 Norrlinia 247 Paragalactinia 325 Phaeocytostroma 350
Myriogenospora 330 Nothopanus 392 Paraglomus 222 Phaeodepas 397
Myriolecis 267 Novomicrothelia 245 Paragyalideopsis 298 Phaeodimeriella 246
Myriophacidium 324 Nummularia 346 Parainoa 295, 434 Phaeographis 76, 312, 313,
Myriospora 256 Paraisaria 338 435
Myriostigma 227 O Parallopsora 282 Phaeoisariopsis 234
Myriotrema 310 Oberwinkleria 368 Paramyrothecium 339 Phaeolepiota 397
Myrmaecium 246 Ocellularia 311 Parapallidocercospora 234 Phaeolus 409
Myxarium 401 Ochroconis 350 Paraphaeosphaeria 238 Phaeomarasmius 395, 452
Ochrolechia 316 Parapithomyces 350 Phaeophlebiopsis 410
N Odonticium 420 Parasiphula 316 Phaeophleospora 234
Nadsonia 225 Oehlia 221 Parasola 393 Phaeophyscia 259
Nadvornikia 310 Oidiodendron 322 Parastagonospora 238 Phaeosaccardinula 247
Naganishia 371 Oidium 319 Parasterina 232 Phaeoscypha 322
Nakazawaea 225 Olivea 352 Parasympodiella 349 Phaeoseptoria 238
Nanostictis 315 Oliveonia 401 Parmelia 273 Phaeosolenia 375
Napicladium 241 Olpidiaster 216 Parmeliella 289 Phaeosphaeria 238
Nawawia 350 Ombrophila 319 Parmelinella 273 Phaeosporis 344
Nebularia 288 Omphalina 397 Parmelinopsis 273 Phaeosporobolus 231
Nectria 73, 336 Oomyces 241 Parmotrema 273–276, 434 Phaeostigme 246
Nectriopsis 329 Opegrapha 228 Parmotremopsis 276 Phaeotrabutia 342
Nemania 347 Opegrapha 312 Parodiella 246 Phaeotremella 371
Neobarya 330 Ophiocordyceps 68, 71, 73, 74, Parodiopsis 246 Phakopsora 116, 117, 353,
Neobulgaria 319 337, 338, 433 Parvothecium 339 354
Neocallimastix 157 Ophiodothella 342 Passalora 234 Phalangispora 350
Neocercosporidium 234 Ophionectria 337 Patouillardina 401 Phallus 48, 402, 452, 511
Neoconidiobolus 216 Ophioparma 318 Paxillus 400 Phanerochaete 410, 452
Neocosmospora 170, 336, Orbilia 324 Peloronectria 339 Phanerodontia 416
337 Orcularia 257 Peltella 236 Phaneroites 416
Neodasyscypha 321 Oropogon 273, 433 Peltigera 290, 434 Phellinidium 405
Neodictyopus 121, 126 Orphanocoela 350 Peltula 324 Phellinopsis 406
Neoeriomycopsis 349 Otospora 219 Penicillifer 337 Phellinotus 125
Phellinus 406, 452 Placopsis 79, 295, 296, 435 Propolis 318 Puccinia 35, 37, 62, 114, 115,
Phellodon 419 Placynthiella 296 Prospodium 117, 365 116, 117, 355–362, 454
Phellorinia 374 Plasmodiophora 423 Protocrea 331 Pucciniosira 364
Phialemoniopsis 349 Plasmopara 424 Protocreopsis 329 Puiggariella 243
Phialocephala 321 Platygramme 313 Protomerulius 401 Pulveroboletus 399
Phialophora 247 Platygraphopsis 302 Protomicarea 280 Punctelia 276, 436
Phillipsia 48, 68, 71, 325, Platypeltella 242 Protoparmelia 276 Purpureocillium 171, 339
435 Platythecium 313, 314 Protoscypha 246 Pustula 423
Phlebia 409, 452, 453 Plectania 325 Protostropharia 394 Pycnoporus 157, 159, 414, 454
Phlebiella 416 Plectocarpon 228 Prototrichia 422 Pycnotrema 314
Phlebiopsis 410, 453 Plectosphaerella 328 Psathyrella 393 Pyramidospora 350
Phlebopus 399 Plenodomus 240 Pseudallescheria 156, 340 Pyrenochaetopsis 237
Phloeomana 393 Pleochaeta 319 Pseudeurotium 324 Pyrenophora 240
Phlyctibasidium 400 Pleotrichocladium 238 Pseudevernia 276 Pyrenula 82, 248–250, 436
Phlyctis 300 Pleurocollybia 375 Pseudobogoriella 246 Pyrgidium 255
Pholiota 394 Pleurocybella 397 Pseudocercosporella 235 Pyricularia 341
Pholiotina 375 Pleuroflammula 376 Pseudochaetosphaeronema Pyrrhoderma 406
Phoma 237 Pleuropedium 350 240 Pyrrhoglossum 376
Phomatospora 349 Pleurophragmium 350 Pseudochapsa 314, 435 Pythium 424
Phomopsis 170, 326 Pleurotus 150, 157, 168, 169, Pseudocosmospora 337 Pyxine 257, 258, 436
Phragmidiella 354 171, 392, 453 Pseudocraterellus 403
Phragmidium 354 Pliariona 314 Pseudocyphellaria 121, 131, Q
Phragmocapnias 248 Pluricellaesporites 420 132, 290, 291, 435 Questieria 246
Phylacia 347 Pluteus 392, 393, 453 Pseudofavolus 414
Phyllachora 342, 343 Pneumocystis 225 Pseudofistulina 397 R
Phyllobaeis 82, 295, 435 Podaxis 374, 453 Pseudogibellula 331 Raciborskiella 242
Phylloblastia 250 Podila 108 Pseudographis 349 Racoplaca 243
Phyllocharis 243 Podoscypha 410 Pseudogymnoascus 324 Radulodontia 420
Phyllocrea 343 Podosordaria 347 Pseudohydnum 401, 454 Radulomyces 393, 394
Phyllogloea 371 Podosphaera 319 Pseudoidium 319 Ramalina 283, 436
Phyllogyalidea 298 Podospora 344 Pseudolagarobasidium 408 Ramaria 128, 157, 401, 454
Phylloporia 406 Podosporium 350 Pseudolostoma 180 Ramaricium 402
Phylloporis 243 Podostictina 121, 131, 132, Pseudoomphalina 397 Ramariopsis 375
Phylloporus 61, 128, 398, 399, 290 Pseudoparmelia 276 Ramboldia 284, 436
453 Podoxyphium 248 Pseudoparodiella 241 Ramularia 235
Phyllopsora 435 Polioma 354, 355 Pseudopestalotiopsis 345 Ramulispora 235
Phyllosticta 241 Polyblastidium 260 Pseudopeziza 319 Ranadivia 408
Phyllotopsis 390, 391 Polycauliona 295, 435 Pseudopithomyces 238 Rasamsonia 253
Physalacria 391 Polycephalomyces 338 Pseudoplectania 325 Ravenelia 117, 364
Physarella 421 Polycoccum 243 Pseudopyrenochaeta 240 Redeckera 219
Physarum 421 Polydesmia 323 Pseudopyrenula 246 Redingeria 314
Physcia 259, 260 Polymeridium 245 Pseudorhytisma 324 Reimnitzia 314
Physcidia 282 Polymyxa 423 Pseudosperma 382 Relicina 276, 277
Physocystidium 397 Polyporoletus 416 Pseudosydowia 236 Remototrachyna 277
Physopella 354 Polyporus 121, 124, 126, 414, Pseudoteratosphaeria 236 Repetobasidium 403
Phytophthora 157, 423, 424 453, 454 Pseudothis 327 Resinicium 406
Piccolia 318 Porina 300–302, 317 Pseudotulostoma 129, 254 Resinomycena 388
Pichia 96, 98, 99, 100, 223 Porodaedalea 406 Pseudozasmidium 235 Resupinatus 392
Picipes 414 Porodisculus 394 Pseudozyma 369 Reticularia 423
Pilatoporus 408 Porogramme 414 Psilachnum 322 Rexia 265
Pilobolus 218 Poronia 68, 347 Psilocybe 153, 159, 381, 382, Rhabdodiscus 314, 436
Pilocintractia 369 Porostereum 410 454 Rhabdospora 235
Piloderma 398 Porphyrellus 157, 399 Psilolechia 280 Rhachomyces 255
Pilophorus 265 Porpidia 285, 435 Psiloparmelia 276 Rhagadostomella 327
Piskurozyma 371 Preussia 240 Psoroglaena 250 Rhinocladiella 248
Pisolithus 400, 453 Prillieuxina 232 Psoroma 289, 436 Rhinotrichella 350
Pithyella 320 Proliferodiscus 321 Pterula 393 Rhizina 436
Placoasterina 232 Pronectria 329 Pterulicium 393 Rhizocarpon 293, 294, 436
Calenia conspersa (Stirt.) R. Sant. Rolueckia conspersa (Stirt.) Candida parazyma Lachance Wickerhamiella parazyma
Papong, Thammath. & Boonpr. (Lachance) C. Vega & Lachance
Calenia laevigata Müll. Arg. Caleniopsis laevigata (Müll. Arg.) Candida rugosa (H.W. Anderson) Diutina rugosa (H.W. Anderson)
Vězda & Poelt Diddens & Lodder Khunnamw., Jindam., Limtong &
Lachance
Calenia submaculans R. Sant. Calenia triseptata Zahlbr.
Candida sergipensis R.C. Trindade, Wickerhamiella sergipiensis (R.C.
Caleniopsis conspersa (Stirt.) Rolueckia conspersa (Stirt.) M.A. Resende, Lachance & C.A. Trindade, M.A. Resende, Lachance
Lücking et al. Papong, Thammath. & Boonpr. Rosa & C.A. Rosa) C. Vega & Lachance
Calicium curtum Turn. & Borr. Calicium abietinum Pers. Canomaculina muelleri (Vain.) Elix Parmotrema muelleri (Vain.) O.
Calicium subquercinum auct. non Calicium lenticulare Ach. & Hale Blanco, A. Crespo, Divakar, Elix &
Asahina Lumbsch
Callopisma conjugens (Nyl.) Müll. Caloplaca conjungens (Nyl.) Zahlbr. Canomaculina subtinctoria (Zahlbr.) Parmotrema subtinctorium (Zahlbr.)
Arg. Elix Hale
Calopeltis tetraspora Toro Caribaeomyces tetrasporus (Toro) Canoparmelia carneopruinata Crespoa carneopruinata (Zahlbr.)
Cif. (Zahlbr.) Elix & Hale Lendemer & B.P. Hodk.
Caloplaca cinnabarina (Ach.) Zahlbr. Brownliella cinnabarina (Ach.) S.Y. Canoparmelia salacinifera (Hale) Parmelinella salacinifera (Hale)
Kondr., Kärnefelt, A. Thell, Elix, Elix & Hale Marcelli & Benatti
Jung Kim, A.S. Kondr. & Hur Cantharellus lateritius var. Cantharellus lateritius (Berk.) Singer
Caloplaca citrina (Hoffm.) Th. Fr. Flavoplaca citrina (Hoffm.) Arup, colombianus R.H. Petersen
Frödén & Søchting Catacauma macroloculatum Phyllachora macroloculata Chardón
Caloplaca conjugens (Nyl.) Zahlbr. Caloplaca erythrantha (Tuck.) Chardón
Zahlbr. Catillaria endochroma (Fée) Zahlbr. Catillochroma endochromum (Fée)
Caloplaca conjungens (Nyl.) Zahlbr. Caloplaca erythrantha (Tuck.) Kalb
Zahlbr. Catinaria versicolor (Fée) Sipman Lopezaria versicolor (Flot.) Kalb &
Caloplaca diducta f. albicans (Nyl.) Caloplaca diducta (Nyl.) Zahlbr. Hafellner
Zahlbr. Cercospora aciotidis Chupp Pseudocercospora aciotidis (Chupp)
U. Braun & Crous
Caloplaca flavovirescens (Wulf.) Gyalolechia flavovirescens (Wulfen)
D.T.& Sarnth. Søchting, Frödén & Arup Cercospora ambrosiae Chupp Passalora ambrosiae (Chupp) Crous
& U. Braun
Caloplaca pollinii (A. Massal.) Jatta Huneckia pollinii (A. Massal.) S.Y.
Kondr., Kärnefelt, Elix, A. Thell, Cercospora coriariae Chupp Pseudocercospora coriariae (Chupp)
Jung Kim, A.S. Kondr. & Hur X.J. Liu & Y.L. Guo
Cercospora lantanicola Chupp Pseudocercosporella lantanicola Chapsa pseudoschizostoma (Hale) Pseudochapsa pseudoschizostoma
(Chupp) U. Braun & Crous Sipman (Hale) Parnmen, Lücking &
Lumbsch
Cercospora liabi Syd. & P. Syd. Passalora liabi (Syd. & P. Syd.) U.
Braun & Crous Chapsa referta (Hale) Lücking Ocellularia referta Hale
Cercospora miconiicola Chupp Pseudocercospora miconiicola Chiodecton decussans Nyl. Syncesia decussans (Nyl.) Tehler
(Chupp) U. Braun & Crous Chiodecton depressum Fée Syncesia depressa (Fée) Tehler
Cercospora polymniae Chupp Pseudocercospora polymniae Chiodecton dilatatum (Nyl.) Vain. Sagenidiopsis undulata (Fée) Egea,
(Chupp) U. Braun & Crous Tehler, Torrente & Sipman
Cercospora ulmifoliae Obreg.-Bot. Pseudocercospora ulmifoliae Chiodecton effusum Fée Syncesia effusa (Fée) Tehler
(Obreg.-Bot.) U. Braun & Crous Chiodecton extenuatum (Nyl.) Sclerophyton extenuatum (Nyl.)
Cerrena caperata (Berk.) Zmitr. Funalia caperata (Berk.) Zmitr. & Zahlbr. Sparrius
Malysheva Chiodecton farinaceum var. Syncesia effusa (Fée) Tehler
Cetraria islandica subsp. Cetraria islandica (L.) Ach. subfibrillosum Nyl.
crispiformis (Räsänen) Kärnefelt Chiodecton leptostictum (Nyl.) Mazosia leptosticta (Nyl.) Sparrius
Cetraria islandica subsp. islandica Cetraria islandica (L.) Ach. Zahlbr.
(L.) Ach. Chiodecton meratii Fée Syncesia effusa (Fée) Tehler
Cetraria laureri Kremp. Nephromopsis laureri (Kremp.) Chiodecton perplexum Nyl. Syncesia graphica (Fr.) Tehler
Kurok.
Chiodecton perplexum var. caesium Syncesia effusa (Fée) Tehler
Cetraria tenuifolia var. columbiana Cetraria arenaria Kärnefelt Nyl.
Räsänen Chiodecton perplexum var. pelinum Syncesia graphica (Fr.) Tehler
Cetraria tenuissima L. Cetraria aculeata (Schreb.) Fr. Nyl.
Cetrariastrum cirrhatum (Fr.) W.L. Hypotrachyna cirrhata (Fr.) Divakar, Chiodecton perplexum var. Syncesia effusa (Fée) Tehler
Culb. & C.F. Culb. A. Crespo, Sipman, Elix & Lumbsch subfibrosum auct. non Nyl.
Cetrariastrum columbiense (Zahlbr.) Hypotrachyna columbiensis (Zahlbr.) Chiodecton pterophorum Nyl. Syncesia decussans (Nyl.) Tehler
W.L. Culb. & C.F. Culb. Divakar, A. Crespo, Sipman, Elix & Chiodecton rubrocinctum (Ehrenb.) Herpothallon rubrocinctum (Ehrenb.)
Lumbsch Nyl. Aptroot, Lücking & G. Thor
Cetrariastrum dubitans Sipman Hypotrachyna dubitans (Sipman) Chiodecton sanguineum (Sw.) Vain. Herpothallon rubrocinctum (Ehrenb.)
Divakar, A. Crespo, Sipman, Elix & Aptroot, Lücking & G. Thor
Lumbsch
Chiodecton separatum Nyl. Sclerophyton syncesioides Sparrius
Cetrariastrum equadoriense (R. Hypotrachyna ecuadorensis (R. Chiodecton seriale Ach. Sclerophyton seriale (Ach.) Sparrius
Sant.) Sipman Sant.) Divakar, A. Crespo, Sipman,
Elix & Lumbsch Chrysoporthella hodgesiana Chrysoporthe hodgesiana Gryzenh.
Gryzenh. & M.J. Wingf. & M.J. Wingf. ex Chungu, Gryzenh.
Cetrariastrum sorocheilum (Vain.) Hypotrachyna sorocheila (Vain.) & M.J. Wingf.
W.L. Culb. & C.F. Culb. Divakar, A. Crespo, Sipman, Elix &
Lumbsch Chytridium schenkii var. dumontii Chytridium schenkii (P.A. Dang.)
Dogma Scherff.
Cetrariastrum vexans Zahlbr. ex Hypotrachyna vexans (Zahlbr. ex
W.L. Culb. & C.F. Culb. W.L. Culb. & C.F. Culb.) Divakar, A. Cintractia occulta Mol.-Val. Kuntzeomyces ustilaginoideus
Crespo, Sipman, Elix & Lumbsch (Henn.) Sacc. & P. Syd.
Chaenocarpus melanurus Lév. Xylaria melanura (Lév.) Sacc. Cintractia pannucea Liro Anthracoidea pannucea (Liro) Vánky
Cintractia vesiculata Mol.-Val. Ustanciosporium standleyanum
Chaenotheca carthusiae (Harm.) Chaenotheca chlorella (Ach.) Müll.
(Zundel) M. Piepenbr.
Lettau Arg.
Cladia fuliginosa Filson Rexiella fuliginosa (Filson) S.
Chaetomium murorum Corda Botryotrichum murorum (Corda)
Stenroos, Pino‐Bodas and Ahti
X.Wei Wang & Samson
Cladina arbuscula subsp. boliviana Cladonia boliviana Ahti
Chaetostigmella meliolinae Toro Phaeodimeriella meliolinae (Toro) (Ahti) Ahti
Toro
Cladina arcuata (Ahti) Ahti & Cladonia arcuata Ahti
Chapsa albomaculata (Sipman) Pseudochapsa albomaculata Follmann
Sipman & Lücking (Sipman) Parnmen, Lücking &
Lumbsch Cladina boliviana (Ahti) Ahti Cladonia boliviana Ahti
Chapsa dilatata (Müll. Arg.) Kalb Pseudochapsa dilatata (Müll. Arg.) Cladina confusa (R. Sant.) Follmann Cladonia confusa R. Sant.
Parnmen, Lücking & Lumbsch & Ahti
Chapsa phlyctidioides (Müll. Arg.) Pseudochapsa phlyctidioides (Müll. Cladina halei Ahti Cladonia halei (Ahti) Ahti & DePriest
Mangold Arg.) Parnmen, Lücking & Lumbsch Cladina peltastica Nyl. Cladonia peltastica (Nyl.) Müll. Arg.
Crocynia gossypina var. mollis (Sw.) Phyllopsora pyxinoides (Nyl.) Dendrothele cyatheae (S. Ito & S. Leptocorticium cyatheae (S. Ito & S.
Hue. Kistenich, Timdal, Bendiksby & S. Imai) N. Maek. Imai) Hjortstam & Ryvarden
Ekman Dermocybe flavotomentosa M.M. Cortinarius flavotomentosus (M.M.
Crocynia pyxinoides Nyl. Phyllopsora pyxinoides (Nyl.) Moser Moser) G. Garnier
Kistenich, Timdal, Bendiksby & S. Diaporthe manihotis Punith. Phomopsis manihot (Speg.)
Ekman Chevaug.
Cryptothecia farkasiae Lücking Amazonomyces farkasiae (Lücking) Dichonema sericeum (Sw.) Mont. Dictyonema sericeum (Sw.) Berk.
Lücking, Sérus. & G. Thor
Dichostereum granulosum (Pers.) Hyphodontia granulosa (Pers.)
Cryptothecia filicina (Ellis & Everh.) Myriostigma filicinum (Ellis & Everh.) Boidin & Lanq. Bernicchia
Lücking & G. Thor Frisch & G. Thor
Dictyonema glabratum (Spreng.) D. Cora glabrata (Spreng.) Fr.
Cryptothecia rubrocincta (Ehrenb.) Herpothallon rubrocinctum (Ehrenb.) Hawksw.
G. Thor Aptroot, Lücking & G. Thor
Dictyonema hirsutum Moncada & Cora hirsuta (Moncada & Lücking)
Cryptothecia rubrocincta (Ehrenb.) Herpothallon rubrocinctum (Ehrenb.) Lücking Moncada & Lücking
Thor Aptroot, Lücking & G. Thor
Dictyonema melvinii Chaves et al. Corella melvinii (Chaves, Lücking
Cryptothelium diplocarpum (Nyl.) Astrothelium diplocarpum Nyl. & L. Umaña) Lücking, Dal-Forno &
Zahlbr. Lawrey
Curvularia geniculata (Tracy & Cochliobolus geniculatus R.R. Dictyonema minus Lücking et al. Cora minor (Lücking, E. Navarro &
Earle) Boedijn Nelson Sipman) Lücking
Cyclomyces tabacinus (Mont.) Pat. Inonotus tabacinus (Mont.) G. Cunn. Dictyonema pavonium (Sw.) Parm. Cora glabrata (Spreng.) Fr.
Cylindrocarpon fusisporum Samuels Nectria fusispora Rossman Dictyonema zahlbruckneri (Schiffn.) Corella zahlbruckneri Schiffn.
& Brayford V. Marcano
Cylindrocladium colombiense Crous Calonectria colombiensis Crous Didymium cancellatum (Batsch) Cribraria cancellata (Batsch) Nann.-
Macbride Bremek.
Cylindrocladium graciloideum Crous Calonectria gracilipes Crous &
& Mchau Mchau Didymosphaeria arxii Aa Aaosphaeria arxii (Aa) Aptroot
Cymatella longipes G.W. Martin Marasmius martinii Singer Dimelaena diffractella (Müll. Arg.) Dimelaena tenuis (Müll. Arg.) H.
Sheard Mayrhofer & Wippel
Cyphelium leptoconium (Nyl.) Pyrgidium montellicum (Beltr.) Tibell
Zahlbr. Dimerella dilucida (Kremp.) R. Sant. Coenogonium dilucidum (Kremp.)
Kalb & Lücking
Cystocoleus niger (Huds.) Har. Cystocoleus ebeneus (Dillwyn)
Thwaites Dimerella epiphylla (Müll. Arg.) Coenogonium subluteum (Rehm)
Malme Kalb & Lücking
Cytidia habgallae (Berk. & Broome) Aleurocystis hakgallae (Berk. &
G.W. Martin Broome) G. Cunn. Dimerella fallaciosa (Müll. Arg.) Coenogonium fallaciosum (Müll.
Vězda Arg.) Kalb & Lücking
Cytospora eucalyptina Speg. Valsa fabianae G.C. Adams, M.J.
Wingf. & Jol. Roux Dimerella flavicans Vězda & Farkas Coenogonium flavicans (Vězda &
Farkas) Kalb & Lücking
Dacrymyces involutus Schwein. Arrhytidia involuta (Schwein.) Coker
Dimerella hypophylla Vězda Coenogonium hypophyllum (Vězda)
Daedalea levis Hook. Trametes elegans (Spreng.) Fr. Kalb & Lücking
Dasyscyphus bambusinus (Bres.) Erioscyphella bambusina (Bres.) Dimerella lutea (Dicks.) Trevis. Coenogonium luteum (Dicks.) Kalb
Dennis Kirschst. & Lücking
Dasyscyphus brasiliensis (Mont.) Erioscyphella brasiliensis (Mont.) Dimerella pyrophthalma (Mont.) Coenogonium pyrophthalmum
Le Gal Baral, Šandová & B. Perić Vězda (Mont.) Lücking, Aptroot & Sipman
Dasyscyphus fimbriifer (Berk. & Lachnum fimbriiferum (Berk. & M.A. Dimerella zonata (Müll. Arg.) R. Coenogonium zonatum (Müll. Arg.)
M.A. Curtis) Sacc. Curtis) J.H. Haines Sant. Kalb & Lücking
Dasyscyphus nudipes (Fuckel) Sacc. Lachnum nudipes (Fuckel) Nannf. Diploschistes muscorum subsp. Diploschistes bartlettii (Lumbsch)
var. nudipes bartletii Lumbsch Lücking
Dasyscyphus varians Rehm Lachnum varians (Rehm) M.P. Dirinaria confusa var. saxicola Dirinaria confusa D.D. Awasthi
Sharma (Räsänen) D.D. Awasthi
Everniastrum columbiense (Zahlbr.) Hypotrachyna columbiensis (Zahlbr.) Galerina cerina var. nebularum A. Galerina cerina A.H. Sm. & Singer
Sipman Divakar, A. Crespo, Sipman, Elix & H. Sm. & Singer
Lumbsch Galerina cerina var. nebularum A.H. Galerina cerina A.H. Sm. & Singer
Everniastrum fragile Sipman Hypotrachyna fragilis (Sipman) Sm. & Singer
Divakar, A. Crespo, Sipman, Elix & Galerina emmetensis var. Galerina emmetensis A.H. Sm. &
Lumbsch intermedia A.H. Sm. & Singer Singer
Everniastrum limaeforme (Taylor) Hypotrachyna limiformis (Taylor) Ganoderma fornicatum (Fr.) Pat. Ganoderma orbiforme (Fr.)
Hale ex Sipman Divakar, A. Crespo, Sipman, Elix & Ryvarden
Lumbsch
Ganoderma hollidayi Steyaert Ganoderma chalceum (Cooke)
Everniastrum lipidiferum (Hale & Hypotrachyna lipidifera (Hale & M. Steyaert
Wirth) Sipman Wirth) Divakar, A. Crespo, Sipman,
Geastrum trichifer Rick Geastrum mirabile Mont.
Elix & Lumbsch
Gerronema cheilocystidiatum Singer Omphalina cheilocystidiata (Singer)
Everniastrum planum Sipman Hypotrachyna plana (Sipman)
Raithelh.
Divakar, A. Crespo, Elix & Lumbsch
Gloeocystidium ochroleucum Bres. Scytinostroma lusitanicum (Trotter)
Everniastrum sorocheilum (Vain.) Hypotrachyna sorocheila (Vain.)
& Torrend P.M. Kirk
Sipman Divakar, A. Crespo, Sipman, Elix &
Lumbsch Gloeosporium vanillae Cooke Colletotrichum coccodes (Wallr.) S.
Hughes
Everniastrum subvexans Sipman Hypotrachyna subvexans (Sipman)
Divakar, A. Crespo, Sipman, Elix & Glomus manihotis R.H. Howeler, Rhizophagus manihotis (R.H.
Lumbsch Sieverd. & N.C. Schenck Howeler, Sieverd. & N.C. Schenck)
C. Walker & A. Schüßler
Everniastrum vexans (W.L. Culb. & Hypotrachyna vexans (Zahlbr. ex
C.F. Culb.) Sipman W.L. Culb. & C.F. Culb.) Divakar, A. Glossodium aversum Nyl. Icmadophila aversa (Nyl.) Rambold
Crespo, Sipman, Elix & Lumbsch & Hertel
Glyphis favulosa Ach. Glyphis cicatricosa Ach. Graphina plurispora (Redinger) Allographa plurispora (Redinger)
Wirth & Hale Lücking & Kalb
Glyphis favulosa var. intermedia Glyphis cicatricosa Ach.
Müll. Arg. Graphina pseudoanaloga (Vain.) Graphis pseudoanaloga Vain.
Zahlbr.
Glyphis labyrinthica Ach. Sarcographa labyrinthica (Ach.)
Müll. Arg. Graphina reniformis (Fée) Müll. Arg. Diorygma reniforme (Fée) Kalb,
Staiger & Elix
Glyphis medusulina Nyl. Sarcographa medusulina (Nyl.) Müll.
Arg. Graphina scribillans (Nyl.) Zahlbr. Thalloloma scribillans (Nyl.) Lücking
Gnomonia ospinae Chardón Apiosphaeria guaranitica (Speg.) Graphina sophistica (Nyl.) Müll. Arg. Graphis platycarpa Eschw.
Höhn. Graphina streblocarpa (Bél.) Müll. Graphis streblocarpa (Bél.) Nyl.
Gomphillus ophiosporus Kalb & Gomphillus hyalinus (Pat.) Lücking, Arg.
Vězda Kalb & Vězda Graphina subserpentina (Nyl.) Müll. Graphis subserpentina Nyl.
Grammothele setulosa (Henn.) Ryv. Megasporoporia setulosa (Henn.) Arg.
Rajchenb. Graphina tetraphora (Nyl.) Zahlbr. Acanthothecis tetraphora (Nyl.)
Graphina acharii (Fée) Müll. Arg. Allographa acharii (Fée) Lücking & Staiger & Kalb
Kalb Graphina triphora (Nyl.) Müll. Arg. Allographa triphora (Nyl.) Lücking
Graphina acharii f. enteroleuca Allographa acharii (Fée) Lücking & & Kalb
(Ach.) Zahlbr. Kalb Graphina virginea (Eschw.) Müll. Diorygma poitaei (Fée) Kalb, Staiger
Graphina acharii f. subducens (Nyl.) Allographa acharii (Fée) Lücking & Arg. & Elix
Zahlbr. Kalb Graphiopsis chlorocephala (Fresen.) Dichocladosporium chlorocephalum
Graphina acharii var. albicans (Nyl.) Allographa acharii (Fée) Lücking & Trail (Fresen.) K. Schub., U. Braun &
Müll. Arg. Kalb Crous
Graphina acharii var. monospora Allographa acharii (Fée) Lücking & Graphis abapha Nyl. Diorygma hieroglyphicum (Pers.)
(Nyl.) Müll. Arg. Kalb Staiger & Kalb
Graphina aggregans (Nyl.) Zahlbr. Anomomorpha aggregans (Nyl.) Graphis acharii Fée Allographa acharii (Fée) Lücking &
Staiger Kalb
Graphina agminalis (Nyl.) Zahlbr. Jocatoa agminalis (Nyl.) Lücking, Graphis adpressa Vain. Allographa adpressa (Vain.) Lücking
Herrera-Camp. & R. Miranda & Kalb
Graphina analoga var. subradiata Allographa subradiata (Nyl.) Lücking Graphis afzelii Ach. Dyplolabia afzelii (Ach.) A. Massal.
(Nyl.) Zahlbr. & Kalb Graphis aggregans Nyl. Anomomorpha aggregans (Nyl.)
Graphina anguina (Mont.) Müll. Arg. Thalloloma anguinum (Mont.) Trevis. Staiger
Graphina chlorocarpa (Fée) Müll. Allographa chlorocarpa (Fée) Graphis agminalis Nyl. Jocatoa agminalis (Nyl.) Lücking,
Arg. Lücking & Kalb Herrera-Camp. & R. Miranda
Graphina chrysocarpa (Raddi) Müll. Allographa chrysocarpa (Raddi) Graphis alborosella Nyl. Chapsa alborosella (Nyl.) Frisch
Arg. Lücking & Kalb Graphis analoga var. monophora Graphis analoga Nyl.
Graphina cleistomma (Nyl.) Müll. Allographa cleistomma (Nyl.) Tuck.
Arg. Lücking & Kalb Graphis analoga var. subradiata Nyl. Allographa subradiata (Nyl.) Lücking
Graphina collospora (Vain.) Zahlbr. Diorygma hieroglyphicum (Pers.) & Kalb
Staiger & Kalb Graphis analoga var. subtecta (Nyl.) Graphis subtecta (Nyl.) Lücking
Graphina columbiana (Nyl.) Müll. Diorygma hieroglyphicum (Pers.) Zahlbr.
Arg. Staiger & Kalb Graphis analoga var. subtecta Nyl. Graphis subtecta (Nyl.) Lücking
Graphina dimidiata (Vain.) Zahlbr. Graphis dimidiata Vain. Graphis anguilliformis Taylor Allographa rhizicola (Fée) Lücking
Graphina haemographa (Nyl.) Müll. Thalloloma haemographum (Nyl.) & Kalb
Arg. Staiger Graphis angustata Eschw. Allographa angustata (Eschw.)
Graphina hololeucoides (Nyl.) Müll. Acanthothecis hololeucoides (Nyl.) Lücking & Kalb
Arg. Staiger & Kalb Graphis bettinae Lücking, L. Allographa bettinae (Lücking,
Graphina incrustans (Fée) Müll. Arg. Fissurina incrustans Fée Umaña, Chaves & Sipman Umaña, Chaves & Sipman) Lücking
& Kalb
Graphina insculpta (Eschw.) Müll. Fissurina insculpta Mont.
Arg. Graphis cabbalistica Nyl. Kalbographa cabbalistica (Nyl.)
Lücking
Graphina malmei Redinger Malmographina plicosa (C.F.W.
Meissn.) M. Cáceres, Rivas Plata Graphis chlorocarpa Fée Allographa chlorocarpa (Fée)
& Lücking Lücking & Kalb
Graphina marcescens (Fée) Müll. Carbacanthographis marcescens Graphis chrysenteron Mont. Pallidogramme chrysenteron (Mont.)
Arg. (Fée) Staiger & Kalb Staiger, Kalb & Lücking
Hypocrea muroiana f. Hypocrea muroiana I. Hino & Lachnum brasiliense (Mont.) J.H. Erioscyphella brasiliensis (Mont.)
dimorphospora Yoshim. Doi Katum. Haines & Dumont Baral, Šandová & B. Perić
Hypocrea muroiana f. minor Hypocrea muroiana I. Hino & Lachnum cerinum (Pers.) Nannf. Neodasyscypha cerina (Pers.)
Yoshim. Doi Katum. Spooner
Hypocrea patella f. tropica Yoshim. Trichoderma patellotropicum Lachnum fuscescens (Pers.) P. Lachnum fuscescens (Pers.) P.
Doi Samuels Karst. var. fuscescens Karst.
Hypocrea subtrachycarpa Yoshim. Trichoderma subtrachycarpum Lachnum nudipes var. minor Dennis Lachnum nudipes (Fuckel) Nannf.
Doi (Yoshim. Doi) Jaklitsch & Voglmayr Lachnum sclerotii (A.L. Sm.) J.H. Erioscyphella sclerotii (A.L. Sm.)
Hypotrachyna aguirrei Sipman, Elix Remototrachyna aguirrei (Sipman, Haines & Dumont Baral, Šandová & B. Perić
& T.H. Nash Elix & T.H. Nash) Flakus, Kukwa & Lactarius annulifer Singer Lactifluus annulifer (Singer)
Sipman Nuytinck
Hypotrachyna costaricensis (Nyl.) Remototrachyna costaricensis (Nyl.) Lactarius brasiliensis Singer Lactifluus brasiliensis (Singer) Silva-
Hale Divakar, Lumbsch, Ferencová, Filho & Wartchow
Prado & A. Crespo
Lactarius deceptivus Peck Lactifluus deceptivus (Peck) Kuntze
Hypotrachyna formosana (Zahlbr.) Hypotrachyna osseoalba (Vain.) Y.S.
Hale Park & Hale Lactarius fragilis (Burl.) Hesler & Lactarius camphoratus (Bull.) Fr.
A.H. Sm.
Hypotrachyna gigas (Kurok.) Hale Hypotrachyna longiloba (H. Magn.)
C.W. Sm. Laeticorticium roseum (Pers.) Donk Corticium roseum Pers.
Hypotrachyna rachista (Hale) Hale Hypotrachyna prolongata (Kurok.) Lagarobasidium detriticum (Bourdot Xylodon detriticus (Bourdot) K.H.
Hale & Galzin) Jülich Larss., Viner & Spirin
Hypotrachyna rhabdiformis (Kurok.) Remototrachyna rhabdiformis Laurera effusa Aptroot & Sipman Astrothelium effusum (Aptroot &
Hale (Kurok.) Divakar & A. Crespo Sipman) Aptroot & Lücking
Lecanactis proximans (Nyl.) Zahlbr. Cresponea proximata (Nyl.) Egea & Lecanora inaequata Nyl. Ochrolechia inaequata (Nyl.) Zahlbr.
Torrente Lecanora insperata Nyl. Orcularia insperata (Nyl.) Kalb &
Lecania heterochroa Müll. Arg. Badimia dimidiata (Bab. ex Leight.) Giralt
Vězda Lecanora laciniosa var. substellata Candelaria concolor (Dicks.) Arnold
Lecania sulphureofusca (Fée) Müll. Sipmaniella sulfureofusca (Fée) (Ach.) Nyl.
Arg. Kalb Lecanora multifera (Nyl.) Vain. Neoprotoparmelia multifera (Nyl.)
Lecanicillium sabanense J.S. Akanthomyces sabanensis (Chir.- Garima Singh, Lumbsch & I.
Chirivi-Salomon, Restrepo & T.I. Salom., Restrepo & Sanjuan) Chir.- Schmitt
Sanjuan Salom., Sanjuan & Restrepo Lecanora multifera Nyl. Neoprotoparmelia multifera (Nyl.)
Lecanora albella var. chlarona Lecanora chlarotera Nyl. Garima Singh, Lumbsch & I.
(Ach.) Nyl. Schmitt
Lecanora albella var. praeferenda Lecanora praeferenda (Nyl.) Nyl. Lecanora murorum var. obliterarum Calogaya saxicola (Hoffm.) Vondrák
Nyl. (Pers.) Nyl.
Lecanora alboatrata Nyl. Lopezaria versicolor (Flot.) Kalb & Lecanora pyracea (Ach.) Nyl. Athallia pyracea (Ach.) Arup, Frödén
Hafellner & Søchting
Lecanora angulosa f. chlarona Lecanora chlarotera Nyl. Lecanora quadrilocularis Nyl. Caloplaca quadrilocularis (Nyl.)
(Ach.) Hue Zahlbr.
Lecanora atra (Huds.) Ach. Tephromela atra (Huds.) Hafellner Lecanora russeola Nyl. Caloplaca russeola (Nyl.) Zahlbr.
Lecanora aurantiaca f. erythrella Gyalolechia flavovirescens (Wulfen) Lecanora subaeruginosa (Nyl.) Nyl. Vainionora flavovirens (Fée) Kalb
(Ach.) Nyl. Søchting, Frödén & Arup Lecanora subferruginea Nyl. Blastenia subferruginea (Nyl.) Müll.
Lecanora aurantiaca var. erythrella Gyalolechia flavovirescens (Wulfen) Arg.
(Ach.) Nyl. Søchting, Frödén & Arup Lecanora subfusca f. chlarona Lecanora chlarotera Nyl.
Lecanora brebissonii (Fée) Nyl. Caloplaca brebissonii (Fée) J. Sant. (Ach.) Körb.
ex Hafellner & Poelt Lecanora subfusca var. argentata Lecanora argentata (Ach.) Röhl.
Lecanora chlarona (Ach.) Nyl. Lecanora chlarotera Nyl. (Ach.) Boistel
Lecanora coarctata (Sm.) Ach. Trapelia coarctata (Turner) M. Lecanora subfusca var. atrynea Lecanora chlarotera Nyl.
Choisy Ach.
Lecanora colobinoides Nyl. Rinodina colobinoides (Nyl.) Müll. Lecanora subfusca var. chlarona Lecanora chlarotera Nyl.
Arg. (Ach.) Ach.
Lecanora conjugens Nyl. Caloplaca conjungens (Nyl.) Zahlbr. Lecanora subfusca var. conferta Myriolecis hagenii (Ach.) Śliwa,
(Duby) Schaer. Zhao Xin & Lumbsch
Lecanora conjungens Nyl. Caloplaca conjungens (Nyl.) Zahlbr.
Lecanora subfusca var. Lecanora tropica Zahlbr.
Lecanora crocantha Nyl. Caloplaca crocantha (Nyl.) Zahlbr.
subcrenulata Nyl.
Lecanora diducta f. albicans Nyl. Caloplaca diducta (Nyl.) Zahlbr.
Lecanora subfusca var. Lecanora subgranulata (Nyl.) Nyl.
Lecanora diducta Nyl. Caloplaca diducta (Nyl.) Zahlbr. subgranulata Nyl.
Lecanora dimidiata Bab. Badimia dimidiata (Bab. ex Leight.) Lecanora subimmersa (Fée) Vain. Lecanora subimmersa (Fée) Vain.
Vězda subsp. subimmersa
Lecidea conspersa Fée Piccolia conspersa (Fée) Hafellner Lecidea millegrana f. Bacidia millegrana (Taylor) Zahlbr.
fusconigrescens Nyl.
Lecidea conspirans Nyl. Buellia conspirans (Nyl.) Vain.
Lecidea parasema Ach. Lecidella elaeochroma (Ach.) M.
Lecidea contigua f. platycarpa Porpidia macrocarpa (DC.) Hertel & Choisy
(Lam.) Nyl. A.J. Schwab
Lecidea parasema var. elaeochroma Lecidella elaeochroma (Ach.) M.
Lecidea contigua var. platycarpa Porpidia macrocarpa (DC.) Hertel & Ach. Choisy
(Lam.) Fr. A.J. Schwab
Lecidea parasema var. entereoleuca Lecidella elaeochroma (Ach.) M.
Lecidea crocantha (Nyl.) Hue Caloplaca crocantha (Nyl.) Zahlbr. Nyl. Choisy
Lecidea cupreorosella var. Aquacidia trachona (Ach.) Aptroot Lecidea parmelioides Hook. Coccocarpia erythroxyli (Spreng.)
chloroticoides Nyl. Swinscow & Krog
Lecidea demutans Nyl. Malmidea demutans (Nyl.) Lücking Lecidea parvifolia Pers. Phyllopsora parvifolia (Pers.) Müll.
Lecidea domingensis f. inexplicata Letrouitia flavidula (Tuck.) Hafellner Arg.
(Nyl.) Hue Lecidea parvifolia var. breviuscula Phyllopsora breviuscula (Nyl.) Müll.
Lecidea domingensis var. Letrouitia flavidula (Tuck.) Hafellner (Nyl.) Nyl. Arg.
inexplicata Nyl. Lecidea parvifolia var. corallina Phyllopsora parvifolia (Pers.) Müll.
Tuck. Arg.
Lecidea endochroma (Fée) Nyl. Catillochroma endochromum (Fée)
Kalb Lecidea parvifolia var. subgranulosa Phyllopsora parvifolia (Pers.) Müll.
Tuck. Arg.
Lecidea endoleuca (Nyl.) Nyl. Bacidia laurocerasi (Delise ex Duby)
Zahlbr. Lecidea perminima Nyl. Catillaria perminima (Nyl.) Zahlbr.
Lecidea erythrantha (Tuck.) Hue Caloplaca erythrantha (Tuck.) Lecidea petraea Ach. Rhizocarpon concentricum (Davies)
Zahlbr. Beltr.
Lobaria pallida (Hook.) Trevis. Lobariella pallida (Hook.) Moncada Marasmiellus subcoracinus (Berk. Marasmiellus subcoracinus (Berk. &
& Lücking & M.A. Curtis) Singer subsp. M.A. Curtis) Singer
subcoracinus
Lobaria patinifera (Taylor) Hue Emmanuelia patinifera (Taylor)
Lücking, M. Cáceres & Ant. Simon Marasmius androsaceus (Bolton) Fr. Gymnopus androsaceus (L.) Della
Magg. & Trassin.
Lobaria patinifera (Taylor) Müll. Arg. Emmanuelia patinifera (Taylor)
Lücking, M. Cáceres & Ant. Simon Marasmius copelandii Peck Mycetinis copelandii (Peck) A.W.
Wilson & Desjardin
Lobaria peltigera (Delise) Vain. Yoshimuriella peltigera (Vain.)
Lücking & Moncada Marasmius corrugatus var. Gymnopus aurantiacus Murrill
aurantiacus (Murrill) Singer
Lobaria subdissecta (Nyl.) Vain. Yoshimuriella subdissecta (Nyl.)
Moncada & Lücking Marasmius floriceps Berk. & M.A. Marasmius leoninus Berk.
Curtis
Lobaria subexornata Yoshim. Lobariella subexornata (Yoshim.)
Yoshim. Marasmius gilvus Pat. Marasmiellus gilvus (Pat.) Singer
Lobaria tenuis Vain. Emmanuelia tenuis (Vain.) Lücking, Marasmius haematocephalus Marasmius haematocephalus
Moncada & Gumboski (Mont.) Fr. var. haematocephalus (Mont.) Fr.
Loflammea flammea (Müll. Arg.) Loflammia epiphylla (Fée) Lücking Marasmius haematocephalus var. Marasmius haematocephalus
Vězda & Vězda leucophyllus Singer (Mont.) Fr.
Lopadium cytharinum (Nyl.) Zahlbr Calopadia lecanorella (Nyl.) Kalb & Marasmius haematocephalus var. Marasmius haematocephalus
Vězda pseudotageticolor Singer (Mont.) Fr.
Lopadium cyttarinum (Nyl.) Zahlbr. Calopadia lecanorella (Nyl.) Kalb & Marasmius pallidocinctus var. Marasmius pallidocinctus Singer
Vězda latisporus Singer
Lopadium newtonianum (Henriq.) Eugeniella newtoniana (Henriq.) Marasmius pseudoniveus Singer Marasmius pseudoniveus Singer
R. Sant. Lücking, Sérus. & Kalb var. pseudoniveus
Lopharia crassa (Lév.) Boidin Phlebiopsis crassa (Lév.) Floudas & Marasmius pseudoniveus var. Marasmius pseudoniveus Singer
Hibbett amylocystis Singer
Marasmius variabiliceps var. Marasmius variabiliceps Singer
Lopharia novae-granatae A.L. Hjortstamia novae-granatae (A.L.
separatus Singer
Welden Welden) Hjortstam & Ryvarden
Maronea multifera (Nyl.) Vain. Neoprotoparmelia multifera (Nyl.)
Lopharia papyracea (Jungh.) D.A. Phlebiopsis friesii (Lév.) Spirin &
Garima Singh, Lumbsch & I.
Reid Miettinen
Schmitt
Lopharia papyrina (Mont.) Boidin Phlebiopsis papyrina (Mont.)
Maronina multifera (Nyl.) Hafellner Neoprotoparmelia multifera (Nyl.)
Miettinen & Spirin
& R.W. Rogers Garima Singh, Lumbsch & I.
Lycoperdon pusillum Batsch Bovista pusilla (Batsch) Pers. Schmitt
Lycoperdon umbrinum Pers. var. Lycoperdon umbrinum Pers. Marssonia agaves Earle Marssonina agaves (Earle) Magnus
umbrinum
Mazosia rotula var. granularis (Müll. Mazosia melanophthalma (Müll.
Macabuna petreae Pardo-Card. Catenulopsora petreae Pardo-Card. Arg.) Müll. Arg. Arg.) R. Sant.
Megalospora sulphurata var. Megalospora sulphurata Meyen Mycoporum pycnocarpum Nyl. Mycoporum compositum (A.
nigricans (Müll. Arg.) Riddle Massal.) R.C. Harris
Megalospora versicolor (Fée) Lopezaria versicolor (Flot.) Kalb & Mycosphaerella flexuosa Crous & Pseudoteratosphaeria flexuosa
Zahlbr. Hafellner M.J. Wingf. (Crous & M.J. Wingf.) Quaedvl. &
Crous
Megalospora versicolor var. Lopezaria versicolor (Flot.) Kalb &
dichroma (Fée) Riddle Hafellner Mycosphaerella perpendicularis Pseudoteratosphaeria
Crous & M.J. Wingf. perpendicularis (Crous & M.J.
Megasporoporiella cavernulosa Cerioporus cavernulosus (Berk.) Wingf.) Quaedvl. & Crous
(Berk.) B.K. Cui, Y.C. Dai & Hai J. Li Zmitr.
Mycosphaerella scytalidii Crous & Phaeophleospora scytalidii (Crous &
Melanographa chionographa (Nyl.) Melaspilea chionographa (Nyl.) M.J. Wingf. M.J. Wingf.) Quaedvl. & Crous
Müll. Arg. Zahlbr.
Myriotrema bahianum (Ach.) Hale Ocellularia bahiana (Ach.) Frisch
Melanographa diplasiospora (Nyl.) Melaspilea diplasiospora (Nyl.) Müll.
Müll. Arg. Arg. Myriotrema calvescens (Fée) Hale Ocellularia calvescens (Fée) Müll.
Arg.
Melanographa interalbicans (Nyl.) Melaspilea interalbicans (Nyl.) Müll.
Müll. Arg. Arg. Myriotrema compunctum (Ach.) Leucodecton occultum (Eschw.)
Hale Frisch
Melanotheca achariana Fée Pyrenula anomala (Ach.) Vain.
Myriotrema dactyliferum (Hale) Leucodecton dactyliferum (Hale)
Melanotheca aciculifera Nyl. Celothelium aciculiferum (Nyl.) Vain.
Gyeln. Lücking
Melanotheca arthonioides (Eschw.) Pyrenula arthoniotheca Upreti
Myriotrema glaucophaenum Glaucotrema glaucophaenum
Müll. Arg.
(Kremp.) Zahlbr. (Kremp.) Rivas Plata & Lumbsch
Meliolina chardonii Toro Asteridiella chardonii (Toro) S.
Hughes Myriotrema myriocarpum (Fée) Hale Austrotrema myriocarpum (Fée) I.
Medeiros, Lücking & Lumbsch
Meruliopsis corium (Pers.) Ginns Byssomerulius corium (Pers.)
Parmasto Myriotrema pachystomum (Nyl.) Fibrillithecis pachystoma (Nyl.)
Hale Sipman
Merulius tremellosus Schrad. Phlebia tremellosa (Schrad.)
Nakasone & Burds. Myriotrema pycnoporellum (Nyl.) Pycnotrema pycnoporellum (Nyl.)
Hale Rivas Plata & Lücking
Micromphale brevipes (Berk. & Gymnopus neobrevipes R.H.
Ravenel) Singer Petersen Myriotrema subcompunctum (Nyl.) Leucodecton subcompunctum (Nyl.)
Hale Frisch
Mollisia caesia var. andina Dennis Mollisia caesia (Fuckel) Sacc.
Myriotrema subwrightii (Hale) Hale Stegobolus anamorphus (Nyl.)
Morganella fulginea Lycoperdon fuligineum Berk. & M.A. Frisch & Kalb
Curtis
Myriotrema terebratulum (Nyl.) Hale Myriotrema clandestinum (Fée) Hale
Morispora tayronensis Salazar- Gerwasia tayronensis Salazar-
Yepes, Pardo-Card. & Buriticá Yepes, Pardo-Card. & Buriticá Myriotrema wightii (Taylor) Hale Sanguinotrema wightii (Taylor)
Lücking
Mortierella ornata W. Gams Dissophora ornata (W. Gams) W.
Gams Myxomycidium flavum G.W. Martin Gloeomucro flavus (G.W. Martin)
Lepidostroma calocerum (G.W. R.H. Petersen
Multiclavula calocera (G.W. Martin)
R.H. Petersen Martin) Oberw. Naematoloma subviride (Berk. & Hypholoma fasciculare (Huds.) P.
Muscodor suturae Kudalkar, Strobel Induratia suturae (Kudalkar, M.A. Curtis) A.H. Sm. Kumm.
& Riy.-Ul-Hass. Strobel & Riy.-Ul-Hass.) Samarak., Naematoloma udum (Pers.) P. Bogbodia uda (Pers.) Redhead
Thongbai, K.D. Hyde & M. Stadler Karst.
Mutatoderma brunneocontextum Hyphoderma romeroae C.E. Gómez, Nectria consors (Ellis & Everh.) Volutellonectria consors (Ellis &
C.E. Gómez Baltazar & Rajchenb. Seaver Everh.) J. Luo, X.M. Zhang & W.Y.
Mycena margarita Murrill Collybia margarita (Murrill) Singer Zhuang
Mycomicrothelia apposita (Nyl.) D. Bogoriella apposita (Nyl.) Aptroot & Nectria pachyderma Rossman Hydropisphaera pachyderma
Hawksw. Lücking (Rossman) Rossman & Samuels
Mycomicrothelia captiosa (Kremp.) Bogoriella captiosa (Kremp.) Nectria parviphialis Samuels Bionectria parviphialis (Samuels)
D.Hawksw. Aptroot & Lücking Schroers
Mycomicrothelia exigua (Müll. Arg.) Bogoriella exigua (Müll. Arg.) Nemania serpens (Pers.) Gray var. Nemania serpens (Pers.) Gray
D.Hawksw. Aptroot & Lücking serpens
Mycomicrothelia hemispherica Bogoriella hemisphaerica (Müll. Neobarya usneae Etayo Lichenobarya usneae (Etayo) Etayo,
(Müll. Arg.) D. Hawksw. Arg.) Aptroot & Lücking Diederich & Lawrey
Mycomicrothelia thelena (Ach.) D. Bogoriella thelena (Ach.) Aptroot & Neuropogon sphacelatus (R. Br.) Usnea sphacelata R. Br.
Hawksw. Lücking D.J. Galloway
Ocellularia microsporum (Zahlbr.) Redingeria microspora (Zahlbr.) M. Opegrapha varia Pers. Alyxoria varia (Pers.) Ertz & Tehler
Hale Cáceres & Lücking Opegrapha viridipruinosa Coppins Alyxoria viridipruinosa (Coppins &
Ocellularia myriotremoides (Nyl.) Myriotrema myriotremoides (Nyl.) & Yahr Yahr) Ertz
Zahlbr. Hale Opegrapha viridis (Ach.) Ach. Zwackhia viridis (Ach.) Poetsch &
Ocellularia recondita (Stirt.) Zahlbr. Rhabdodiscus reconditus (Stirt.) Schied.
Rivas Plata, Lücking & Lumbsch Opegrapha viridis (Ach.) Behlen & Zwackhia viridis (Ach.) Poetsch &
Ocellularia remanens (Nyl.) Müll. Myriotrema clandestinum (Fée) Hale Desberger Schied.
Arg. Ophiocordyceps amazonica (Henn.) Paraisaria amazonica (Henn.)
Ocellularia sinuosa Sipman Gyrotrema sinuosum (Sipman) G.H. Sung, J.M. Sung, Hywel-Jones Luangsa-ard, Mongkols. & Samson
Frisch & Spatafora
Ocellularia subemersa Müll. Arg. Rhabdodiscus subemersus (Müll. Ophiocordyceps blattarioides Paraisaria blattarioides (Sanjuan &
Arg.) Rivas Plata, Lücking & Sanjuan & Spatafora Spatafora) Luangsa-ard, Mongkols.
Lumbsch & Samson
Ocellularia terebratula (Nyl.) Müll. Myriotrema clandestinum (Fée) Hale Oropogon loxensis var. atroalbicans Oropogon loxensis (Fée) Zukal
Arg. (Nyl.) Hue
Omphalina foliacea P.M. Jørg. Agonimia foliacea (P.M. Jørg.) Oudemansiella glutinosa Singer Dactylosporina glutinosa (Singer)
Lücking & Moncada R.H. Petersen
Pannoparmelia hypotrachyna (Nyl.) Remototrachyna costaricensis (Nyl.) Parmelia densirhizinata Kurok. Hypotrachyna densirhizinata
R. Sant. Divakar, Lumbsch, Ferencová, (Kurok.) Hale
Prado & A. Crespo Parmelia dilatata Vain. Parmotrema dilatatum (Vain.) Hale
Pannularia nigrocincta (Mont.) Nyl. Parmeliella nigrocincta (Mont.) Müll. Parmelia dominicana Vain. Parmotrema dominicanum (Vain.)
Arg. Hale
Paraphelaria colombiana A.L. Aphelariopsis colombiana (A.L. Parmelia eborina Hale Parmotrema eborinum (Hale) Hale
Welden Welden) Jülich
Parmelia endorubra f. Hypotrachyna microblasta (Vain.)
Parathelium indutum Nyl. Pyrenula adacta Fée imbricatiformis Gyeln. Hale
Parathelium polysemum Nyl. Lithothelium polysemum (Nyl.) Parmelia endosulphurea (Hillmann) Parmotrema endosulphureum
Aptroot Hale (Hillmann) Hale
Parmelia abstrusa Vain. Relicina abstrusa (Vain.) Hale Parmelia fasciculata Vain. Parmotrema fasciculatum (Vain.)
Parmelia acanthifolia Pers. Parmotrema cetratum (Ach.) Hale Hale
Parmelia andina Müll. Arg. Parmotrema andinum (Müll. Arg.) Parmelia flaventior Stirt. Flavopunctelia flaventior (Stirt.) Hale
Hale
Parmelia flavescens (Kremp.) Nyl. Parmotrema flavescens (Kremp.)
Parmelia andreana Müll. Arg. Flavopunctelia flaventior (Stirt.) Hale Hale
Parmelia angustior Nyl. Imshaugia angustior (Nyl.) Sipman Parmelia flavotincta Hale Parmotrema flavotinctum (Hale)
Parmelia atrichella Nyl. Bulbothrix atrichella (Nyl.) Hale Hale
Parmelia aurata (Ach.) Eschw. Crocodia aurata (Ach.) Link Parmelia fracta Hale Parmotrema fractum (Hale) Hale
Parmelia austrosinensis Zahlbr. Parmotrema austrosinense (Zahlbr.) Parmelia gigas Kurok. Hypotrachyna longiloba (H. Magn.)
Hale C.W. Sm.
Parmelia bangii Vain. Parmotrema bangii (Vain.) Hale Parmelia granatensis Nyl. Bulbothrix goebelii (Zenker) Hale
Parmelia blanchetii Hue Parmotrema blanchetianum (Müll. Parmelia granulosa Vain. Hypotrachyna columbiensis (Zahlbr.)
Arg.) Kalb Divakar, A. Crespo, Sipman, Elix &
Lumbsch
Parmelia bogotensis Vain. Hypotrachyna bogotensis (Vain.)
Hale Parmelia hypotrachyna Nyl. Remototrachyna costaricensis (Nyl.)
Divakar, Lumbsch, Ferencová,
Parmelia borreri (Sm.) Turner Punctelia borreri (Turner) Krog Prado & A. Crespo
Parmelia borreri f. rudecta (Ach.) Punctelia rudecta (Ach.) Krog
Parmelia kamtschadalis var. Hypotrachyna vexans (Zahlbr. ex
Nyl.
americana f. tenuis Müll. Arg. W.L. Culb. & C.F. Culb.) Divakar, A.
Parmelia caperata (L.) Ach. Flavoparmelia caperata (L.) Hale Crespo, Sipman, Elix & Lumbsch
Parmelia caperata f. ramealis Nyl. Flavoparmelia caperata (L.) Hale Parmelia laevigata (Sm.) Ach. Hypotrachyna laevigata (Sm.) Hale
Parmelia microblasta Vain. Hypotrachyna microblasta (Vain.) Parmelia zollingeri Hepp Parmotrema zollingeri (Hepp) Hale
Hale Parmeliella incrassata P.M. Jørg. Nebularia incrassata (P.M. Jørg.)
Parmelia microspora Müll. Arg. Xanthoparmelia microspora (Müll. P.M. Jørg.
Arg.) Hale Parmeliella mariana (Fr.) P.M. Jørg. Lepidocollema marianum (Fr.) P.M.
Parmelia mougeotii Schaer. Xanthoparmelia mougeotii (Schaer. & D.J. Galloway Jørg.
ex D. Dietr.) Hale Parmelina cleefi Sipman Hypotrachyna cleefii (Sipman)
Parmelia negata Nyl. Punctelia negata (Nyl.) Krog Divakar, A. Crespo, Sipman, Elix &
Lumbsch
Parmelia osteoleuca Nyl. Hypotrachyna osteoleuca (Nyl.) Hale
Parmelina lindmanii (Lynge) Hale Myelochroa lindmanii (Lynge) Elix
Parmelia perforata (Jacq.) Ach. Parmotrema perforatum (Jacq.) A. & Hale
Massal.
Parmelina melanochaeta (Kurok.) Parmotrema melanochaetum
Parmelia perlata (Huds.) Ach. Parmotrema perlatum (Huds.) M. Hale (Kurok.) O. Blanco, A. Crespo,
Choisy Divakar, Elix & Lumbsch
Parmelia perlata f. sorediifera Müll. Parmotrema reticulatum (Taylor) M. Parmelina minarum (Vain.) Hale Hypotrachyna minarum (Vain.) Krog
Arg. Choisy & Swinscow
Parmelia perlata var. ciliata Duby Parmotrema reticulatum (Taylor) M. Parmelina muelleri (Vain.) Hale Parmotrema muelleri (Vain.) O.
Choisy Blanco, A. Crespo, Divakar, Elix &
Parmelia physodalica Hale Hypotrachyna physodalica (Hale) Lumbsch
Hale Parmelina spumosa (Asahina) Hale Hypotrachyna spumosa (Asahina)
Parmelia prolongata Kurok. Hypotrachyna prolongata (Kurok.) Krog & Swinscow
Hale Parmelina swinscowii (Hale) Hale Hypotrachyna swinscowii (Hale)
Parmelia propagulifera Vain. Hypotrachyna microblasta (Vain.) Krog & Swinscow
Hale Parmelinopsis cleefii (Sipman) V. Hypotrachyna cleefii (Sipman)
Parmelia rachista Hale Hypotrachyna prolongata (Kurok.) Marcano & Sipman Divakar, A. Crespo, Sipman, Elix &
Hale Lumbsch
Parmelia rampoddensis Nyl. Parmotrema rampoddense (Nyl.) Parmelinopsis horrescens (Taylor) Hypotrachyna horrescens (Taylor)
Hale Elix & Hale Krog & Swinscow
Parmelia reducens Nyl. Hypotrachyna reducens (Nyl.) Hale Parmelinopsis melanochaeta Parmotrema melanochaetum
(Kurok.) Elix & Hale (Kurok.) O. Blanco, A. Crespo,
Parmelia relicina var. coronata (Fée) Bulbothrix coronata (Fée) Hale Divakar, Elix & Lumbsch
Nyl.
Parmelinopsis minarum (Vain.) Elix Hypotrachyna minarum (Vain.) Krog
Parmelia reticulata Taylor Parmotrema reticulatum (Taylor) M. & Hale & Swinscow
Choisy
Parmelinopsis spumosa (Asah.) Elix Hypotrachyna spumosa (Asahina)
Parmelia revoluta Flörke Hypotrachyna revoluta (Flörke) Hale & Hale Krog & Swinscow
Parmelia rudecta Ach. Punctelia rudecta (Ach.) Krog Parmelinopsis spumosa (Asahina) Hypotrachyna spumosa (Asahina)
Parmelia sancti-angeli Lynge Parmotrema sancti-angelii (Lynge) Elix & Hale Krog & Swinscow
Hale Parmelinopsis swinscowii (Hale) Elix Hypotrachyna swinscowii (Hale)
Parmelia sinuosa (Sm.) Ach. Hypotrachyna sinuosa (Sm.) Hale & Hale Krog & Swinscow
Parmelia sorocheila Vain. Hypotrachyna sorocheila (Vain.) Parmentaria astroidea Fée Pyrenula astroidea (Fée) R.C. Harris
Divakar, A. Crespo, Sipman, Elix & Parmentaria chilensis Fée Pyrenula chilensis (Fée) R.C. Harris
Lumbsch
Parmotrema blanchetii (Hue) Hale Parmotrema blanchetianum (Müll.
Parmelia subcrinita auct. non Nyl. Parmotrema ultralucens (Krog) Hale
Arg.) Kalb
Parmelia sublaevigata (Nyl.) Nyl. Hypotrachyna sublaevigata (Nyl. ex
Parmotrema chinense (Osbeck) Parmotrema perlatum (Huds.) M.
Tuck.) Hale
Hale & Ahti Choisy
Parmelia sublaevigata var. texana Canoparmelia texana (Tuck.) Elix
Parmotrema leucosemothetum Parmotrema reticulatum (Taylor) M.
(Tuck.) Nyl. & Hale
(Hue) Hale Choisy
Parmelia subsinuosa Nyl. Hypotrachyna pulvinata (Fée) Hale
Parmotrema subarnoldii (Abbayes) Parmotrema arnoldii (Du Rietz) Hale
Parmelia taeniata Nyl. Anzia parasitica (Fée) Zahlbr. Hale
Peltigera spuria (Ach.) DC. Peltigera didactyla (With.) J.R. Phaeographina caesiopruinosa Platygramme caesiopruinosa (Fée)
Laundon (Fée) Müll. Arg. Fée
Peltigera tomentosa Vain. Peltigera laciniata (G. Merr.) Gyeln. Phaeographina chrysenteron Pallidogramme chrysenteron (Mont.)
(Mont.) Müll. Arg. Staiger, Kalb & Lücking
Penicillium funiculosum Thom Talaromyces funiculosus (Thom)
Samson, N. Yilmaz, Frisvad & Phaeographina chrysocarpa (Raddi) Allographa chrysocarpa (Raddi)
Seifert Redinger Lücking & Kalb
Penicillium purpurogenum Talaromyces purpureogenus Phaeographina colubrosa (Nyl.) Platygramme colubrosa (Nyl.)
Samson, N. Yilmaz, Houbraken, Müll. Arg. Staiger
Spierenb., Seifert, Peterson, Varga Phaeographina dividens (Nyl.) K.P. Phaeographis dividens (Nyl.) Kr.P.
& Frisvad Singh & D.D. Awasthi Singh & Swarnal.
Peniophora gemmea D.P. Rogers Hyphoderma gemmeum (D.P. Phaeographina dolichographa (Nyl.) Allographa dolichographa (Nyl.)
Rogers) Donk Zahlbr. Lücking & Kalb
Pertusaria acroscyphoides Sipman Lepra acroscyphoides (Sipman) I. Phaeographina internigricans Graphis internigricans Nyl.
Schmitt, B.P. Hodk. & Lumbsch (Leight.) Zahlbr.
Pertusaria albescens (Huds.) M. Lepra albescens (Huds.) Hafellner Phaeographina mesographa (Nyl.) Phaeographis mesographa (Nyl.)
Choisy & Werner Müll. Arg. Müll. Arg.
Pertusaria alpina Hepp ex Ahles Pertusaria leioplaca (Ach.) DC. Phaeographina scalpturata (Ach.) Phaeographis scalpturata (Ach.)
Pertusaria amara (Ach.) Nyl. Lepra amara (Ach.) Hafellner Müll. Arg. Staiger
Pertusaria culbersonii Vězda Varicellaria culbersonii (Vězda) I. Phaeographis cinnabarina (Fée) Thalloloma cinnabarinum (Fée)
Schmitt & Lumbsch Müll. Arg. Staiger
Pertusaria leioplaca var. octospora Pertusaria leioplaca (Ach.) DC. Phaeographis dimorpha (Nyl.) Platythecium leiogramma (Nyl.)
Nyl. Zahlbr. Staiger
Pertusaria leioplaca var. pycnocarpa Pertusaria leioplaca (Ach.) DC. Phaeographis glaucoleucoides (Nyl.) Diorygma junghuhnii (Mont. &
Nyl. Zahlbr. Bosch) Kalb, Staiger & Elix
Pertusaria leioplaca var. Pertusaria trypetheliiformis Nyl. Phaeographis inusta var. Phaeographis medusiformis
trypetheliiformis (Nyl.) Nyl. medusuliformis (Nyl.) Müll. Arg. (Kremp.) Müll. Arg.
Pertusaria leioplaca var. turgida Pertusaria leioplaca (Ach.) DC. Phaeographis leiogramma (Nyl.) Platythecium leiogramma (Nyl.)
Müll. Arg. Zahlbr. Staiger
Pertusaria leioplacoides var. rhodiza Pertusaria tetrathalamia (Fée) Nyl. Phaeographis patellula (Nyl.) Müll. Leiorreuma patellulum (Fée) Staiger
(Nyl.) Müll. Arg. Arg.
Pertusaria leioplacoides var. Pertusaria tetrathalamia (Fée) Nyl. Phaeographis serpentinella (Nyl.) Platythecium serpentinellum (Nyl.)
tetraspora Müll. Arg. Zahlbr. Staiger
Phellinus calcitratus (Berk. & M.A. Inonotus calcitratus (Berk. & M.A. Phyllachora antioquensis Chardón Phyllachora oxyspora Starbäck
Curtis) Ryvarden Curtis) Gomes-Silva & Gibertoni Phyllachora cornispora-necrotica Phyllachora acutispora Speg.
Phellinus callimorhus Lév. Fuscoporia callimorpha (Lév.) Chardón
Groposo, Log.-Leite & Góes-Neto Phyllachora molinae Chardón Phyllachora urvilleana Speg.
Phellinus callimorphus (Lév.) Fuscoporia callimorpha (Lév.) Phyllachora murilloi Garcés Phyllachora acutispora Speg.
Ryvarden Groposo, Log.-Leite & Góes-Neto
Phyllachora ruelliae Chardón Telimena ruelliae (Chardón)
Phellinus conchatus (Pers.) Quél. Phellinopsis conchata (Pers.) Y.C. Mardones, Trampe & M. Piepenbr.
Dai
Phyllobathelium epiphyllum (Müll. Phyllobathelium firmum (Stirt.)
Phellinus contiguus (Pers.) Pat. Fuscoporia contigua (Pers.) G. Arg.) Müll. Arg. Vězda
Cunn.
Phyllophiale alba R. Sant. Porina alba (R. Sant.) Lücking
Phellinus ferreus (Pers.) Bourdot & Fuscoporia ferrea (Pers.) G. Cunn.
Galzin Phyllophthalmaria coccinea (Leight.) Chroodiscus coccineus (Leight.)
Zahlbr. Müll. Arg.
Phellinus ferruginosus (Schrad.) Fuscoporia ferruginosa (Schrad.)
Pat. Murrill Phylloporina platypoda (Müll. Arg.) Phylloporis platypoda (Müll. Arg.)
Müll. Arg. Vězda
Phellinus linteus (Berk. & M.A. Tropicoporus linteus (Berk. & M.A.
Curtis) Teng Curtis) L.W. Zhou & Y.C. Dai Phylloporina rubentior (Stirt.) Müll. Porina rubentior (Stirt.) Müll. Arg.
Arg.
Phellinus pachyphloeus (Pat.) Pat. Inonotus pachyphloeus (Pat.) T.
Wagner & M. Fisch. Phylloporina rufula var. obscurata Porina limbulata (Kremp.) Vain.
(Müll. Arg.) Müll. Arg.
Phellinus punctatus (Fr.) Pilát Fomitiporia punctata (P. Karst.)
Murrill Phylloporina spruceana Müll. Arg. Porina limbulata (Kremp.) Vain.
Phellinus punctatus (P. Karst.) Pilát Fomitiporia punctata (P. Karst.) Phyllopsora buettneri var. glauca (B. Phyllopsora buettneri (Müll. Arg.)
Murrill de Lesd.) Brako Zahlbr.
Phellinus rufitinctus (Berk. & M.A. Phellinidium rufitinctum (Berk. Phyllopsora corallina var. Phyllopsora ochroxantha (Nyl.)
Curtis ex Cooke) Pat. & M.A. Curtis ex A.L. Sm.) ochroxantha (Nyl.) Brako Zahlbr.
Bondartseva & S. Herrera Phyllopsora corallina var. santensis Phyllopsora santensis (Tuck.)
Phellinus tropicalis M.J. Larsen & Tropicoporus tropicalis (M.J. Larsen (Eschw.) Brako Swinscow & Krog
Lombard & Lombard) L.W. Zhou & Y.C. Dai Phyllopsora labriformis Timdal Parallopsora labriformis (Timdal)
Phlebia livida (Pers.) Bres. Mycoacia livida (Pers.) Zmitr. Kistenich, Timdal & Bendiksby
Phlebia lividina Hjortstam Phlebia fascicularia (Rick) Phyllopsora leucophyllina (Nyl.) Parallopsora leucophyllina (Nyl.)
Nakasone & Burds. Timdal Kistenich, Timdal & Bendiksby
Phlebia subcretacea (Litsch.) M.P. Cabalodontia subcretacea (Litsch.) Phyllopsora parvifolia var. Phyllopsora canoumbrina (Vain.)
Christ. Piątek subgranulosa (Tuck.) Müll. Arg. Brako
Phlebiella tulasnelloidea (Höhn. & Xenasma tulasnelloideum (Höhn. & Phyllosticta noackiana Allesch. Boeremia noackiana (Allesch.)
Litsch.) Oberw. Litsch.) Donk Aveskamp, Gruyter & Verkley
Phlebiella vaga (Fr.) P. Karst. Xenasmatella vaga (Fr.) Stalpers Physcia acromela (Pers.) Nyl. Teloschistes flavicans (Sw.) Norman
Phlebiopsis ravenelii (Cooke) Phaeophlebiopsis ravenelii (Cooke) Physcia adglutinata f. minor (Fée) Hyperphyscia minor (Fée) Kalb
Hjortstam Zmitr. Nyl.
Phlebopus brunneoruber (Beeli) Phlebopus braunii (Bres.) Heinem. Physcia aegialita (Ach.) Nyl. Dirinaria aegialita (Afzel. ex Ach.)
Heinem. & Rammeloo B.J. Moore
Phloeospora mimosae-pigrae Evans Sphaerulina mimosae-pigrae H.C. Physcia albicans (Pers.) J.W. Heterodermia albicans (Pers.)
& G. Carrión Evans & G. Carrión Thomson Swinscow & Krog
Pyrenula costaricensis Müll. Arg. Pyrenula aggregata (Fée) Fée Ramalina yemensis var. ecklonii f. Ramalina celastri (Spreng.) Krog &
sublinearis Nyl. Swinscow
Pyrenula discolor Ach. Ampliotrema discolor (Ach.) Kalb
Ramaria botrytis (Pers.) Bourdot Ramaria botrytis (Pers.) Bourdot
Pyrenula laii Aptroot Pyrenula immissa (Stirt.) Zahlbr. var. botrytis
Pyrenula macrocarpa A. Massal. Pyrenula complanata (Mont.) Trevis. Ramaria botrytis (Pers.) Ricken var. Ramaria botrytis (Pers.) Bourdot
Pyrenula marginata Hook. Pyrenula mamillana (Ach.) Trevis. botrytis
Pyrenula marginatula Müll. Arg. Pyrenula adacta Fée Ramaria botrytis var. Ramaria aurantiiramosa (Marr &
aurantiiramosa Marr & D.E. Stuntz D.E. Stuntz) Franchi & M. Marchetti
Pyrenula mastophoroides var. Pyrenula mastophoroides (Nyl.)
flavicans (Nyl.) Zahlbr. Zahlbr. Ramaria zippelii (Lév.) Corner Phaeoclavulina zippelii (Lév.)
Overeem
Pyrenula mucosa (Vain.) R.C. Harris Pyrenula papillifera (Nyl.) Aptroot
Ramularia endophylla Verkley & U. Mycosphaerella punctiformis (Pers.)
Pyrenula nova-granadensis Upreti & Pyrenula ravenelii (Tuck.) R.C.
Braun Starbäck
Ajay Singh Harris
Resinicium friabile Hjortstam & Melo Resinicium luteosulphureum (Rick)
Pyrenula obvoluta (Nyl.) Aptroot Pyrenula dermatodes (Borrer)
Baltazar & Rajchenb.
Schaer.
Pyrenula obvoluta (Nyl.) R.C. Harris Pyrenula dermatodes (Borrer) Rexia fuliginosa (Filson) S. Stenroos, Rexiella fuliginosa (Filson) S.
& Aptroot Schaer. Pino-Bodas & Ahti Stenroos, Pino‐Bodas and Ahti
Pyrenula pinguis Fée Pyrenula quassiicola Fée Rhodocybe nitellina (Fr.) Singer Rhodophana nitellina (Fr.) Papetti
Pyrenula santensis (Nyl.) Müll. Arg. Pyrenula balia (Kremp.) R.C. Harris Rhodocybe testacea Dennis Clitopilus testaceus (Dennis)
Noordel. & Co-David
Pyrenula subaggregata Müll. Arg. Pyrenula aspistea (Afzel. ex Ach.)
Ach. Rhodophyllus ferrugineogranulatus Entoloma ferrugineogranulatum
Singer (Singer) E. Horak
Pyrenula subducta var. retracta Pyrenula subducta (Nyl.) Müll. Arg.
(Nyl.) Zahlbr. Rhodophyllus lyophylliformis Singer Entoloma lyophylliforme (Singer) E.
Horak
Pyrenula verruculosa Upreti & Ajay Pyrenula schiffneri (Zahlbr.) Aptroot
Singh Rhodotorula colostri (T. Castelli) Rhodosporidiobolus colostri (T.
Lodder Castelli) Q.M. Wang, F.Y. Bai, M.
Pyrrhospora russula (Ach.) Hafellner Ramboldia russula (Ach.) Kalb, Groenew. & Boekhout
Lumbsch & Elix
Rhodotorula rubra (Schimon) F.C. Rhodotorula mucilaginosa (A. Jörg.)
Pyxine caesiopruinosa (Tuck.) Pyxine albovirens (G. Mey.) Aptroot Harrison F.C. Harrison
Imshaug
Rhodotorula taiwanensis F.L. Lee & Rhodotorula taiwanensis F.L. Lee
Pyxine cocoes f. isidiophora Müll. Pyxine isidiophora (Müll. Arg.) C.H. Huang & C.H. Huang ex Denchev & T.
Arg. Imshaug Denchev
Pyxine cocoes var. sorediata (Ach.) Pyxine sorediata (Ach.) Mont. Ricasolia corrosa (Ach.) Nyl. Yoshimuriella corrosa (Ach.)
Tuck. Moncada & Lücking
Pyxine isidiophora (Müll. Arg.) Pyxine coralligera Malme Ricasolia crenulata (Hook.) Nyl. Lobariella crenulata (Hook.)
Imshaug Yoshim.
Pyxine meissneri Nyl. Pyxine berteriana (Fée) Imshaug Ricasolia excisa (Müll. Arg.) Stizenb. Emmanuelia excisa (Müll. Arg.)
Pyxine microspora Vain. Pyxine pyxinoides (Müll. Arg.) Kalb Lücking, Moncada & Ant. Simon
Pyxine sorediata f. caesiopruinosa Pyxine albovirens (G. Mey.) Aptroot Ricasolia fendleri (Mont. & Tuck.) Yoshimuriella fendleri (Tuck. &
(Tuck.) Hue Nyl. Mont.) Moncada & Lücking
Racodium rupestre Pers. Cystocoleus ebeneus (Dillwyn) Ricasolia pallida (Hook.) Nyl. Lobariella pallida (Hook.) Moncada
Thwaites & Lücking
Rimelia commensurata (Hale) Hale Parmotrema commensuratum Scytinostroma galactinum (Fr.) Donk Baltazaria galactina (Fr.) Leal-Dutra,
& A. Fletcher (Hale) Hale Dentinger & G.W. Griff.
Rimelia reticulata (Taylor) Hale & A. Parmotrema reticulatum (Taylor) M. Scytinostroma ochroleucum (Bres. Scytinostroma lusitanicum (Trotter)
Fletcher Choisy & Torrend) Donk P.M. Kirk
Rimelia simulans (Hale) Hale & A. Parmotrema simulans (Hale) Hale Segestria octomera (Müll. Arg.) R.C. Porina octomera (Müll. Arg.) F.
Fletcher Harris Schill.
Rimelia subisidiosa (Müll. Arg.) Hale Parmotrema subisidiosum (Müll. Segestria rubentior (Stirt.) R.C. Porina rubentior (Stirt.) Müll. Arg.
& A. Fletcher Arg.) Hale Harris
Rinodina insperata (Nyl.) Malme Orcularia insperata (Nyl.) Kalb & Sesquicillium pseudosetosum Clonostachys pseudosetosa
Giralt Samuels (Samuels) Schroers
Roccella decipiens Darb. Roccella verruculosa Follmann Sphaeria depressa Hook. Phacidium depressum Hook. ex
Berk.
Rogersella asperula Liberta & A.J. Lyomyces griseliniae (G. Cunn.)
Navas Riebesehl & Langer Sphaeria depressa J.M. Hook Phacidium depressum Hook. ex
Berk.
Rogersella griseliniae (G. Cunn.) Lyomyces griseliniae (G. Cunn.)
Stalpers Riebesehl & Langer Sphaeria melanococca Lév. Dothidotthia melanococca (Lév.)
Rosellinia bunodes (Berk. & Dematophora bunodes (Berk. & Aptroot
Broome) Sacc. Broome) C. Lambert, Wittstein & Sphaerodothis columbiensis Phyllachora columbiensis (Chardón)
M. Stadler Chardón P.F. Cannon
Rosellinia obregonii L.E. Petrini Dematophora obregonii (L.E. Sphaerophoron compressum Ach. Bunodophoron melanocarpum (Sw.)
Petrini) C. Lambert, Wittstein & M. Wedin
Stadler
Sphaerophorus compressus Ach. Bunodophoron melanocarpum (Sw.)
Rozites colombiana Halling & Cortinarius colombianus (Halling & Wedin
Ovrebo Ovrebo) Peintner, E. Horak, M.M.
Moser & Vilgalys Sphaerophorus formosanus Bunodophoron melanocarpum (Sw.)
(Zahlbr.) Asahina Wedin
Rozites colombianus Halling & Cortinarius colombianus (Halling &
Ovrebo Ovrebo) Peintner, E. Horak, M.M. Sphaerophorus melanocarpus (Sw.) Bunodophoron melanocarpum (Sw.)
Moser & Vilgalys DC. Wedin
Russula emetica subsp. lacustris Russula emetica (Schaeff.) Pers. Sphaerostilbe gracilipes Tul. & C. Nectria gracilipes (Tul. & C. Tul.)
Singer Tul. Wollenw.
Saccomorpha icmalea (Ach.) Placynthiella icmalea (Ach.) Coppins Spirechina columbiensis F. Kern & Gerwasia columbiensis (F. Kern &
Clauzade & Cl. Roux & P. James Whetzel Whetzel) Buriticá
Sarcographa actinobola (Nyl.) Müll. Sarcographa cinchonarum Fée Squamacidia janeirensis (Müll. Arg.) Phyllopsora cinchonarum (Fée)
Arg. Brako Timdal
Sarcographa decolorascens (Nyl.) Phaeographis decolorascens (Nyl.) Steccherinum subochraceum Steccherinum hydneum Rick ex
Zahlbr. Lücking Bononi & Hjortstam Maas Geest.
Sarcographa intricans (Nyl.) Müll. Phaeographis intricans (Nyl.) Stegobolus anamorphoides (Nyl.) Rhabdodiscus anamorphoides (Nyl.)
Arg. Staiger Lücking Vain.
Sarcographa leprieurii (Mont.) Müll. Phaeographis leprieurii (Mont.) Stegobolus anamorphus (Nyl.) Rhabdodiscus anamorphoides (Nyl.)
Arg. Staiger Frisch & Kalb Vain.
Schaereria tenebrosa (Flot.) Hertel Schaereria fuscocinerea (Nyl.) Stegobolus auberianus (Mont.) Rhabdodiscus auberianus (Mont.)
& Poelt Clauzade & Cl. Roux A.Frisch & Kalb Vain.
Schismatomma flavescens (Nyl.) Syncesia flavescens (Nyl.) Tehler Stegobolus auberianus (Mont.) Rhabdodiscus auberianus (Mont.)
Zahlbr. Frisch & Kalb Vain.
Stereocaulon denudatum var. Stereocaulon vesuvianum Pers. Sticta peruviana (Delise) Nyl. Sticta weigelii (Ach.) Vain.
vesuvianum (Pers.) Laur. Sticta tomentosa f. latior Nyl. Sticta cometia Ach.
Stereocaulon lecanoreum Nyl. Stereocaulon strictum Th. Fr. Sticta tomentosa f. leucoblepharis Sticta leucoblepharis Tuck. & Mont.
Stereocaulon mixtum Nyl. Stereocaulon ramulosum Raeusch. (Mont.) Nyl.
Stereocaulon proximum Nyl. Stereocaulon ramulosum Raeusch. Sticta tomentosa f. leucoblepharis Sticta leucoblepharis Tuck. & Mont.
(Mont.) Zahlbr.
Stereocaulon proximum var. Stereocaulon strictum Th. Fr.
compressum (Nyl.) Nyl. Sticta tomentosa f. leucoblepharis Sticta leucoblepharis Tuck. & Mont.
(Tuck. & Mont.) Zahlbr.
Stereocaulon proximum var. Stereocaulon ramulosum Raeusch.
gracilius Müll. Arg. Sticta tomentosa f. ornata (Müll. Sticta dilatata (Nyl.) Vain.
Arg.) Hue
Stereocaulon ramulosum f. Stereocaulon strictum Th. Fr.
compressum Nyl. Sticta tomentosa var. dilatata (Nyl.) Sticta dilatata (Nyl.) Vain.
Hue
Stereocaulon ramulosum f. Stereocaulon meyeri Stein
farinosum Th. Fr. Sticta weigelii f. beauvoisii (Delise) Sticta beauvoisii Delise
Hue
Stereocaulon ramulosum var. Stereocaulon strictum Th. Fr.
compressum (Nyl.) Nyl. Sticta weigelii var. beauvoisii Sticta beauvoisii Delise
(Delise) Hue
Stereocaulon ramulosum var. Stereocaulon ramulosum Raeusch.
elegans Th. Fr. Sticta weigelii var. peruviana Sticta weigelii (Ach.) Vain.
(Delise) Vain.
Stereocaulon tomentosum Fr. var. Stereocaulon tomentosum Th. Fr.
tomentosum Stictina andensis Nyl. Sticta andensis (Nyl.) Trevis.
Stereocaulon vesuvianum var. Stereocaulon vesuvianum Pers. Stictina andreana Müll. Arg. Sticta andreana (Müll. Arg.) Zahlbr.
denudatum (Flörke) I.M. Lamb. Stictina cometia (Ach.) Nyl. Sticta cometia Ach.
Stereocaulon violascens Müll. Arg. Stereocaulon vesuvianum Pers. Stictina filicinella Nyl. Sticta filicinella (Nyl.) Zahlbr.
Sticta aurata Ach. Crocodia aurata (Ach.) Link Stictina fuliginosa (Dicks.) Nyl. Sticta fuliginosa (Dicks.) Ach.
Sticta colombiana Moncada & Sticta andina Moncada, Lücking & Stictina gyalocarpa Nyl. Sticta gyalocarpa (Nyl.) Trevis.
Lücking Sérus.
Stictina humboldtii (Hook.) Nyl. Sticta humboldtii Hook.
Sticta damaecornis f. rudiuscula Sticta rudiuscula (Vain.) Moncada
Stictina impressula (Nyl.) Nyl. Sticta impressula (Nyl.) Zahlbr.
Vain. & Lücking
Stictina kunthii (Hook.) Nyl. Sticta kunthii Hook.
Sticta damicornis f. subscrobiculata Sticta subscrobiculata (Nyl.) Gyeln.
(Nyl.) Stizenb. Stictina kunthii var. pilosella Nyl. Sticta kunthii Hook.
Stictina limbata f. andensis (Nyl.) Sticta andensis (Nyl.) Trevis. Strigula viridiseda (Nyl.) R.C. Harris Dichoporis viridiseda (Nyl.) S.H.
Jiang, Lücking & Sérus.
Stictina peltigerella Nyl. Sticta peltigerella (Nyl.) Trevis.
Strigula vulgaris (Müll. Arg.) Lücking Phylloporis vulgaris (Müll. Arg.) S.H.
Stictina peruviana (Delise) Nyl. Sticta weigelii (Ach.) Vain. Jiang, Lücking & J.C. Wei
Stictina quercizans var. beauvoisii Sticta beauvoisii Delise Stropharia semiglobata (Batsch) Protostropharia semiglobata
(Delise) Müll. Arg. Quél. (Batsch) Redhead, Moncalvo &
Stictina quercizans var. ornata Müll. Sticta tomentosa (Sw.) Ach. Vilgalys
Arg. Stypella minor Möller Protomerulius minor (A. Møller) V.
Stictina quercizans var. peruviana Sticta weigelii (Ach.) Vain. Spirin & O. Miettinen
(Delise) Nyl. Teloschistes chrysophthalmus (L.) Niorma chrysophthalma (L.) S.Y.
Stictina tomentosa (Sw.) Nyl. Sticta tomentosa (Sw.) Ach. Th. Fr. Kondr., Kärnefelt, Elix, A. Thell,
M.H. Jeong & Hur
Stictina tomentosa var. dilatata Nyl. Sticta tomentosa (Sw.) Ach.
Teloschistes hypoglaucus (Nyl.) Niorma hypoglauca (Nyl.) S.Y.
Stictina tomentosa var. impressula Sticta impressula (Nyl.) Zahlbr.
Zahlbr. Kondr., Kärnefelt, Elix, A. Thell,
Nyl.
M.H. Jeong & Hur
Stictina tomentosa var. ornata Sticta tomentosa (Sw.) Ach.
Tephromela aglaea (Sommerf.) Calvitimela aglaea (Sommerf.)
(Müll. Arg.) Müll. Arg.
Hertel & Rambold Hafellner
Stigmatidium extenuatum (Nyl.) Nyl. Sclerophyton extenuatum (Nyl.)
Sparrius Thanatephorus cucumeris (A.B. Rhizoctonia solani J.G. Kühn
Frank) Donk
Stigmatidium granulatum auct. non Erythrodecton granulatum (Mont.)
(Mont.) Nyl. G. Thor Thecaria montagnei (Bosch) Staiger Pliariona montagnei (Bosch) A.
Massal.
Stigmatidium interruptum (Fée) Nyl. Platygraphopsis interrupta (Fée)
Müll. Arg. Thelenella cinerascens (Vain.) R.C. Aspidothelium cinerascens Vain.
Harris
Stigmatidium leptostictum Nyl. Mazosia leptosticta (Nyl.) Sparrius
Thelenella fugiens (Müll. Arg.) R.C. Aspidothelium fugiens (Müll. Arg.)
Stirtonia sprucei R. Sant. Amazonomyces sprucei (R. Sant.) Harris R. Sant.
Lücking, Sérus. & G. Thor
Theloschistes flavicans var. Teloschistes flavicans (Sw.) Norman
Strigula actinoplaca Nyl. Actinoplaca strigulacea Müll. Arg. acromela (Pers.) Müll. Arg.
Strigula agyronema var. confluens Puiggariella nemathora (Mont.) S.H. Theloschistes flavicans var. exilis Teloschistes exilis (Michx.) Vain.
Müll. Arg. Jiang, Lücking & J.C. Wei (Michx.) Müll. Arg.
Strigula complanata (Fée) Mont. Phyllocharis orbicularis (Fr.) S.H.
Thelotrema albidum Nyl. Chapsa albida (Nyl.) Lücking &
Jiang, Lücking & Sérus.
Sipman
Strigula elegans (Fée) Müll. Arg. Strigula smaragdula Fr.
Thelotrema albomaculatum Sipman Pseudochapsa albomaculata
Strigula janeirensis (Müll. Arg.) Raciborskiella janeirensis (Müll. (Sipman) Parnmen, Lücking &
Lücking Arg.) R. Sant. Lumbsch
Strigula maculata (Cooke & Racoplaca maculata (Cooke & Thelotrema auberianoides Nyl. Ocellularia auberianoides (Nyl.)
Massee) R. Sant. Massee) S.H. Jiang, Lücking & J.C. Müll. Arg.
Wei
Thelotrema bahianum (Ach.) Ach. Ocellularia bahiana (Ach.) Frisch
Strigula melanobapha (Kremp.) R. Racoplaca melanobapha (Kremp.)
Sant. S.H. Jiang, Lücking & J.C. Wei Thelotrema bahianum var. Ocellularia obturascens (Nyl.) Hale
obturascens Nyl.
Strigula nemathora Mont. Puiggariella nemathora (Mont.) S.H.
Jiang, Lücking & J.C. Wei Thelotrema bahianum var. ruptum Ocellularia bahiana (Ach.) Frisch
Nyl.
Strigula nigrocincta var. soluta Müll. Puiggariella nemathora (Mont.) S.H.
Arg. Jiang, Lücking & J.C. Wei Thelotrema berkeleyanum (Mont.) Stegobolus berkeleyanus Mont.
Brusse
Strigula obducta (Müll. Arg.) R.C. Phylloporis obducta (Müll. Arg.) R.
Harris Sant. & Tibell Thelotrema brasilianum Hale Acanthotrema brasilianum (Hale)
Frisch
Strigula orbicularis Fr. Phyllocharis orbicularis (Fr.) S.H.
Jiang, Lücking & Sérus. Thelotrema calvescens Fée Ocellularia calvescens (Fée) Müll.
Arg.
Strigula phaea (Ach.) R.C. Harris Dichoporis phaea (Ach.) S.H. Jiang,
Lücking & Sérus. Thelotrema cavatum Ach. Ocellularia cavata (Ach.) Müll. Arg.
Strigula phyllogena (Müll. Arg.) R.C. Phylloporis phyllogena (Müll. Arg.) Thelotrema cavatum f. amplius Nyl. Ampliotrema amplius (Nyl.) Kalb
Harris Clem. Thelotrema cavatum f. obturatum Ocellularia cavata (Ach.) Müll. Arg.
Strigula plana Müll. Arg. Strigula smaragdula Fr. (Ach.) Nyl.
Strigula platypoda (Müll. Arg.) R.C. Phylloporis platypoda (Müll. Arg.) Thelotrema cavatum var. amplius Ampliotrema amplius (Nyl.) Kalb
Harris Vězda Nyl.
Thelotrema laevigans Nyl. Clandestinotrema clandestinum Tricharia albostrigosa R. Sant. Aderkomyces albostrigosus (R.
(Ach.) Rivas Plata, Lücking & Sant.) Lücking, Sérus. & Vězda
Lumbsch Tricharia couepiae (Bat.) Lücking Aderkomyces couepiae Bat.
Thelotrema leucocarpoides Nyl. Ocellularia leucocarpoides (Nyl.) Tricharia heterella (Stirt.) Lücking Aderkomyces heterellus (Stirt.)
Lücking Lücking, Sérus. & Vězda
Thelotrema leucomelanum Nyl. Clandestinotrema leucomelanum Tricharia subalbostrigosa Lücking Aderkomyces subalbostrigosus
(Nyl.) Rivas Plata, Lücking & (Lücking) Lücking, Sérus. & Vězda
Lumbsch
Tricharia vulgaris (Müll. Arg.) R. Gyalideopsis vulgaris (Müll. Arg.)
Thelotrema leucomelanum var. Clandestinotrema cathomalizans Sant. Lücking
cathomalizans Nyl. (Nyl.) Rivas Plata, Lücking &
Tricholoma cystidiosum Cifuentes Tricholoma cifuentesii Courtec.
Lumbsch
& Guzmán
Thelotrema metaphoricum Nyl. Rhabdodiscus metaphoricus (Nyl.)
Trichosporon gamsii Middelhoven, Apiotrichum gamsii (Middelhoven,
Vain.
Scorzetti, Sigler & Fell Scorzetti, Sigler & Fell) Yurkov &
Thelotrema microporoides Nyl. Myriotrema rugiferum (Harm.) Hale Boekhout
Thelotrema monosporum var. Thelotrema lepadodes Tuck. Trichothelium rubescens Lücking Porina rubescens (Lücking)
patulum Nyl. Hafellner & Kalb
Thelotrema myriocarpum Fée Austrotrema myriocarpum (Fée) I. Triclinum cinchonarum Fée Phyllopsora cinchonarum (Fée)
Medeiros, Lücking & Lumbsch Timdal
Trypethelium variolosum Ach. Astrothelium variolosum (Ach.) Müll. Uromyces trifolii (R. Hedw.) Lév. Puccinia trifolii R. Hedw.
Arg. Uromyces variabilis Mayor Gerwasia variabilis (Mayor) Buriticá
Tubeufia eriodermatis Etayo Lichenotubeufia eriodermatis Usnea antillarum (Vain.) Zahlbr. Eumitria baileyi Stirt.
(Etayo) Etayo
Usnea baileyi (Stirt.) Zahlbr. Eumitria baileyi Stirt.
Tubeufia pannariae Etayo Lichenotubeufia pannariae (Etayo)
Etayo Usnea barbata f. ceratina (Ach.) Usnea ceratina Ach.
Schaer
Tuckneraria laureri (Kremp.) Nephromopsis laureri (Kremp.)
Randlane & A. Thell Kurok. Usnea barbata var. rubiginea Meyen Usnea rubicunda Stirt.
& Flot.
Tylophorella polyspora f. compressa Tylophorella pyrenocarpoides (Müll.
Nádv. Arg.) Egea Usnea barbata var. scabrida (Taylor) Usnea rubicunda Stirt.
Müll. Arg.
Tylophorella polyspora Vain. Tylophorella pyrenocarpoides (Müll.
Arg.) Egea Usnea barbata var. strigosa (Ach.) Usnea strigosa (Ach.) Pers.
Flot.
Tylophoron rufocapillatum Nádv. Tylophoron protrudens Nyl.
Usnea caespitia Motyka Usnea durietzii Motyka
Tylophoron simile Nádv. Tylophoron protrudens Nyl.
Usnea ceratina var. scabrosa Ach. Usnea ceratina Ach.
Tylopilus indecisus (Peck) Murrill Porphyrellus indecisus (Peck) E.-J.
Gilbert Usnea chaetophora Stirt. Usnea dasopoga (Ach.) Nyl.
Tylopilus umbrosus (G.F. Atk.) A.H. Porphyrellus umbrosus (G.F. Atk.) Usnea finkii Zahlbr. Usnea transitoria Motyka
Sm. & Thiers Singer, J. García & L.D. Gómez Usnea florida var. comosa Ach. Usnea subfloridana Stirt.
Tyromyces duracinus (Pat.) Murrill Trullella duracina (Pat.) Zmitr.
Usnea florida var. mollis (Stirt.) Usnea mollis Stirt.
Tyromyces subcaesius A. David Cyanosporus subcaesius (A. David) Vain.
B.K. Cui, L.L. Shen & Y.C. Dai
Usnea florida var. perplexans (Stirt.) Usnea perplexans Stirt.
Uleodothis andina Chardón Achorella andina (Chardón) Chardón Vain.
Usnea rubricornuta Truong & P. Usnea moreliana Motyka Verrucaria mastophoroides Nyl. Pyrenula mastophoroides (Nyl.)
Clerc Zahlbr.
Usnea solida Motyka Usnea ceratina Ach. Verrucaria mastophoroides var. Pyrenula mastophoroides (Nyl.)
flavicans Nyl. Zahlbr.
Usnea spinulifera (Vain.) Motyka Usnea dasaea Stirt.
Verrucaria myriomma Nyl. Astrothelium annulare (Fée) Aptroot
Usnea spinulifera (Vain.) Motyka ex Usnea dasaea Stirt. & Lücking
B. de Lesd.
Verrucaria nitida var. aspistea (Ach.) Pyrenula aspistea (Afzel. ex Ach.)
Usnea subcomosa Vain. Usnea ceratina Ach. Nyl. Ach.
Usnea subhirta (Vain.) Motyka Usnea cornuta Körb. Verrucaria nucula (Ach.) Nyl. Porina nucula Ach.
Usnea sulcata Motyka Usnea alata Motyka Verrucaria nucula var. endochrysea Porina endochrysea C. Bab.
(Mont.) Nyl.
Usnea sulcata var. neutra (Motyka) Usnea alata Motyka
Motyka. Verrucaria obvoluta Nyl. Pyrenula dermatodes (Borrer)
Schaer.
Usnea superba Motyka Usnea arthroclada Fée
Verrucaria ochraceoflava Nyl. Pyrenula ochraceoflava (Nyl.) R.C.
Usnea trichodea f. gracilis (Ach.) Usnea gracilis Ach.
Harris
Hue
Verrucaria operta Nyl. Pyrenula globifera (Eschw.) Aptroot
Ustilago concelata Zundel Sporisorium concelatum (Zundel)
M. Piepenbr. Verrucaria papilligera Leight. Pyrenula papilligera (Leight.) Müll.
Arg.
Ustilago garcesii Zundel Anthracocystis panici-leucophaei
(Bref.) McTaggart & R.G. Shivas Verrucaria papulosa Nyl. Astrothelium papulosum (Nyl.)
Aptroot & Lücking
Verrucaria americana Fée Porina americana Fée
Verrucaria pleiomera Nyl. Pyrenula pleiomera (Nyl.) Zahlbr.
Verrucaria apposita Nyl. Bogoriella apposita (Nyl.) Aptroot &
Lücking Verrucaria porinoides Mont. Astrothelium pupula (Ach.) Aptroot
& Lücking
Verrucaria aspistea Ach. Pyrenula aspistea (Afzel. ex Ach.)
Ach. Verrucaria punctella var. exstans Pyrenula quassiicola Fée
Nyl.
Verrucaria astroidea (Fée) Nyl. Pyrenula astroidea (Fée) R.C. Harris
Verrucaria pupula (Ach.) Nyl. Astrothelium pupula (Ach.) Aptroot
Verrucaria belonospora Nyl. Porina belonospora (Nyl.) Müll. Arg. & Lücking
Verrucaria cartilaginea Fée Astrothelium pupula (Ach.) Aptroot Verrucaria pyrenuloides (Mont.) Nyl. Pyrenula pyrenuloides (Mont.) R.C.
& Lücking Harris
Verrucaria catapasta Nyl. Polymeridium catapastum (Nyl.) Verrucaria santensis Tuck. Pyrenula balia (Kremp.) R.C. Harris
R.C. Harris
Verrucaria subducta f. retracta Nyl. Pyrenula subducta (Nyl.) Müll. Arg.
Verrucaria contendens Nyl. Polymeridium contendens (Nyl.) R.C. Verrucaria subducta Nyl. Pyrenula subducta (Nyl.) Müll. Arg.
Harris
Verrucaria subducta var. retracta Pyrenula subducta (Nyl.) Müll. Arg.
Verrucaria convexa (Nyl.) Nyl. Pyrenula subducta (Nyl.) Müll. Arg. Nyl.
Verrucaria cryptostoma Nyl. Pyrenula cryptostoma (Nyl.) Müll. Verrucaria subprostans Nyl. Anisomeridium subprostans (Nyl.)
Arg. R.C. Harris
Verrucaria diffluens Nyl. Astrothelium pulcherrimum (Fée) Verrucaria thelena Ach. Bogoriella thelena (Ach.) Aptroot &
Aptroot & Lücking Lücking
Verrucaria diluta Fée Pseudopyrenula diluta (Fée) Müll. Verrucaria thelotremoides Nyl. Astrothelium thelotremoides (Nyl.)
Arg. Aptroot & Lücking
Verrucaria dirempta Nyl. Pseudopyrenula diluta (Fée) Müll. Verrucaria tropica Ach. Nigrovothelium tropicum (Ach.)
Arg. Lücking, M.P. Nelsen & Aptroot
Verrucaria dolichophora Nyl. Porina dolichophora (Nyl.) Müll. Arg. Wardia vastatrix J.F. Hennen & Hemileia vastatrix Berk. & Broome
Verrucaria duplicans Nyl. Pyrenula duplicans (Nyl.) Aptroot M.M. Hennen
Woessia pseudohyphophorifera Bacidina pseudohyphophorifera
Verrucaria heterochroa Mont. Astrothelium aeneum (Eschw.)
Lücking & Sérus. (Lücking & Sérus.) Lücking
Aptroot & Lücking
Xanthomendoza weberi (S.Y. Kondr. Gallowayella weberi (S.Y. Kondr. &
Verrucaria hypophyta Nyl. Pyrenula dermatodes (Borrer)
& Kärnefelt) L. Lindblom Kärnefelt) S.Y. Kondr., Fedorenko,
Schaer.
S. Stenroos, Kärnefelt, Elix, Hur &
Verrucaria intrusa Nyl. Polyblastiopsis intrusa (Nyl.) Zahlbr. A. Thell
Astrothelium hypoxylon auct. non Astrothelium sulphureum (Eschw.) Cladonia bacillaris Nyl. Cladonia didyma (Fée) Vain.
(Fée) Nyl. Nyl. Cladonia bellidiflora (Ach.) Schaer. Cladonia microscypha Ahti & S.
Bacidia consimilis auct. non (Müll. Loflammiopsis brasiliensis Lücking Stenroos
Arg.) R. Sant. & Kalb Cladonia boryi Tuck. not occurring in Colombia
Badimia galbinea (Kremp.) Vězda Badimia vezdana Lücking, Farkas Cladonia calycantha auct. non Nyl. Cladonia rappii var. exilior (Abbayes)
& V. Wirth Ahti
Baeomyces placophyllus auct. non Phyllobaeis erythrella (Mont.) Kalb Cladonia carassensis Vain. not occurring in Colombia
Ach.
Cladonia cervicornis (Ach.) Flot. Cladonia rappii var. exilior (Abbayes)
Bryocaulon divergens (Ach.) not occurring in Colombia Ahti
Kärnefelt
Cladonia clathrata Ahti & L. Xavier not occurring in Colombia
Bryoria bicolor (Ehrh.) Brodo & D. Oropogon bicolor Essl.
Cladonia coniocraea (Flörke) Spreng. not occurring in Colombia
Hawksw.
Buellia disciformis (Fr.) Mudd Cratiria obscurior (Stirt.) Marbach Cladonia cornucopioides auct. non Fr. Cladonia corallifera (Kunze) Nyl.
& Kalb Cladonia cornuta (L.) Hoffm. not occurring in Colombia
Buellia tetrapla (Nyl.) Müll. Arg. not occurring in Colombia Cladonia degenerans f. trachyna Cladonia squamosa (Scop.) Hoffm.
Bunodophoron insigne (Laurer) Bunodophoron melanocarpum (Sw.) auct. non (Ach.) Flörke
Wedin Wedin Cladonia degenerans var. Cladonia ramulosa (With.) J.R.
Byssoloma subpolychromum auct. Byssoloma minutissimum Kalb & gracilescens auct. non (Flörke) Laundon
non Vězda Vězda Spreng.
Caloplaca crocea (Kremp.) Hafellner Caloplaca erythroleuca (Nyl.) Cladonia degenerans var. trachyna Cladonia squamosa (Scop.) Hoffm.
& Poelt Zahlbr. auct. non Nyl.
Caloplaca lucifera Thor Caloplaca granularis (Müll. Arg.) Cladonia dilleniana Flörke not occurring in Colombia
Zahlbr. Cladonia dissecta Ahti not occurring in Colombia
Caloplaca pallidior (Nyl.) Zahlbr. not occurring in Colombia Cladonia fimbriata (L.) Fr. Cladonia subsquamosa Kremp.
Candelaria fibrosa (Fr.) Müll. Arg. Candelaria concolor (Dicks.) Arnold Cladonia flagellaris Ahti & Marcelli not occurring in Colombia
Celothelium dominicanum auct. non Celothelium cinchonarum (Müll. Cladonia furfuracea auct. non Vain. Cladonia furfuraceoides Ahti &
(Vain.) M.B. Aguirre Arg.) Vain. Sipman
Cetraria crispa auct. non (Ach.) Nyl. Cetraria arenaria Kärnefelt Cladonia hypoxantha auct. non Cladonia microscypha Ahti & S.
Cetraria islandica var. crispa auct. Cetraria arenaria Kärnefelt Tuck. Stenroos
non Ach. Cladonia macilenta var. carcata Cladonia miniata G. Mey.
Cetrelia olivetorum (Nyl.) W.L. Culb. not occurring in Colombia auct. non (Ach.) Nyl.
& C.F. Culb. Cladonia mitis Sandst. Cladonia boliviana Ahti
Cresponea premnea (Ach.) Egea & not occurring in Colombia Heterodermia andina Moberg not occurring in Colombia
Torrente Hyperphyscia adglutinata auct. non Hyperphyscia confusa Essl., C.A.
Cryptothecia atropunctata G. Thor not occurring in Colombia (Flörke) H. Mayrhofer & Poelt Morse & S.D. Leav.
Cryptothecia bartlettii G. Thor not occurring in Colombia Hypogymnia austerodes auct. non Hypogymnia bitteri (Lynge) Ahti
(Nyl.) Räsänen
Cryptothecia candida (Kremp.) R. Myriostigma filicinum (Ellis & Everh.)
Sant. Frisch & G. Thor Hypotrachyna thysanota (Kurok.) not occurring in Colombia
Hale
Cryptothecia candida auct. non Myriostigma filicinum (Ellis & Everh.)
(Kremp.) R. Sant. Frisch & G. Thor Laurera chrysoglypha auct. non Astrothelium sphaerioides (Mont.)
(Vain.) Zahlbr. Aptroot & Lücking
Cryptothecia cf. exilis G. Thor not occurring in Colombia
Lecanora allophana (Ach.) Nyl. not occurring in Colombia
Cyanisticta obvoluta (Sw.) C.W. not occurring in Colombia
Dodge Lecanora aurantiaca auct. non Caloplaca xanthopa (Hue) Zahlbr.
(Lightf.) Nyl.
Dibaeis sorediata Kalb & Gierl not occurring in Colombia
Lecanora cancriformis auct. non Lecanora caesiorubella Ach.
Dichosporidium boschianum (Mont.) not occurring in Colombia (Hoffm.) Vain.
G. Thor
Lecanora cenisia Ach. not occurring in Colombia
Diploschistes gypsaceus auct. non Diploschistes diacapsis (Ach.)
(Ach.) Zahlbr. Lumbsch Lecanora miculata Ach. Caloplaca brebissonii (Fée) J. Sant.
ex Hafellner & Poelt
Dirinaria coccinea (Müll. Arg.) D.D. not occurring in Colombia
Awasthi Lecanora pallidior Nyl. not occurring in Colombia
Dirinaria palmarum (Vain.) C.W. not occurring in Colombia Lecanora populicola (DC.) Duby not occurring in Colombia
Dodge Lecanora pulicaris (Pers.) Ach. not occurring in Colombia
Dirinaria subpicta (Nyl.) C.W. Dodge not occurring in Colombia Lecanora subcrenulata auct. non Lecanora tropica Zahlbr.
Echinoplaca argentea auct. non Aderkomyces couepiae Bat. Müll. Arg.
(Mont.) R. Sant. Lecidea anomala auct. non Ach. Biatora globulosa (Flörke) Fr.
Echinoplaca heterella auct. non Echinoplaca handelii (Zahlbr.) Lecidea atroalbella Nyl. Buellia aethalea (Ach.) Th. Fr.
(Stirt.) R. Sant. Lücking
Lecidea domingensis auct. non Letrouitia flavidula (Tuck.) Hafellner
Entyloma amaranthi Mol.-Val. Albugo bliti (Biv.) Kuntze (Pers.) Nyl.
Erioderma physcioides auct. non Erioderma divisum P.M. Jørg. & Arv. Lecidea endoleuca f. laurocerasi Bacidia proposita (Nyl.) Zahlbr.
Vain. auct. non (Delise) Nyl.
Leptogium inflexum auct. non Nyl. Leptogium burgessii (L.) Mont. Peltigera polydactyla auct. non Peltigera dolichorhiza (Nyl.) Nyl.
(Neck.) Hoffm.
Leptogium menziesii auct. non Leptogium andinum P.M. Jørg.
(Ach.) Mont. Petractis hypoleuca (Ach.) Vězda not occurring in Colombia
Leptogium pulchellum auct. non Leptogium corticola (Taylor) Tuck. Phaeographis subtigrina (Vain.) not occurring in Colombia
Ach. Zahlbr.
Leucocintractia leucoderma (Berk.) Leucocintractia scleriae (DC.) M. Phaeophyscia hirsuta (Mereschk.) not occurring in Colombia
M. Piepenbr. Piepenbr., Begerow & Oberw. Essl.
Lobaria discolor auct. non (Bory) Yoshimuriella deplanata (Nyl.) Phaeophyscia pusilloides (Zahlbr.) not occurring in Colombia
Hue Moncada & Lücking Essl.
Lobaria pulmonaria (L.) Hoffm. not occurring in Colombia Physcia adglutinata (Flörke) Nyl. Hyperphyscia confusa Essl., C.A.
Morse & S.D. Leav.
Lobaria quercizans Michx. not occurring in Colombia
Physcia adscendens H. Olivier Heterodermia sp.
Mazosia pseudobambusae auct. Mazosia bambusae (Vain.) R. Sant.
non Kalb & Vězda Physcia dactylifera Elix not occurring in Colombia
Microbotryum bistortarum (DC.) Sphacelotheca hydropiperis Physcia endochrysea auct. non Phaeophyscia endococcinodes
Vánky (Schumach.) de Bary (Nyl.) Hampe (Poelt) Essl.
Physcia endococcina auct. non Phaeophyscia endococcinodes
Mycoblastus glabrescens (Nyl.) not occurring in Colombia
(Körb.) Nyl. (Poelt) Essl.
Zahlbr.
Physcia obscura var. endococcina Phaeophyscia endococcinodes
Mycomicrothelia melanospora not occurring in Colombia
auct. non (Körb.) B. de Lesd. (Poelt) Essl.
(Hepp) D. Hawksw.
Physcia stellaris (L.) Fr. not occurring in Colombia
Myriostigma miniatum (Vain. ex not occurring in Colombia
Lücking) Aptroot, Ertz, Grube & M. Physcia stellaris (L.) Nyl. not occurring in Colombia
Cáceres Physcia stellaris f. rosulata auct. Physcia aipolia (Ehrh. ex Humb.)
Nephroma helveticum Ach. Nephroma tropicum (Müll. Arg.) non (Ach.) Nyl. Fürnr.
Zahlbr. Physcia tribacioides Nyl. not occurring in Colombia
Ocellularia interposita (Nyl.) Hale not occurring in Colombia Physcia verrucosa Moberg not occurring in Colombia
Ochrolechia pallescens (L.) A. Ochrolechia africana Vain. Platygramme impudica (A.W. Platygramme sp.
Massal. Archer) A.W. Archer
Ochrolechia pallescens (L.) A. Ochrolechia subpallescens Polycauliona phlogina (Ach.) Arup, Solitaria chrysophthalma (Degel.)
Massal. Verseghy Frödén & Søchting Arup, Søchting & Frödén
Ochrolechia upsaliensis (L.) A. not occurring in Colombia Polymeridium simulans R.C. Harris not occurring in Colombia
Massal.
Polymeridium subcinereum auct. Polymeridium pyrenuloides (Fée)
Pannaria mariana auct. non (Fr.) Pannaria mosenii C.W. Dodge non (Nyl.) R.C. Harris Aptroot
Müll. Arg.
Porina elegans Malme not occurring in Colombia
Parmelia americana auct. non Hypotrachyna vexans (Zahlbr. ex
(Meyen & Flot.) Mont. W.L. Culb. & C.F. Culb.) Divakar, A. Porina farinosa C. Knight not occurring in Colombia
Crespo, Sipman, Elix & Lumbsch Pseudevernia furfuracea (L.) Zopf Everniastrum sp.
Parmelia erumpens Kurok. not occurring in Colombia Pseudocyphellaria crocata (L.) Vain. Pseudocyphellaria citrina (Pers.)
Parmelia gracilis auct. non (Müll. Hypotrachyna physcioides (Nyl.) Lücking, Moncada & S. Stenroos
Arg.) Vain. Hale Pseudocyphellaria crocata (L.) Vain. Pseudocyphellaria sandwicensis
Parmelia kamtschadalis auct. non Hypotrachyna cirrhata (Fr.) Divakar, (Zahlbr.) Moncada & Lücking
(Ach.) Eschw. A. Crespo, Sipman, Elix & Lumbsch Pseudocyphellaria crocata (L.) Vain. Pseudocyphellaria xanthosticta
(Pers.) Moncada & Lücking
Parmelia kamtschadalis var. Hypotrachyna cirrhata (Fr.) Divakar,
americana auct. non (Meyen & A. Crespo, Sipman, Elix & Lumbsch Pseudocyphellaria faveolata (Delise) not occurring in Colombia
Flot.) Nyl. Malme
Rafael F. de Almeida is a biologist
Almeida is a biologist with expertise in Taxonomy and Systematics. He served as
Colombia (UPFC) project. Robert Lücking is a
research editor for the Useful Plants and Fungi of Colombia (UPFC) project. Robert
lichenologist at the Botanischer Garten und Botanisches Museum in Berlin, where he works as
shop.kew.org/kewbooksonline
curator for lichens, fungi, and bryophytes, with emphasis on the funga of South America. Aída
Vasco-Palacios is
Vasco-Palacios is a professor at the Faculty of Microbiology at the University of Antioquia, where ISBN 978–1–84246–790–9
she studies fungal biodiversity in Colombia and its biotechnological potential. Also, she is the
president of the Colombian Association of Mycologists. Ester Gaya is a Senior Research Leader in
the Trait Diversity and Function department at the Royal Botanic Gardens, Kew (RBG Kew), where
she leads the Comparative Fungal Biology team. Mauricio Diazgranados is
Diazgranados is a research leader
in the Ecosystem Stewardship department at RBG Kew. He focuses on investigating how useful
plants and fungi can contribute to developing nature-based solutions to tackle environmental
and societal challenges. He was also the principal investigator of the UPFC project. 9 781842 467909