INSTITUTO DE PESQUISAS JARDIM BOTÂNICO DO RIO DE JANEIRO

Transcription

1 Volume 66(4): Outubro Dezembro 2015

2 INSTITUTO DE PESQUISAS JARDIM BOTÂNICO DO RIO DE JANEIRO Rua Jardim Botânico Jardim Botânico - Rio de Janeiro - RJ - CEP JBRJ ISSN Rodriguésia A revista Rodriguésia uma publicação trimestral do Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, a qual foi criada em A Revista publica artigos científicos originais, de revisão, de opinião e notas científicas em diversas áreas da Biologia Vegetal (taxonomia, sistemática e evolução, fisiologia, fitoquímica, ultraestrutura, citologia, anatomia, palinologia, desenvolvimento, genética, biologia reprodutiva, ecologia, etnobotânica e filogeografia), bem como em História da Botânica e atividades ligadas a Jardins Botânicos. Ficha catalográfica Rodriguésia: revista do Jardim Botânico do Rio de Janeiro. -- Vol.1, n.1 (1935) -.- Rio de Janeiro: Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, v. : il. ; 28 cm. Trimestral Inclui resumos em português e inglês ISSN Botânica I. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro CDD CDU - 58(01) Indexação DOAJ EBSCO JSTOR Scopus SciELO Index of Botanical Publications (Harvard University Herbaria) Latindex Referativnyi Zhurnal Review of Plant Pathology Ulrich s International Periodicals Directory Esta publicação é afiliada à ABEC-Brasil Edição eletrônica ISSN:

3 Rodriguésia 66(4): Editorial/Sumário DOI: / Editorial Rodriguésia was created in the winter of 1935, as a joint managerial initiative of the Instituto de Biologia Vegetal, Jardim Botânico and the Estação Biológica de Itatiaya. The Comissão de Redacção in charge of editing the new journal was formed by P. Campos Porto, Fernando R. da Silveira and Leonam de A. Penna. The Amazonian tree from the coffee-family known as pau-mulato (Calycophyllum spruceanum), planted in one of the famous walkways within the Botanical Garden, was in the cover of the first volume. From that point onwards, the journal had the objective to serve the studious classes and to disseminate all subjects that referred to the life of the three above-mentioned institutions it represented, coming out four times a year to mark the start of each season. During its eight decates of live, Rodriguésia has played the role of disseminating scientific papers regarding the Brazilian and Neotropical Flora, featuring different lines of research. Over time, the journal strived to publish high quality articles, improve submission and evaluation procedures and to promote wider reach both for the national and international botanical community. Today Rodriguésia s editorial board comprises 30 researchers as well as a team of two professional editors. This quarterly journal is processed online and its content is freely available electronically and indexed by the SciELO Citation Index (SciELO CI), which has now been integrated into the Web of Science platform (WoS), Scopus and by the CAPES Periodical Portal. We are immensely pleased that the eightieth year anniversary of Rodriguésia is marked by an edition showcasing the World Flora Online (WFO) project, a direct response to the first target of the Global Strategy for Plant Conservation for 2020 and the conclusion of the Brazilian List of Plants and Fungi. The latter project ennummerates the species of plants and fungi in Brazil and becomes, from 2016, the framework for the new Brazilian Flora Online project. The life-long objective to accummulate and disseminate plant knowledge has always been at the heart of our journal, and it is a very appropriate vehicle for the publication of five articles of synthesis, focusing on the advances brought by the Brazilian List to the knowledge of the fungi, algae, bryophytes, ferns and lycophytes, and seed plants over the last five years. Therefore this volume counts with 15 articles presenting the current state of the World Flora Project, the situation of the floras of several biodiverse countries and regions, such as Malaysia, Cameroon, North America, Flora Neotropica, Bolivia and the Southern Cone of South America, besides the five articles mentioned above, a paper on Espirito Santo state s flora and another on allien plants in Brazil. The involvement and collective effort of the international botanical community towards preparing floras is expressed through the multiplicity of authors who collaborated in this volume of Rodriguésia. We invite everyone to read the articles and wish the results and interesting results presented in this commemorative volume play a role in stimulating botanists from Brazil and worldwide to participate in the Brazil Flora 2020 Project, with enthusiasm extending to other countries so that together we may meet the first target of the Global Strategy for Plant Conservation for Vidal de Freitas Mansano, Marli Pires Morim and Daniela Zappi

4 Rodriguésia 66(4): Editorial/Sumário DOI: / FOREWORD [Rodriguésia volume 80] It is with great pleasure and admiration that I write this foreword to the 80 th anniversary volume of the Rio de Janeiro Botanic Garden in-house journal Rodriguésia, which was chosen to become the vehicle for a series of articles in English setting the individual role of Brazil and other biodiverse countries and regions in relation with the Flora of the World project. In particular, this volume commemorates and assess the progress made since the launching in 2010 of the Lista de Plantas e Fungos do Brasil. During the last 5 years, 574 contributors have been responsible for the list that culminated this year with 45,352 species recorded, a number over 10% higher than what was published in the first edition, in The Lista do Brasil is fully established as the main reference point for Brazilian plant (both vascular and non-vascular) and fungal names and is widely used. As it continues to be available for the wide public, behind the scenes Brazilian botanists are organizing an even more ambitious endeavour, the launch of the Flora do Brasil online. In my previous role as Director for Biodiversity Conservation and later National Secretary for Biodiversity and Forests at the Brazilian Ministry of the Environment, I had the privilege to represent Brazil in many negotiations under the Convention on Biological Diversity, among which I would like to mention the role I played as coordinator of the negotiations at the Jardín Botánico Canario Viera y Clavijo in Las Palmas de Gran Canaria and at the Linnean Society in London that led to the establishment of the Global Strategy for Plant Conservation at COP 6, in the Hague, in 2002, which included as its first target to establish by 2010 a widely accessible working list of known plant species, as a step towards a complete world flora. In 2008 I led internal negotiations within the Brazilian Ministry of the Environment and its five institutes which led to the creation that year of the Centro Nacional de Conservação da Flora (CNCFlora) as part of the structure of the Rio de Janeiro Botanic Garden Research Institute and the transfer to this new center of resources from the National Biodiversity Mainstreaming Project (PROBIO II) funded by the Global Environment Facility (GEF), the Financial Mechanism of the Convention on Biological Diversity. The Lista do Brasil List was the first project which benefited from this funding through the CNCFlora. The launch in 2010 of the Lista de Plantas e Fungos do Brasil was a great satisfaction to all Brazilians as for the first time in a century we had access to a consolidated and updated list of all Brazilian plants, partially replacing the old Flora Brasiliensis published between 1840 and 1906 at the initiative of the great Bavarian botanist Carl Friedrich Philipp von Martius, who collected extensively in Brazil between 1817 and 1820! Given the richness of the Brazilian flora and the lack before 2010 of a consolidated list of Brazilian plants, this Lista do Brasil also represented a significant contribution for the achievement of GSPC initial target 1 at global level in Shortly after, in October 2010 in Nagoya, Japan, the Parties to the CBD adopted at COP 10 the current Strategic Plan for Biodiversity with its 20 Aichi Biodiversity Targets which has been recognized as the overarching agenda and framework for biodiversity for all the United Nations, all the biodiversity-related conventions and all major international organizations such as IUCN. An updated GSPC was also adopted at COP10 and recognized as part of the Strategic Plan for Biodiversity The updated GSPC has as its revised target 1 the commitment to establish by 2020 an online flora of all known plants. I am proud to confess that I was one of those that pushed for more ambitious targets in the revised GSPC, including this one, despite the disbelief of some botanists. Targets have to be ambitious and help mobilize efforts and support to achieve results which we would not normally achieve under business-as-usual scenarios.

5 This volume is introduced by Peter Wyse-Jackson s contribution about the World Flora online Project, followed by invited contributions from different countries and regions of the world, such as Malaysia, Cameroon, North America, Argentina and Southern Cone to share their experience regarding the organization of their checklists and floras. An essay further exposing the challenge to the Brazilian botanists regarding the size of their task still to come is followed by papers originated from data collected by the Lista do Brasil These papers analyse our current knowledge regarding the fungi and all plant groups in Brazil, stating the progress attained in five years of the project and the areas where more studies are still needed in order to have a complete snapshot of our fungal and plant biodiversity. I would like to end this foreword with a call to the Brazilian Government and its partners to continue to provide political and financial support to the continuous updating of the Lista de Plantas e Fungos do Brasil and to the ongoing efforts towards the Flora do Brasil online. Additionally, I would like to strongly recommend that Brazil takes the lead to discover the rich and mostly untapped flora of the Amazon Region, certainly the least studied and potentially the richest flora of the world - such endeavour will require significant increase in field work beyond the relatively well known floras around Belém and Manaus and the training of many new botanists, specially taxonomists, to dedicate their lives to unravel the flora of the Amazon Region. I believe this is an endeavour worth the highest priorities in Brazil and in the world which will certainly require the support of a consortium of the most capable botanical institutions in the world. The recent adoption by the Brazilian National Congress of the new Brazilian ABS Law (known in Brazil as the Biodiversity Law ) has removed the prior barriers and hurdles that impeded or delayed access to the rich Brazilian biodiversity. What Brazil needs now is an ambitious research program to discover its rich Amazonian biodiversity and transform it through research and technology into a major contributor to generate jobs and wealth for the country and its people. Braulio Ferreira de Souza Dias Executive Secretary of the Convention on Biological Diversity

6 Sumário/Contents Artigos Originais / Original Papers Developing a World Flora Online - a 2020 challenge to the world s botanists from the international community. Peter Wyse Jackson & James S. Miller The flora of Malaysia projects L.G. Saw & R.C.K. Chung The World Flora Online 2020 project: will Cameroon come up to the expectation? Jean Michel Onana Flora of North America North of Mexico Nancy R. Morin, Luc Brouillet & Geoffrey A. Levin Hard Copy to Digital: Flora Neotropica and the World Flora Online William Wayt Thomas & Melissa Tulig The Catalogue of Vascular Plants of the Southern Cone and the Flora of Argentina: their contribution to the World Flora Fernando O. Zuloaga & Manuel J. Belgrano Flora of Bolivia - where do we stand? RI. Meneses, S. Beck, E. García, M. Mercado, A. Araujo & M. Serrano Diversity of Brazilian Fungi Maia, L.C et al. Update of the Brazilian floristic list of Algae and Cyanobacteria Meneses, M. et al. Bryophytes diversity in Brazil Denise P. Costa & Denilson F. Peralta Diversity of ferns and lycophytes in Brazil Prado, J. et al. Growing knowledge: an overview of Seed Plant diversity in Brazil The Brazil Flora Group

7 Flora of Brazil Online: Can Brazil s botanists achieve their 2020 vision? Marli Pires Morim & Eimear M. Nic Lughadha The naturalized flora of Brazil: a step towards identifying future invasive non-native species Rafael Dudeque Zenni Angiosperm Checklist of Espírito Santo: using electronic tools to improve the knowledge of an Atlantic Forest biodiversity hotspot Valquíria Ferreira Dutra, Anderson Alves-Araújo & Tatiana Tavares Carrijo

8 Rodriguésia 66(4): DOI: / Developing a World Flora Online - a 2020 challenge to the world s botanists from the international community Peter Wyse Jackson 1,2 & James S. Miller 1 Abstract The Global Strategy for Plant Conservation of the Convention on Biological Diversity adopted as its first target in 2010 the preparation of An online flora of all known plants by This target was subsequently adopted by a broadly-based international consortium of botanical institutions that have committed themselves to undertaking this ambitious project. The preparation of a world flora will be the first modern and large-scale and comprehensive attempt to produce a comprehensive overview and baseline of knowledge on the world s plant diversity. This article outlines previous historic efforts to document the world s known flora. It also describes the ways in which the World Flora Online Consortium was created, how it is organized and its plans to compile diverse datasets available in digital formats into a single online portal available and open to all. Such data are being combined from geographical floristic accounts, such as the Flora of China, the Flora of North America and many other regional and national projects, together with relevant monographic treatments. Key words: Biodiversity, conservation, Global Strategy for Plant Conservation, taxonomy, world flora. Resumo A Estratégia Global para a Conservação das Plantas e a Convenção da Biodiversidade adotaram em 2010, como a primeira meta para 2020, a produção de uma flora online de todas as espécies conhecidas. Tal meta foi assumida por um amplo grupo de participantes provenientes de diversos institutos botânicos e esta comunidade comprometeu-se a atingir essa meta. O primeiro Consórcio da Flora On-line foi criado com a finalidade de preparação de uma flora mundial moderna e em larga-escala, provisionando a primeira tentativa de produzir uma estrutura para embasar a totalidade do conhecimento botânico mundial. O presente artigo expõe esforços históricos prévios realizados para documentar a flora do mundo, assim como a presente organização do Consórcio da Flora Online, cujo objetivo é compilar diversas bases de dados disponíveis em formatos digitais distintos num portal de acesso aberto ao público. Os dados estão sendo combinados a partir de tratamentos regionais distintos, como a Flora da China, Flora da América do Norte e muitos outros projetos regionais e nacionais, combinados a monografias relevantes. Palavras-chave: Biodiversidade, Conservação, Estratégia Global para a Conservação das Plantas, Taxonomia, Flora Mundial. Overview In 2002 the Convention on Biological Diversity (CBD) adopted a Global Strategy for Plant Conservation (GSPC), with the aim of halting the loss of plant diversity worldwide. It has now been endorsed by almost all of the countries of the world, and was renewed and updated on 29 th October, 2010 for the period up to 2020 by decision x/17 of the Conference of the Parties to the Convention on Biological Diversity. Incorporating 16 global targets, the Strategy includes within its objectives the need to document and understand plant diversity, to provide a baseline of knowledge on which plant conservation actions can be based. The Global Strategy for Plant Conservation was developed within the context of a serious crisis impacting on plant diversity worldwide. There are estimated to be approximately 300,000 known vascular plant species, perhaps another 50, ,000 await discovery (Joppa et al. 2010; Miller 2011). Currently there are known to be approximately 100,000 plant species that are 1 Missouri Botanical Garden, P.O. Box 299, St. Louis, MO , U.S.A. 2 Author for correspondence: Peter.Wysejackson@mobot.org

9 940 Wyse Jackson, P. & Miller, J.S. threatened by extinction in the wild. Preservation, management, and sustainable use of the world s biological resources is critical to maintain a careful balance between nature and development; otherwise, we will lose forever the biodiversity that supports us and our environment. Today, the world s biodiversity is being reduced at an accelerating rate without regard to its actual use and the potential it holds. Plants provide the basis for all terrestrial ecosystems and are a fundamental requirement in the maintenance of ecosystem services on which our survival depends. The rationale for the development of the GSPC was that plants are universally recognized as a vital part of the world s biological diversity and an essential resource for the planet. In addition to the small number of crop plants used for basic food and fibers, many thousands of wild plants have great economic and cultural importance and potential, providing food, medicine, fuel, clothing and shelter for vast numbers of people throughout the world. Plants play a key role in maintaining the planet's basic environmental balance and ecosystem stability and provide an important component of the habitats for the world's animal life. A driver for the development of the GSPC was the concern that many are in danger of extinction, threatened by habitat transformation, over-exploitation, alien invasive species, pollution and climate change. It was recognized that the potential disappearance of such vital and large amounts of biodiversity sets one of the greatest challenges for the world community: to halt the destruction of the plant diversity that is so essential to meet the present and future needs of humankind. The global strategy for plant conservation was proposed to address this challenge. While the entry point for the strategy is conservation, aspects of sustainable use and benefit-sharing are also included (Wyse Jackson & Kennedy 2009). As recognized in the GSPC, there is a tremendous need for a comprehensive documentation resource on the world s plant species, as a baseline to guide conservation effort and for plant resource planning and management. Information on the world s plants is presently dispersed in hundreds of journals and thousands of books, and indeed is by no means complete. It is generally retrieved by geographic location in various Floras that cover particular regions, which vary from small sites to continental. But the geographic ranges of plant species generally don t often conform with geo-political boundaries, so Floras don t usually present a complete set of information about the species that are included. There are also monographic works that do cover individual plant groups, generally genera or, less frequently, families, in a comprehensive manner. But there is only up-to-date information for a small percentage of the world s plant genera. As a consequence of the dispersed nature of available literature, it is often difficult for secondary users of information about plants to find what they need. The conservation community, for example, cannot easily retrieve information about the conservation status or comprehensive geographic distributional information of plants. Recent efforts to scan historical literature about plants and make it available electronically via the internet, such as the Biodiversity Heritage Library (< biodiversitylibrary.org/>), have improved access to the relevant literature, but it is still generally necessary to consult large numbers of references to retrieve any comprehensive information for the plants of a particular area. History of Global Plant Inventories Historically, many early botanists made efforts to comprehensively catalog the world s plants. When the Swedish botanist, Carl Linnaeus ( ) published Species Plantarum in 1753, it did in effect aim to be a world flora, but Linnaeus only knew about just short of 6,000 species in about 1,000 genera, a tiny percentage of the c.300,000 species known today. Linnaeus eventually expanded his knowledge to cover about 7,700 species of plants and believed that the world had no more than 10,000 species (Davis & Heywood 1973). The successors to Linnaeus built upon his work and their efforts greatly increased the known number of plant species but, within about a hundred years, the number was becoming insurmountable for an individual botanist to handle in a single publication, even in multiple volumes. The work of Augustin Pyramus De Candolle ( ), a Swiss botanist (born in France), was among the last efforts to comprehensively catalog the world s flora. His Prodromus Systematis Naturalis Regni Vegetabilis, widely known simply as the Prodromus, began in 1824 and eventually ran to 17 volumes, some published by other botanists after De Candolle s death (he was personally only able to finish seven volumes). It was near comprehensive for the time in relation to plant species known, but Rodriguésia 66(4):

10 Developing a World Flora Online a few parts were never completed. Collectively, the volumes of the Prodromus provide treatment for 58,975 species of Dicotyledons and Gymnosperms (Davis & Heywood 1973). The monocots and the genus Ficus were never completed though. Near the end of the 19th century, Adolf Engler ( ), a German botanist, began two monumental works intended to cover the whole plant kingdom. Die Naturlichen Pflanzenfamilien, which he started in 1887 with his colleague Karl Prantl ( ), provided a summary of plant families and genera and Das Pflanzenreich, from 1900 onwards, was a series of detailed generic monographs. Engler and his collaborators completed much of this work, but by the time they were working the number of plant species had grown so much that neither work was comprehensive when he passed away in 1930 (Davis & Heywood 1973). Following Engler, the number of known plant species had grown to a point where it was no longer considered an approachable project for traditional publication and subsequent botanists generally restricted publications to individual groups of plants or plants of regions substantially smaller than the whole world. Major Regional Plant Inventories Notable milestones in early regional completed plant inventories include The Flora of West Tropical Africa (Hutchinson & Dalziel ), prepared and published by the Royal Botanic Gardens, Kew, completed in 1936, world s first completed tropical regional flora. The revised second edition, covering 7,072 species in 1,742 genera, was finished in Another floristic milestone was Flora Europaea (Tutin et al ), a five volume comprehensive overview of the plants of Europe published between 1964 and 1993, the first continental Flora published. More recently the completion of The Flora of China (Wu & Raven ) represented another such milestone. The Flora of China was a 25 year an international collaborative project begun in 1988 to publish a comprehensive catalog of all Chinese wild vascular plants, with full descriptions of 31,500 species and illustrations of about 20,000 species. The first volume appeared in 1994 and, by March of 2013, 22 volumes of text and 22 volumes of illustrations have been published. The Flora of China was jointly published by the Missouri Botanical Garden Press (St. Louis) and Science Press (Beijing). The last 50 years have seen our knowledge of the plant kingdom grow to where there are presently 941 about 300,000 known and accepted plant species (Prance et al. 2000; Miller 2011). These have been described via both floristic and monographic works and most parts of the world have had significant floristic activity in recent years so information on most of the world s plants is there to be synthesized. Yet collectively the information is highly dispersed in the literature. But in the last 30 years, the assembly of various electronic databases and the development of the software to run them has again made the possibility of a comprehensive catalog of plants possible. Today the information exists to be compiled from modern floras and its rigorous review by specialists on various plant groups is an approachable project. The tools and content to produce a World Flora Online, called for in Target 1 of the Global Strategy for Plant Conservation, exist today. The Delivery of Target 1 of the GSPC, a World Flora Online The first phase of the Global Strategy for Plant Conservation, up to 2010, included as its first target to prepare A working list of known plant species, as a step towards a complete world flora. This target was completed through the collaboration of the Missouri Botanical Garden with the Royal Botanic Gardens, Kew, U.K. when The Plant List was launched at the end of 2010 (< theplantlist.org/>). This represented the culmination of major efforts made by both institutions and many other organizations too, to bring together multiple projects into this first ever global checklist. It aims to be comprehensive for species of Vascular plant (flowering plants, conifers, ferns and their allies) and of Bryophytes (mosses and liverworts). It does not include algae, fungi or plants only known as fossils. Version 1.1 of The Plant List was released in September It replaced Version 1.0 and included new data sets, updated versions of the original data sets and improved algorithms to resolve logical conflicts between those data sets. The Plant List provides the accepted Latin name for most species, with links to all synonyms by which that species has been known. Around 20% of plant names are still unresolved, indicating that the data sources included provided no evidence or view as to whether the name should be treated as accepted or not, or there were conflicting opinions that could not be readily resolved. It is fully recognized that The Plant List has limitations and that quality of content varies from one plant group to another in a Rodriguésia 66(4):

11 942 Wyse Jackson, P. & Miller, J.S. manner reflective of our present state of knowledge and what contributions were included. The Plant List is clearly a work in progress that aims to be a best effort list, to demonstrate progress and to stimulate further work. The Plant List includes 1,064,035 scientific plant names of species rank. Of these 350,699 (33.0%) are accepted species names. 470,624 (44.2%) names included are synonyms and 242, (22.8%) are unresolved. It currently contains contributions from the Royal Botanic Gardens, Kew, the Missouri Botanical Garden, Global Compositae Checklist, the International Legume Database and Information Service (ILDIS), the Royal Botanic Gardens, Edinburgh, the South African National Botanical Institute, Conservatoire et Jardin Botanique, Ville de Genève. A key tool used by the Missouri Botanical Garden to support its plant systematic research is the Tropicos database (< org/>), which will be a fundamental resource for contributions to a World Flora Online. Tropicos is the world s largest database of plant identification information, containing extensive research information for over 1.2 million plant names and over 4 million plant specimens. It is based upon and incorporates nearly 30 years of scientific research and data compilation by Garden staff and many others. Thousands of scientists at the Garden and around the world depend upon it to support their research in plant and conservation science. Botanical information from Tropicos represents a key component of the data content of The Plant List. When the GSPC was updated in 2010, a new 1 st target was adopted, to prepare An online flora of all known plants by 2020 (< int/gspc/strategy.shtml>). The World Flora Online (WFO) is a critical initiative required in order to meet the world s needs for knowledge on which to base plant conservation, ecological restoration and to sustain human use of plant species for a multitude of socio-economic purposes. It will also provide a fundamental resource to clarify where conservation needs are greatest and what gaps in knowledge exist. In January 2012 in St Louis, Missouri, U.S.A., representatives from four institutions: the Missouri Botanical Garden, the New York Botanical Garden, the Royal Botanic Garden Edinburgh, and the Royal Botanic Gardens, Kew - all members of the Global Partnership for Plant Conservation (GPPC) (< took the initiative to meet and discuss how to achieve GSPC Target 1 by The meeting resulted in a proposed outline of the scope and content of a World Flora Online, as well as a decision to form an international consortium of institutions and organizations to collaborate on providing that content (< meetings/sbstta/sbstta-16/information/sbstta-16- inf-38-en.pdf>). The World Flora Online project was subsequently launched in India, at an event held during the 11th Conference of the Parties (COP) to the Convention on Biological Diversity in October, 2012 where the COP also adopted a decision welcoming the World Flora Online initiative: The 11 th Conference of the Parties of the Convention on Biological Diversity Welcomes the initiative of the Missouri Botanical Garden, the New York Botanical Garden, the Royal Botanic Garden, Edinburgh, and the Royal Botanic Gardens, Kew, and their partner organizations and supporters worldwide, to lead the development of a World Flora Online by 2020 to facilitate the achievement of Target 1 of the Global Strategy for Plant Conservation. (UNEP/CBD/COP/DEC/ XI/26) (< cop-11-dec-26-en.pdf>). In January, 2013, a Memorandum of Understanding (MOU) for the World Flora Online was opened for signature. As of the end of August 2015, 29 institutions and organizations had signed the MOU, becoming part of the project and members of the World Flora Online Consortium (< worldfloraonline.org/>) (Tab. 1). A range of other institutions and organizations worldwide is also being invited to participate in the WFO Consortium. Through the MOU, a WFO Council was created, including representatives from each of the members of the Consortium. Council Members have met regularly since 2012: at the Missouri Botanical Garden (July 2012), the Royal Botanic Garden, Edinburgh (November 2013), the Komarov Institute, St. Petersburg, Russia (June 2014), and the Conservatoire et Jardin Botanique, Ville de Genève (February 2015) (Miller et al. 2014). Future meetings are currently planned for Rio de Janeiro (October 2015), New York (early 2016), and Capetown, South Africa (Fall 2016). The Council currently comprises 29 member institutions (Tab. 1) and is chaired by Peter Wyse Jackson, President of the Missouri Botanical Garden. Initially structured with four working groups including Governance, Technology, Taxonomy, and Data Gathering, there has been no continuing need for the Governance working group and the latter two have been combined. Rodriguésia 66(4):

12 Developing a World Flora Online 943 Table 1 Members of the World Flora Online Consortium* Australian Biological Resources Study Canberra, Australia Botanic Garden and Botanical Museum, Berlin-Dahlem, Dahlem Centre of Plant Science (DCPS), Berlin, Germany Botanic Garden Meise, Belgium Conservatoire et Jardin Botaniques, Genève, Switzerland Flora Iberica Project, Madrid, Spain Flora of North America Association, US & Canada Forest Research Institute Malaysia, Kuala Lumpur, Malaysia Global Biodiversity Information Facility, Copenhagen, Denmark Institute of Botany, Academy of Sciences of the Czech Republic, Prague, Czech Republic Institute of Botany, Azerbaijan National Academy of Sciences. Baku, Azerbaijan Institute of Botany, Chinese Academy of Sciences Beijing, China Institute of Botany, Slovak Academy of Sciences Bratislava, Slovakia Instituto de Botánica Darwinion, Buenos Aires, Argentina Instituto de Pesquisas Jardim Botânico do Rio de Janeiro Rio de Janeiro, Brazil Instituto Nacional de Biodiversidad (INBio), Costa Rica Komarov Institute of Botany, Russian Academy of Sciences, St. Petersburg, Russia Kunming Institute of Botany, Chinese Academy of Sciences, Kunming, China Missouri Botanical Garden, St. Louis, Missouri, USA Muséum National d'histoire Naturelle, Paris, France Natural History Museum, London, UK Naturalis Biodiversity Center, Leiden, Netherlands Royal Botanic Garden Edinburgh, Edinburgh, UK Royal Botanic Gardens, Kew, London, UK Smithsonian National Museum of Natural History, Washington, DC, USA South African National Biodiversity Institute, Pretoria, South Africa The New York Botanical Garden, New York, NY, USA Botany Department of Trinity College Dublin, Dublin, Ireland Tsitsin Main Botanical Garden, Russian Academy of Sciences, Moscow, Russia UNESCO Chair in Plant Conservation and Biodiversity in Macaronesia and in Western Africa Gran Canaria, Spain * Members, as of 31 st May, 2015 Currently the Technology working group is chaired by Chuck Miller (Missouri Botanical Garden) and Mark Watson (Royal Botanic Garden, Edinburgh) and the Taxonomy working group is chaired by Thomas Borsch (Berlin-Dahlem Botanical Garden) and David Simpson (Royal Botanic Garden, Kew). Scope and Content of the World Flora Online The World Flora Online is planned to be an open-access, web-based compendium of the world s plant species. The World Flora Online will consist of a nomenclatural backbone, placing all names not accepted in proper synonomy. Associated with the name of each accepted species will be descriptions, primarily from previously published works, illustrations, photos, conservation assessments when available, and other information. Descriptions and keys will also be included with families and genera. Information to be included has been discussed at each of the previous WFO Council meetings and the target audiences anticipated to benefit from consulting the WFO are summarized in Table 2. Rodriguésia 66(4):

13 944 Wyse Jackson, P. & Miller, J.S. It was agreed in the earliest organizational meetings that it would be impossible to generate new information, particularly descriptions, for the approximately 300,000 currently known plant species therefore it would need to rely primarily on existing descriptions, distributional information and identification tools. In line with The Plant List, the focus of the WFO will be on comprehensive species level information for species of Vascular plant (flowering plants, conifers, ferns and their allies) and of Bryophytes (mosses and liverworts), but not including algae, fungi or plants only known as fossils. Already very extensive data are held by a wide variety of botanical organizations and institutions worldwide which can be incorporated to provide a solid basis for the WFO development. For example, the Missouri Botanical Garden can contribute information that results from more than 40 years of activity in efforts to complete floristic inventories of various parts of the world. This will include contributions from the Flora of North America, Flora Mesoamericana, Flora of the Venezuelan Guayana, Flora of China, the Madagascar Catalog (including the other islands of the SW Indian Ocean), and checklists of Ecuador, Peru, Bolivia, and the South American Southern Cone (Argentina, Chile, Paraguay, and Uruguay). Another example of existing data of value to the WFO project is the emonocot project, led by the Royal Botanic Gardens, Kew (RBG Kew) (< emonocot has been developed by a collaborative team from the Table 2 WFO Potential Audiences Conservationists Plant taxonomists Other scientists General interest groups Contributors Primary data providers Information converters and taxonomic curators Expert taxonomic reviewers Technical data and systems managers Other stakeholders Institutional interests Taxonomic research planners and evaluators UK, consisting of RBG Kew, the Natural History Museum (NHM) and Oxford University. Monocots constitute approximately 23% (70,000 species) of all higher plants and include numerous groups with large numbers of species of economic, conservation or ecological importance (such as palms, orchids, aroids, grasses, and sedges). The objective of this project has been to produce a web-based treatment of monocot plants. The WFO Council has decided to use software prepared for emonocot, provided by RBG Kew, for the development of a public portal for the World Flora Online. Another important initiative relevant to the World Flora Online is The Global Plants Initiative (GPI) that is making available more than 1.8 million plant type specimens and other resources online to support floristic research around the world (< global-plants>). In 2013, JSTOR released Global Plants, a new community-contributed online database for scientific researchers, conservationists and others engaged in studying the world s plant biodiversity. The GPI has been a significant international undertaking by leading herbaria to digitize and make available via JSTOR plant type specimens and other holdings used by botanists and others especially those working in botany and botanical research. Funded for many years by the Andrew W. Mellon Foundation, its partners include more than 300 institutions in more than 70 countries. Founding partners include the Missouri Botanical Garden; Muséum National d Histoire Naturelle (MNHN), Paris; The National Herbarium, Addis Ababa University; New York Botanical Garden; the Royal Botanic Gardens Kew; and the South African National Biodiversity Institute. Efforts to make digital images of type specimens for Global Plants, have also encouraged additional digitization, one major example being the work of MNHN where this herbarium has digitized over 6 million vascular plant specimens plus an additional 3 million cryptogam specimens (< institution/mnhn/search>). The team at MNHN also coordinated the GPI project in Francophone West Africa, including in Mauritania, Senegal, Guinea, Burkina Faso and Togo. At national levels, many floristic treatments can also be incorporated into the World Flora Online. An example is the Flora of Nepal project that is being coordinated by the Royal Botanic Garden Edinburgh, in collaboration with the Nepal Rodriguésia 66(4):

14 Developing a World Flora Online government s Department of Plant Resources and Tribhuvan University in Kathmandu and the University of Tokyo. The accounts of Nepal s 7,000 species are being published in ten volumes, and all this information and the data from which they have been compiled is also available through the project s website (< In Brazil, the Brazilian Council for Research in Science and Technology, the Jardim Botânico do Rio de Janeiro (JBRJ) and a range of Brazil-based sponsors and other organizations are cooperating to deliver the REFLORA programme, designed to facilitate the creation of an on-line Flora do Brasil Digital, Brazil s response to GSPC Target 1 (< reflora.jbrj.gov.br/jabot/principaluc/principaluc. do>; Forzza et al. 2012). To support this project several herbaria internationally are creating high resolution images of herbarium specimens of most relevance for study of the Brazilian flora and making these available to JBRJ for inclusion in the Reflora Virtual Herbarium. The Reflora Virtual Herbarium is fully integrated with the List of the Brazilian Flora first published in Achieving the World Flora Online and its Potential Impact This ambitious collaborative international effort to achieve a World Flora Online will build upon existing floras, checklists, monographs, and other published research, but will also collect and generate information on poorly-known plants and plant groups and unexplored regions by engaging botanists with the most field experience and expertise in these plants or regions. The WFO will be built on a nomenclatural backbone, including all names from the International Plant Names Index (IPNI) as organized in The Plant List. This will be reviewed by taxonomic specialists for each family/genus to determine if names represent accepted species or where they should be placed in synonomy. Hopefully the percentage of unresolved names will be significantly reduced. Basic information on geographic distribution will be included. A particular emphasis will be on providing baseline data at country level, which will support the use of such information by countries implementing national-level biodiversity conservation programmes. The WFO will also include descriptions, presumably with the vast majority from previously published sources, with their source clearly identified. Keys may be included when available, but it was also agreed that dichotomous keys to large taxa with world-wide distributions are difficult if not near impossible and 945 of limited value. Vouchered images of living plants, images of specimens, and images of illustrations will also be included. Conservation assessments will be included when available. The WFO will be primarily in English, but will include data in the languages in which it is provided. It is not intended that it will be comprehensively multi-lingual - at least in the first phases. The WFO is not intended to provide critical, monographic treatments of all of the world s plant species, nor will it be a detailed local Flora with vouchered distributional data. Nevertheless it will provide the first widely-accessible and all-inclusive global overview of the world s plants. It has been more than 150 years since any attempt came close to assembling a comprehensive catalog of all of the world s plant species. The WFO will provide for the first time in modern history, a single information resource that presents basic information on all of the world s plants. This will undoubtedly become the most regularly consulted resource by the world s plant research community and will probably serve the ongoing function of continuing to incorporate all new advances in plant science. But it will also serve many other important functions. For the first time ever, it will be available to the conservation community to provide information at the species level and it will be the foundation for achieving the second target of the Global Strategy for Plant Conservation, a comprehensive list of the world s threatened plant species. It is further anticipated that the WFO will also be regularly consulted as the definitive source of information on plants by broad audiences including educators, gardeners, and those interested in nature in general. The entire botanical research community will benefit greatly from having a comprehensive information resource on the world s plants available and it would be of immediate and direct benefit to advancing research efforts. The World Flora Online will also be of immediate and benefit to the conservation community and will support vastly better efforts to ensure future survival of all plant species. The WFO Council has analyzed the potential users of the WFO (Table 2) and, through its work, it will aim to develop the project so that it addresses best the uses to which it will be put. A primary purpose is to provide the baseline of information required for the achievement of many other targets of the GSPC. For example, Target two of the Global Strategy for Plant Conservation calls for the development of a comprehensive global list of Rodriguésia 66(4):

15 946 Wyse Jackson, P. & Miller, J.S. endangered species and this cannot be generated without a comprehensive resource on the world s plant species. The World Flora Online will also benefit any other scientific research aimed at the discovery and development of products from plants including nutritional, pharmaceutical, and agricultural products (Miller 2011). The world continues to face extremely daunting and challenging environmental problems and many feel that human activities are triggering the largest extinction event in the history of the planet. The Global Strategy for Plant Conservation provides sixteen targets that aim to stem and control the negative impact humans are having on the planet, but achieving some meaningful success for all sixteen targets is heavily dependent on achieving targets 1 and 2, and 2 cannot be achieved without 1. While the WFO will be a great tool for plant scientists and a wonderful resource for anyone with an interest in plants, it will also be a critical tool to help meet the conservation goals essential to preserve the biological integrity of the earth. References Biodiversity Heritage Library. Available at < biodiversitylibrary.org/>. Access on 7 July Davis, P.H. & Heywood, V.H Principles and Angiosperm Taxonomy. Robert E. Krieger Publishing Company, Huntington, New York. 558p. emonocot. An online resource for monocot plants. Available at < Access on 7 July Flora of Nepal. Available at < Access on 4 August Forzza, R.C.; Baumgratz, J.F.A.; Bicudo, C.E.M.; Canhos, D.A.L.; Carvalho Jr., A.A.; Coelho, M.A.N.; Costa, A.F.; Costa, D.P.; Hopkins, M.G.; Leitman, P.M.; Lohmann, L.G.; Lughadha, E.N.; Maia, L.C.; Martinelli, G.; Menezes, M.; Morim, M.P.; Peixoto, A.L.; Pirani, J. R.; Prado, J.; Queiroz, L.P., Souza, S.; Souza, V. C.; Stehmann, J.R.; Sylvestre, L.S.; Walter, B.M.T. & Zappi, D.C New Brazilian floristic list hightlights conservation challenges. Bioscience 62: Global Plants. Available at < content/global-plants>. Access on 7 July Global Strategy for Plant Conservation (GSPC). Available at < shtml>. Access on 4 August Global Strategy for Plant Conservation (GSPC). Available at < Access on 4 August Global Strategy for Plant Conservation (GSPC): World Flora online by Available at < cbd.int/doc/meetings/sbstta/sbstta-16/information/ sbstta-16-inf-38-en.pdf>. Access on 7 July Hutchinston, J. & Dalziel, J.M Flora of West Tropical Africa: the British west African territories, Liberia, the French and Portuguese territories south of latitude 18 N to Lake Chad and Fernando Po. 3 vols. Crown Agents for Overseas Governments and Administrations, London. Vol. 1 & 3, 2 parts, Vol. 1 part 1, pp , part 2, pp Vol. 2 pp Vol. 3 part 1, pp , part 2, pp Joppa, L.N; Roberts, D.L. & Pimm, S.L How many species of flowering plants are there? Proceedings of the Royal Society B: Biological Sciences 278: Miller, J.S The discovery of medicines from plants: a current biological perspective. Economic Botany 65: Miller, J.S.; Thomas, W.W.; Watson, M.; Simpson, D. & Wyse Jackson, P World Flora Online Council met in St. Petersburg. Taxon 63: 959. Muséum National d Histoire Naturelle (MNHN). Available at < mnhn/search>. Access on 4 August Prance, G.T.; Beentje, H.; Dransfield, J. & Johns, R The tropical flora remains under-collected. Annals of the Missouri Botanical Garden 87: Reflora. Available at < PrincipalUC/PrincipalUC.do>. Access on 4 August The Plant List. Available at < org/>. Access on 7 July Tropicos database. Available at < org/>. Access on 4 August Tutin, T.G.; Heywood, V.H.; Barges, N.A.; Valentine, D.H.; Winters, S.M. & Webb, D.A Flora Europaea. 5 vols. Cambridge University Press, Cambridge. World Flora Online. Available at < worldfloraonline.org/>. Access 4 August Wu, Z. & Raven, P.H Flora of China. 25 vols. Missouri Botanical Garden Press, St. Louis; Science Press, Beijing. Wyse Jackson, P.W. & Kennedy, K The Global Strategy for Plant Conservation: a challenge and opportunity for the international community. Trends in Plant Science: Special Issue: Plant science research in botanic gardens 14: Artigo recebido em 07/07/2015. Aceito para publicação em 22/09/2015. Rodriguésia 66(4):

16 Rodriguésia 66(4): DOI: / The flora of Malaysia projects L.G. Saw 1,2 & R.C.K. Chung 1 Abstract Malaysia has an estimated 15,000 species of vascular plants. Located in the Malesian region, its affinity is Sundaic, having common elements with Sumatra, Java and Palawan. The two halves of Malaysia, Peninsular Malaysia extending from mainland Asia and East Malaysian states of Sabah and Sarawak on the island of Borneo have their own distinct floristic components. Peninsular Malaysia has about 8,200 species of vascular plants and Sabah and Sarawak have an estimated 12,000 species. The flora of Sabah and Sarawak is generally richer than that of Peninsular Malaysia. Due to historical reasons, the flora of Malaysia project is planned in a phased approach. The Tree Flora of Sabah and Sarawak project, initiated in 1991, represents the first systematic modern attempt to document some of the important plant families of these two states. It would take at least another ten years to complete this project. It is estimated that the Flora of Peninsular Malaysia project, initiated in 2005, will take at least 20 years to complete. To achieve these ambitions, there must be continual support in skilled manpower and fund allocations. Key words: Flora, Malaysia, botanical revisions. Resumo A Malásia tem um número estimado de espécies de plantas vasculares. Localizada na região da Malesia, possui afinidade com a região conhecida como Sundaica, dividindo elementos em comum com a Sumatra, Java e Palawan. A Malásia é constituída por duas partes, sendo que a Malásia peninsular se estende a partir da Ásia continental, enquanto os estados malaios orientais de Sabah e Sarawak, localizados na ilha de Bornéu, têm componentes florísticos particulares e distintos. A Malásia peninsular tem cerca de espécies de plantas vasculares e Sabah e Sarawak têm um número estimado de espécies. A flora de Sabah e Sarawak é geralmente mais rica do que a península da Malásia. Devido a razões históricas, o projeto flora da Malásia preve uma abordagem em fases distintas. O Projeto Flora Arbórea de Sabah e Sarawak, iniciado em 1991, representa a primeira tentativa moderna e sistemática para documentar algumas das famílias de plantas importantes destes dois estados. Estima-se que sejam necessários de pelo menos mais 10 anos para concluir este projeto. Enquanto isso o Projeto Flora da Maláisa peninsular, iniciado em 2005, vai demorar pelo menos 20 anos para ser concluído. Para concretizar estes planos, deve haver apoio contínuo para treinar e manter mão de obra qualificada e obtenção de financiamento. Palavras-chave: Biodiversidade, ficologia, lista de plantas e fungos do Brasil. Introduction Malaysia is located just north of the equator, straddling between 0 51 N and 7 25 N with a total landmass of 329,847 square kilometres. The country is separated by the South China Sea into two regions of similar size; Peninsular Malaysia connected to mainland Asia on the western half and the states of Sabah and Sarawak on the island of Borneo on the east. It has a warm equatorial climate with little seasonality, hot and humid throughout the year (Fig. 1). The two geographical halves of Malaysia pose interesting challenges towards documenting the Flora of Malaysia. Peninsular Malaysia, part of the Malay Peninsula (here includes Singapore and Peninsular Thailand) contains the floristic elements of the Sunda Self and also of the mainland Asiatic species from seasonal climates (Wong 1998). Borneo, with its greater isolation from Malay Peninsula, has a flora of Sundaic element; however it has local floristic influences. Therefore these two regions result from quite different botanical past histories. A very brief historical perspective is provided here, highlighting 1 Forest Research Institute Malaysia, Kepong, Selangor, Malaysia. 2 Author for correspondence: sawlg@frim.gov.my

17 948 Saw, L.G. & Chung, R.C.K. Figure 1 Map of Malaysia. only the major works that are significance to the flora of these two regions. A more detailed account of the historical works relating to the flora of both Borneo and Malay Peninsula can be obtained from the introductory volumes of Flora Malesiana (de Wit 1949, van Steenis-Kruseman 1950, van Steenis, 1955). Wong (1987, 1995a) and Soepadmo (1999) also provided reviews with additional updates from de Wit, van Steenis and van Steenis-Kruseman of the botanical collection and documentation of the flora of both Peninsular Malaysia and Borneo. We present here an updated paper in 2005 describing the modern floristic projects we have implemented in Malaysia (see Saw & Chung 2007). Methods This paper reviewed botanical collections, research, administration and history based on literature, experience for the flora of Malaysia. Acronym of herbaria mentioned in the paper follows Index Herbariorum (Thiers 2015 < nybg.org/science2/indexherbariorum.asp>). Results and Discussion Botanical History of Peninsular Malaysia Peninsular Malaysia, with a more direct former British Colonial rule had a longer and more sustained period of botanical exploration and enumeration. Its botanical history dates back to the first British settlement in the early 1800 s in Penang, when the island was important for the spice trade. One of the most important collectors during this period include N. Wallich whose collection, organized in a catalogue (Wallich s catalogue), included contributions from G. Porter, W. Jack and G. Finlayson. Numbering about 8,000 species, Wallich s catalogue became the basis of many plant names for Penang and Singapore in Malaya (British colonial boundaries of present day Peninsular Malaysia and Singapore), and India (van Steenis-Kruseman 1950; or see < summary>). W. Griffith, Wallich s predecessor, collected large numbers of specimens particularly from Malacca and also form the basis of botanical work in Malaya. Many collectors followed included L. Wray Jr., Father Scortechini, H. Kunstler (often labelled as King s Collector), A.C. Maingay, C. Curtis, C.B. Kloss, R. Derry, T. Oxley, J.S. Goodenough, I.H. Burkill, Mohamad Haniff, N. Cantley, F.W. Foxworthy, etc.). A full list of these collectors has been presented by Burkill (1927) with some details of their background while their collection itineraries can be obtained from van Steenis-Kruseman (1950). H.N. Ridley s arrival to Malaya is a very significant moment for Malayan botany. Between 1888 and 1900, he was appointed as Director of Rodriguésia 66(4):

18 The flora of Malaysia projects Gardens and Forests, Straits Settlements and in , Director of the Gardens, Singapore. Ridley was a man of great ability who contributed most significantly towards the botany of Peninsular Malaysia. In his career, he described over 4,200 plant species. He amassed a huge collection amounting to about 50,000 numbers, of which the main set is at Kew with duplicates in Singapore and other herbaria (van Steenis-Kruseman 1950). No other collector has amassed collections at that rate for for Malaya ever since. Subsequent directors and curators of the herbarium at Singapore Botanic Gardens continued to build upon the foundation set up by Ridley. Of particular importance were the contributions from I.H. Burkill, M.R Henderson, E.H.J. Corner, R.E. Holttum and C.X. Furtado. All of them contributed in the exploration, carried out collections and publications, giving us a better understanding of the flora of Peninsular Malaysia. At the turn of the twentieth century, A.M. Burn-Murdoch set up a forest herbarium in Kuala Lumpur, with the aim of producing an account of the commercially important tree species of Malaya (Wong 1987). The specimens were collected as reference specimens and duplicates were submitted to H.N. Ridley for identification. The small herbarium was at the office of the Conservator of Forests, Strait Settlements and Federated Malay States. His successor, G.E.S. Cubitt continued with the collection although at a slower rate. In 1916, the Wray Herbarium of the Agriculture Department was transferred to the Forest Department in Kuala Lumpur (Cubitt 1919). In 1918 Cubitt secured the services of F.W. Foxworthy, as the first Forest Research Officer of the Federated Malay States and Straits Settlements. Under Foxworthy the herbarium grew quickly. By the end of 1920, the herbarium contained over 6000 numbers (Wong 1987). With the decision to form a Forest Research Institute (FRI), an area of about 324 hectares was acquired at Kepong in 1926 and the main office building was constructed in 1929 with the herbarium moving into the east wing of the building. C.F. Symington joined FRI in 1929 and began to assist the running of the herbarium. He contributed a large collection to the Kepong herbarium focusing on the important timber family Dipterocarpaceae for which he was preparing a foresters manual. By the time second World War broke out in Malaya with J.G. Watson having succeeded Foxworthy, the herbarium had about 43,000 specimens. Unfortunately, during the 949 war many of the specimens were badly damaged when looters plundered the herbarium. With the internment of the British officers, V.L. Bain, a Eurasian being exempted from detention was appointed acting State Forest Officer of Selangor. He was able to reappoint several local staff members at Kepong. Aziz Budin went on to restore the damaged collection and attempted to replace some of the lost specimens either by obtaining duplicates from other herbaria or by making new collections. After the war, J. Wyatt-Smith took charged of the herbarium and the collection gained momentum. With the formation of the Federation of Malaya in 1957 and subsequently the Federation of Malaysia in 1963, the transition towards Malaysianisation came into being. K.M. Kochummen who joined FRI as Assistant Botanist in 1953 subsequently took charge of the herbarium in In his career, he collected over 5,000 specimens; Kochummen was an important contributor towards many tree family revisions for both the Tree Flora of Malaya and Tree Flora of Sabah and Sarawak. In 1964, F.S.P. Ng was recruited as Forest Botanist. By 1965, the collection at FRI numbered over 74,000 specimens (Wong 1987). In 1965, T.C. Whitmore was engaged under the Colombo Plan to lead the Tree Flora of Malaya project (Whitmore 1972). In the years following, Whitmore conducted large collecting expeditions into many parts of Peninsular Malaysia, in particular places not collected previously. The Tree Flora project completed its last volume with the publication of Volume 4 in 1989 (Ng 1989). By then the herbarium has accumulated about 130,000 specimens. In 1980, K.M. Wong joined Kochummen managing the FRI collection. L.G. Saw joined the institute in 1982 as Hill Forest Silviculturist, later in 1985 joined the herbarium to study under Kochummen as he was soon to retire. In 1985, the Forest Research Institute Malaysia (FRIM) was formed as a statutory body from FRI and in the years following; the mandate of FRIM was to expand beyond the forestry related flora research it traditionally worked on to now include the study of the entire flora of Malaysia. In this much summarised collections survey we had to leave out many collectors which would feature in a wider review. Later botanists to join the much expanded role of FRIM included Farah Ghani (deceased), Idris Mohd. Said (since left), L.S.L. Chua, R.C.K. Chung, Y.Y. Sam, E. Soepadmo and Ruth Kiew. Since the initiation of the Tree Flora of Rodriguésia 66(4):

19 950 Saw, L.G. & Chung, R.C.K. Sabah and Sarawak and later the Flora of Peninsular Malaysia projects, the herbarium at the Forest Research Institute Malaysia has seen a tremendous increase in its botanical staff and capabilities. Bibliography of the Flora of Peninsular Malaysia In the following account, we have restricted the discussion to the main floristic publications that pertain to the flora of the Malay Peninsula. Other incidental accounts of local checklists and revisions of genera can be obtained from the bibliography found in the general chapter of Flora Malesiana Volume 5 (van Steenis 1955) and Turner (1997). The Flora of British India was the first major account covering all the families of Malay Peninsula. The scope of the volumes was to include plants within the British territories of India, together with those of Kashmir and Western Tibet, and Malaya (Hooker in 7 volumes) as part of the British colony. Plants from Borneo however, were not included in the revisions. Although the Flora of India included treatment of the Flora of the Malay Peninsula, it became apparent that it was not warranted from a phytogeographic perspective and the manner of treatment produced from limited data available at the time had produced an unsatisfactory revision (de Wit 1949). As a result, King (1889), working from the Calcutta Herbarium, initiated the series Materials for a Flora of the Malayan Peninsula. The revisions, written by various authors, were originally published as separate papers in the Journal of the Asiatic Society of Bengal. King and Gamble subsequently compiled these instalments into 4 volumes. King died after completion of Volume 4 and the work of editorship was passed on to J.S. Gamble who continued the series to instalment 26 which were, together with the last instalment published in 1936 (Ng & Jacobs 1983) compiled into Vol. 5. Nevertheless, these volumes covered only the dicotyledonous families and even so, the Urticales viz. Cannabinaceae, Moraceae, Ulmaceae, Urticaceae and most of the Euphorbiaceae never appeared in print (Ng & Jacobs 1983). Ridley (1907) published separately in Singapore, in three parts, the Materials for a Flora of the Malayan Peninsula completing the monocotyledons. These publications were very important ground-breaking works and they become the basis for subsequent work on the Flora of the Malayan Peninsula. Using the Materials as foundation for the Flora of the Malay Peninsula, Ridley upon his retirement completed the Flora of Malay Peninsula and published them in 5 volumes between 1922 and 1925 (Ridley ) for the angiosperms and a separate final fern instalment in 1926 (Ridley 1926). Following Ridley s publication of the Flora of Malay Peninsula, botanical work continued in more detail and from different perspectives. Burkill (1935), succeeding Ridley, subsequently produced two volumes of A Dictionary of the Economic Products of the Malay Peninsula. Other important publications from Singapore included Corner s (1940) Wayside Trees of Malaya, Henderson s (1959, 1954) Malayan Wild Flowers. By the 1950s, a revised Flora of Malaya was initiated as knowledge of the Malayan flora improved with more explorations and collections. A number of publications followed, mainly Zingiberaceae (Holttum 1950), Marantaceae (Holttum 1951), bamboos (Holttum 1958), orchids (Holttum 1964) and ferns (Holttum 1968). The volume on grasses was published by Gilliland (1971). Following the retirement of Holttum from active research due to his old age, the revised Flora of Malaya was more or less discontinued. Piggott (1988) produced a popular photographic account for ferns. The orchid flora was again subjected to another revision by Seidenfaden & Wood (1992). Turner (1997) collated a checklist of Peninsular Malaysian flora based on literature. More recently, Clarke (2001) published the Nepenthes of Sumatra and Peninsular Malaysia and Kiew (2005) revised the Begonias of Peninsular Malaysia in richly illustrated volumes. At the Forest Research Institute, Kepong, interest was towards tree species and identification manuals for foresters for the more important timber tree families and other minor forest products. Burn-Murdoch (1911, 1912) initiated the first publications of such foresters manual with the publication of the Trees and Timbers of the Malay Peninsula. The Malayan Forest Records series was started and Foxworthy published a number of volumes on commercial timbers and minor forest products (Foxworthy 1921, 1922, 1932). In 1934, C.F. Symington was appointed the first Forest Botanist and he envisaged producing a foresters tree manual comprising all the Malayan timberproducing families. However, it was obvious that much research was still required and that a great deal of instability still existed in the botanical Rodriguésia 66(4):

20 The flora of Malaysia projects nomenclature. He then concentrated on the most important timber family, the Dipterocarpaceae, which he completed in 1940 and was published as the Foresters Manual of Dipterocarps in 1943 in his absence (Symington 1943). Symington had since left Malaya just before the Japanese occupation of Malaya during the Second World War (Symington 1974). After the War, John Wyatt-Smith served as Forest Botanist. Wyatt-Smith also saw the importance of Symington s work and the need to document similar information on timber trees of other families. However, it became evident that the botanical knowledge of the many non-dipterocarp trees was inadequate for a similar treatment. In the interim, Wyatt-Smith (1952) produced a booklet listing the more common timber species found in Malaya. K.M. Kochummen subsequently revised this Pocket Check List three times to include new information. The Pocket Check List has now become an important identification reference for students and foresters for the common Peninsular Malaysian timber species. The book also standardized vernacular names for timber species. Wyatt-Smith also produced a series of other more taxonomic publications on some of the important timber families such as Burseraceae (Wyatt-Smith 1953a), Leguminosae (Wyatt-Smith 1953b), Myristicaceae (Wyatt-Smith 1953c), Sapotaceae (Wyatt-Smith 1954a), Lauraceae (Wyatt-Smith 1954b) and Sapindaceae (Wyatt-Smith 1954c), and the genus Calophyllum (Guttiferae, Henderson & Wyatt-Smith 1956). The Tree Flora of Malaya project under T.C. Whitmore as editor published two volumes (Whitmore 1972, 1973) followed by another two volumes with Ng (1978, 1989) as editor. The final four volumes covered over 2,800 species of trees found in Malaya. Interests in non-timber but commercially important groups resulted in the production of Dransfield s (1979) A manual of the rattans of the Malay Peninsula and K.M. Wong s (1995b) The bamboos of Peninsular Malaysia. Botanical History of Sabah and Sarawak (and Borneo) Sabah and Sarawak lacked the collection intensity of Malaya in the early years. However, recent years saw both herbaria at the Forest Research Centres of Sandakan and Kuching steadily increasing their collections. Wong (1995a) has amply summarised the collection history and 951 bibliography of Bornean flora in the introductory chapters of the Tree Flora of Sabah and Sarawak Volume 1 and we shall not elaborate further here. Suffice to add since that review, the Tree Flora of Sabah and Sarawak has published eight volumes (Soepadmo & Wong 1995; Soepadmo et al.1996; Soepadmo & Saw 2000; Soepadmo et al. 2002, 2004, 2007, 2011, 2014). The Plants of Kinabalu project led by Beaman published five volumes (Parris et al. 1992; Wood et al. 1993; Beaman & Beaman 1998; Beaman et al. 2001; Beaman & Anderson 2004). Modern identification manuals, amounting to floristic enumerations, of the rattans of Sabah and Sarawak ( Dransfield 1984, 1992), and the bamboos of Sabah ( Dransfield 1992) have been published. More charismatic groups such as orchids and Nepenthes continue to attract interest with publication of checklist of the Orchids of Borneo (Wood & Cribb 1994), Slipper Orchids of Borneo (Cribb 1997) and the Orchids of Borneo (Beaman et al. 2001), and Nepenthes of Borneo (Clarke 1997). Richly illustrated Etlingera (Zingiberaceae) of Borneo (Poulsen 2006) and Rhododendron (Ericaceae) of Sabah (Argent et al. 2007) were also recently published. The Flora of Malaysia - What Do We Know? Currently there is no comprehensive and up-to-date checklist for the flora of Malaysia. A number of checklists exist as a result of the different botanical history of the two main regions of Malaysia. For Peninsular Malaysia, the now outdated work of Ridley ( ) provided the first complete enumeration of the vascular plants of the Malay Peninsula; the angiosperms were published in the five volumes between 1922 and Subsequently, Ridley published a separate checklist of the ferns (Ridley 1926). Turner s (1997) publication of A Catalogue of the Vascular Plants of Malaya serves as the most recent checklist for Peninsular Malaysia based on an existing literature survey. In this catalogue Turner enumerated 8,198 species (Tab. 1). Parris & Latiff (1997) published a further update on the ferns and fern allies with some additions and nomenclatural changes to the group (Tab. 2). In this checklist, ferns and fern allies of Sabah and Sarawak were included to provide the first complete checklist of the group for Malaysia. For Sabah and Sarawak, no checklist exists but two important compilations were made for Rodriguésia 66(4):

21 952 Saw, L.G. & Chung, R.C.K. Table 1 Summary of the checklist of the flora of Peninsular Malaysia comparing Ridley s ( , 1926) enumeration and Turner s (1997) catalogue. Enumeration Groups Families Genera Species Ridley ( , 1926) Turner (1997) Ferns Gymnosperms Dicots 132 1,048 5,009 Monocots ,734 Total 182 1,493 7,183 Ferns & fern allies Gymnosperms Dicots 165 1,092 5,529 Monocots ,010 Total 248 1,651 8,198 Borneo (Merrill 1921; Masamune 1942, 1945). Masamune s compilations provided a more critical checklist and in that enumeration, 8,164 species of Bornean vascular plants were listed (Tab. 3). Other and more current accounts for the flora of Borneo were mostly foresters manuals and checklists often on selected groups in the region or states of Brunei, Kalimantan, Sabah and Sarawak (e.g. Anderson 1980; Argent et al. 1997; Ashton 1964, 1968, 1988; Browne 1955; Burgess 1966; Cockburn 1976, 1980; Hasan & Ashton 1964; Keith 1947; Kessler & Sidiyasa 1994; Newman et al. 1996; Primack 1983; Smythies 1965; Whitmore et al. 1990a, 1990b, 1990c; Wood & Agama 1956; Wood & Meijer 1964). The other checklists and revisions have been reviewed in the previous section. The launch of the Tree Flora of Sabah and Sarawak in 1991 was very significant and for the first time, used a modern systematic approach to enumerate the trees species (Soepadmo & Wong 1995). Apart from these enumerations, the other sources of information on the Flora of Malaysia are from the Flora Malesiana Series I & II for seed plants and ferns and other scattered publications. Based upon the information above, the flora of Peninsular Malaysia now stands over 8,300 species with recent updates from Turner (1997) (e.g., Turner 2000; Latiff & Turner 2001a, 2001b, 2001c, 2001d, 2002a, 2002b, 2003; Kamarudin & Turner 2004). This is probably an accurate estimate. For Sabah and Sarawak it is more difficult to arrive to an accurate figure with most estimates being for Borneo as a whole Merrill (1921) estimated about 9,000 species, Masamune (1942, 1945) enumerated about 8,200 species and more recently Wong (1995a) estimated a flora of between Merrill s 9,000 and 15,000 species. Kiew (1984) stressed the urgency for projects addressing the yet not studied and little understood Bornean flora. At present the Tree Flora of Sabah and Sarawak is the most important modern taxonomic project for Borneo. Since its inception in 1991, eight volumes have been published and provide an indication of the diversity of the Bornean flora. Table 4 provides a comparison of the tree flora of Sabah and Sarawak with the tree flora of Malaya comparing families and their enumeration. On average, the tree flora of Sabah and Sarawak contain about 38.5% more species than the tree flora of Malaya. If this proportion is maintained for the rest of the tree flora, considering that the tree flora of Malaya counts with 2,830 species (Ng et al. 1990), it is estimated that the tree flora of Sabah and Sarawak will contain about 4,000 species. Based upon this estimation also, with about 8,300 species of vascular plants in Peninsular Malaysia, it is estimated that the flora of Sabah and Sarawak will contain about 11,500 species. In Table 4, there are 889 species common to both Sabah and Sarawak, and Peninsular Malaysia (i.e., 55.1% overlap with Peninsular Malaysia). Based upon this overlap and using the estimated ratios, the total tree flora of Malaysia is estimated to be around 5,200 species and the total flora of vascular plants of Malaysia around 15,200 species. Rodriguésia 66(4):

22 The flora of Malaysia projects Table 2 Ferns and fern allies checklist of Parris & Latiff (1997). Region Species Total Malay Peninsula 647 Sabah 750 Sarawak 615 Total 1,165* * Species distribution overlap between the regions; as a result the total is smaller. Herbaria, Collections and Specimens Specimens are essential for the documentation of the Flora of Malaysia. Today, the collection at the herbarium of the Forest Research Institute Malaysia (KEP) comprises about 400,000 specimens. The other large herbarium holdings include the Forest Research Centre at Sandakan (SAN) with 300,000 specimens and the Forest Research Centre at Kuching (SAR) with about 300,000 specimens (Tab. 5). Other important Malaysian collections are found at the herbaria at University of Malaya (KLU) and Universiti Kebangsaan Malaysia (UKMB). The herbarium at the Singapore Botanic Gardens (SING) is particularly important for the Peninsular Malaysian flora. Many type specimens for plants described from Peninsular Malaysia are found amongst its 750,000 specimens (Tab. 5). Other important collections for the Malaysian flora include Brunei Forest Department (BRUN), Herbarium Bogoriense (BO), the Forest Herbarium (BKF), Bangkok, Thailand, National Herbarium of Netherlands, Leiden (L), Royal Botanic Gardens, Kew (K), Royal Botanic Garden, Edinburgh (E), UK, Arnold Arboretum (A), Harvard University, USA, and Central National Herbarium (CAL), Calcutta, India. For Sabah and Sarawak, O. Beccari s collection in the Herbarium Beccarianum (FI-B), Florence, Italy is particularly important. State of Knowledge for a Flora of Malaysia Among the key resources for speeding up the documentation of a flora of Malaysia is the availability of recent revisions that provide the foundation for flora writing. The vascular flora of Malaysia will include 250 families in Peninsular Malaysia and 253 families in Sabah and Sarawak. In working towards the flora of Malaysia, we have continued to separate the two regions, simply out of convenience from the historical perspective, but 953 also as there is a general tendency in many revisions to maintain the two regions as separate. We have compiled a review of published revisions against the families such as those included in the Flora Malesiana series, Tree Flora of Malaya, the revised Flora of Malaya, Tree Flora of Sabah and Sarawak, and other journal articles or series. In the analysis, out of the 250 families occurring in Peninsular Malaysia, 207 families (83%) have revisions. For Sabah and Sarawak, the coverage is much lower with 164 families out of 253 (64%) having revisions. In recent years, world checklists are being generated and these are being made available on the internet, for example, the checklists available from Royal Botanic Gardens, Kew (< wcb/>). Such checklists provide a useful preliminary starting point for flora accounts, especially for Borneo where existing list by Masamune (1942, 1945) have long become obsolete. Towards a Flora of Malaysia In the last two decades, Malaysia has been very fortunate in terms of the resources available to document its floristic diversity. The Tree Flora of Malaya published its final volume in 1989 (Ng 1989), but it was clear that the botanical work of documenting the flora of Malaysia needed to continue. It was an obvious decision to extend the well tested formula of the Tree Flora of Malaya and to extend it to Sabah and Sarawak. L.G. Saw, the first author of this paper, was asked to prepare proposals for funding towards a Tree Flora of Sabah and Sarawak project. The project was launched in 1991 with five years funding from the Malaysian Government, the Overseas Development Administration (ODA) of the United Kingdom and the International Tropical Timber Organisation (ITTO). It was originally estimated that the project would run for ten years, cover about 3,000 species (Soepadmo 1995) and be published in eight volumes. Having already completed eight volumes of the tree flora (Soepadmo & Saw 2000; Soepadmo & Wong 1995; Soepadmo et al. 1996, 2002, 2004, 2007, 2011, 2014) and with a current estimate of about 4,000 species of trees, we now estimate the Tree Flora of Sabah and Sarawak will need another ten years to complete the remaining estimated 2,000 species at a revision rate of about 200 species per year using present resources. In realising the flora of Malaysia, a pragmatic approach is to review our existing commitment towards the Tree Flora of Sabah and Sarawak Rodriguésia 66(4):

23 954 Saw, L.G. & Chung, R.C.K. Table 3 The flora of Borneo based on Masamune s checklist (Masamune 1942). Checklists Groups Families Genera Species Masamune (1945) Ferns & fern allies Masamune (1942) Gymnosperms Dicots ,997 Monocots ,170 Total 167 1,428 8,164 Table 4 Revisions of families of the Tree Flora of Sabah and Sarawak compared to the Tree Flora of Malaya. The figures for Tree Flora of Sabah and Sarawak were extracted from volumes 1-8 including 2 single-species families not found in the Tree Flora of Malaya; the figures for the Tree Flora of Malaya were extracted from volumes 1-4 with updates from Turner (1997). TFSS Volumes Tree Flora of Sabah & Sarawak Tree Flora of Malaya Species common to both regions Families Genera Species Families Genera Species Total , Table 5 Important herbarium holdings for Malaysia and Singapore. Country Herbaria Specimens Malaysia Forest Research Institute Malaysia 400,000 Forest Research Centre, Sandakan, Sabah 300,000 Forest Research Centre, Kuching, Sarawak 300,000 University Malaya 65,000 Universiti Kebangsaan Malaysia 72,000 Singapore Singapore Botanic Gardens 750,000 project and attempt to extend it into a full national flora project. The institutions currently engaged in the Tree Flora of Sabah and Sarawak, i.e. the Forest Research Institute Malaysia, and the Forest Departments of Sabah and Sarawak are keen to complete the Tree Flora project and it is clear from the information above that the Flora of Malaysia is best completed in a phased approach. In this pragmatic approach, the flora of Malaysia can be tackled as two regional projects, revisions for Peninsular Malaysia and revisions for Sabah and Sarawak. This is the way forward towards plans to complete the Flora of Malaysia. In April 2004, the Ministry of Natural Resources and Rodriguésia 66(4):

24 The flora of Malaysia projects Environment was formed. With the creation of the ministry, it became a national priority supported by government to document the biodiversity of the country. The work of documenting the flora of Malaysia quickly became a national need and no more an academic wish-list of botanists in Malaysia. Quickly identified was the need for a checklist of its entire flora as Peninsular Malaysia already has a checklist; the immediate need was for Sabah and Sarawak to have an updated list. In 2005, an application to fund a Flora of Peninsular Malaysia project was made. It was thought that the time was right for such a project as the Tree Flora of Sabah and Sarawak had already been running well for about 15 years and over that period the flora of Peninsular Malaysia had been relatively neglected. Furthermore, as explained earlier, a phased approach to realise the flora of Malaysia seemed a viable option for Malaysia. Subsequently the Flora of Peninsular Malaysia project received five years of funding at the end of The project has since published six volumes in two series - Ferns & Lycophytes (Parris et al. 2010, 2013) and Seed Plants (Kiew et al. 2010, 2011, 2012, 2013). For Sabah and Sarawak, the next phase (after the completion of the Tree Flora of Sabah and Sarawak project) will be to start the Flora of Sabah and Sarawak project so that all vascular plants will have been covered. Funding opportunities will need to be sourced in the future. Collaborations, Contributors and Rates of Revision Flora projects are always collaborative involving both local and foreign experts. The experiences from the Tree Flora of Malaya and Tree Flora of Sabah and Sarawak and the Flora of Peninsular Malaysia projects have shown that contributions from experts are essential for their success. Experts often produce revisions at a much faster pace and can tackle groups that are systematically more difficult. At the same time, Malaysian botanists must be trained to become experts so that they will continue to work within the country. Such strategy must be used for a Flora of Malaysia. Currently, the Tree Flora of Sabah and Sarawak and the Flora of Peninsular Malaysia are using such a strategy. Collaborations are at many different levels. At institutional level our traditional partners include local institutes such as Forest Research Centre, Sandakan, Forest Research Centre, Kuching, University 955 of Malaya, Universiti Kebangsaan Malaysia, Universiti Malaysia Sarawak and Universiti Malaysia Sabah. At regional level we collaborate with many herbaria including Singapore Botanic Gardens, Herbarium Bogoriense and the Royal Forest Herbarium, Bangkok. Internationally collaboration includes, the Royal Botanic Gardens Kew, Royal Botanic Garden Edinburgh, the Natural History Museum, London, the National Herbarium of the Netherlands, Leiden, and the Arnold Arboretum, Harvard University, USA. These collaborating institutes are important as they support our research by providing herbarium specimen loans, sourcing of literature, and a wide range of expert advice and support. At present we have over 25 collaborators actively contributing to revisions of families. To develop and build local expertise two essential elements must be in place; opportunities to build careers in botanical sciences and availability of training regimes for those interested. The Flora of Peninsular Malaysia project when it was first envisage included these elements. We are also very fortunate that, in the last few years, the Forest Research Institute Malaysia has committed to increase the number of botanists to do floristic work. In the last ten years, FRIM has recruited seven new botanists and taken on seven contract researchers for the two projects. Together with existing staff, FRIM now has 16 botanists working on both projects. Training of these new and aspiring botanists became a very important element of both projects. We are confident that, if the current institutional and financial support is maintained, both the Tree Flora of Sabah and Sarawak and the Flora of Peninsular Malaysia projects will be successful and will produce not just the revisions that contribute towards a Flora of Malaysia but also ensure that Malaysia will maintain a pool of botanists trained in understanding the its National flora. How many years will it take for the current flora projects to complete? It is very important that in planning the delivery of a Flora of Malaysia, we have a realistic estimate of manpower and financial cost. The Tree Flora of Malaya took 24 years to complete. Kiew (1984) made some projections regarding the rate of revision production by botanists in different types of floras (identification and information floras) in past flora projects. They ranged from 250 species per taxonomist per year to per year. Taking the Rodriguésia 66(4):

25 956 Saw, L.G. & Chung, R.C.K. example of the late K.M. Kochummen, in his seven years tenure as a full-time experienced botanist, he managed to revise 54 species per year for the Tree Flora of Sabah and Sarawak (Tab. 6). For the Flora of Peninsular Malaysia, Table 7 provides the different rates of revision against the number of full-time staff working on the flora revisions. The matrix estimates the number of years needed to complete the flora of Peninsular Malaysia with the estimated flora of 8,300 species. Using the example of Kochummen, we estimated that for a budding botanist, it would be very difficult to maintain a revision of over 50 species per year. A more realistic figure of about 40 species may be feasible for a relatively grounded botanist. If our current manpower strength is maintained with about 10 fulltime botanists working for the project, we envisage that it will take just over twenty years to complete the Flora of Peninsular Malaysia. This estimate ignores the contributions from other collaborators. For the Tree Flora of Sabah and Sarawak we have also worked out the rates using similar formulas (Tab. 8). The project with 5 full-time staff will take over twelve years to complete. Table 6 Families revised by K.M. Kochummen ( March 1999) during his tenure with the Tree Flora of Sabah and Sarawak project, his average revision rate was 54 species per year. Families Genera Species Anacardiaceae Burseraceae 8 59 Celestraceae Moraceae Ochnaceae 5 7 Total In our experience, training young botanists is a challenging task. Over the last twenty three years for both the flora projects we have directly trained twenty three individuals in the science of botanical revisions. We consider out of this number only seven have enough experience to independently revise families. Another four still require guidance in their work. The remaining have since left the project to take up other careers not related to botany. Finances and Institutional Commitment One of the constant challenges in any flora project is to ensure long-term commitment and sustainability in funding for the continuity of project. Projects are financed in fixed time-frame (e.g., 5 years for the Flora of Peninsular Malaysia), following which it is often difficult to obtain extensions. The Tree Flora of Sabah and Sarawak project has gone through a number of funding changes over the last 15 years; ODA, ITTO, Government of Malaysia (Intensification of Research and Development Priority Areas (IRPA) funding); 2006-present - research development funds from the Ninth and Tenth Malaysian Plans. Similarly, we expect the Flora of Peninsular Malaysia to go through various funding challenges as the project develops. In the future, we are not certain of how we can continue but it is up to the project to develop different ways to maintain funding continuity. We are now working towards funding for the two flora projects as a consolidated project under the Eleventh Malaysian Plan. Whoever the funding comes from it is important such projects have strong institutional commitment, failing which it would be almost impossible to secure continuity in finances and manpower commitment. It is essential that funding bodies see quality products arising from the project. There is a need to Table 7 Rate of revision based on about 8,300 species of vascular for the Flora of Peninsular Malaysia. Table 8 Revision rates based on about 2,500 species of tree species for the Tree Flora of Sabah and Sarawak. Number of Full-time Staff Number of Full-time Staff Revision Rates/ Staff/ Year Revision Rates/ Staff/ Year Rodriguésia 66(4):

26 The flora of Malaysia projects be creative in selling the products from the project, outside the standard flora volumes and more innovative methods must be used to disseminate the results of the projects to a wider audience so that they can be seen as pertinent and relevant to both national and scientific needs. Conclusion The Tree Flora of Sabah and Sarawak has produced eight volumes to date, FRIM is currently making information from the project available on the internet thus disseminating the results of the project to the wider public. The Flora of Peninsular Malaysia is being implemented together with a conservation project of threatened plants of Peninsular Malaysia, thus extending the taxon information with distribution to be used in conservation. Here too, the project has been productive and has now generated six volumes in two series. We are now in the process of including all revisions available on the web. This will be launched in the near future using software BRAHMS (Botanical Research And Herbarium Management System) online software. Such achievements will certainly complement the global effort towards the World Flora Online project. Based upon the discussion above, the phased approach towards a Flora of Malaysia the following points are reiterated. The inventory for a Flora of Malaysia can be done with resources in Malaysia and collaboration with our traditional partners; Based on current Tree Flora of Sabah and Sarawak and the Flora of Peninsular Malaysia projects, the Flora of Malaysia to continue with the geographical division of Peninsular Malaysia and Sabah & Sarawak; The project to be phased into the immediate short-term needs (checklists) and revisions of the two geographical floras; and Project must be seen as long-term as it requires long-term institutional and financial commitments in order to be brought into completion. References Anderson, J.A.R A Checklist of the trees of Sarawak. Forest Department, Sarawak. 364p. Argent, G.; Campbell, E.J.F. & Wilkie, P. (eds.) Manual of the larger and more important nondipterocarp trees of Central Kalimantan, Indonesia. Vol. 1 & 2. Forest Research Institute, Samarinda. 685p. 957 Argent, G.; Lamb, A. & Phillipps, A The Rhododendron of Sabah Malaysian Borneo. Natural History Publications (Borneo), Malaysia. Kota Kinabalu, Sabah. 290p. Ashton, P.S Manual of dipterocarp trees of Brunei State. Oxford University Press, London. 242p. Ashton, P.S A manual of the dipterocarp trees of Brunei State and of Sarawak. Borneo Literature Bureau for Sarawak Forest Department, Kuching. 129p. Ashton, P.S Manual of the non-dipterocarp trees of Sarawak. Vol. 2. Dewan Bahasa dan Pustaka for Forest Department Sarawak, Kuching. 490p. Beaman, J.H. & Beaman, R.S The plants of Mount Kinabalu 3. Gymnosperm and Non-Orchid Monocotyledons. Natural History Publications (Borneo) in association with Royal Botanic Gardens, Kew. 220p. Beaman, J.H.; Anderson, C. & Beaman, R.S The plants of Mount Kinabalu 4. Dicotyledon Families Acanthaceae to Lythraceae. Natural History Publications (Borneo) in association with Royal Botanic Gardens, Kew. 570p. Beaman, J.H. & Anderson, C The plants of Mount Kinabalu 5. Dicotyledon Families Magnoliaceae to Winteraceae. Natural History Publications (Borneo) in association with Royal Botanic Gardens, Kew. 609p. Beaman, J.H. & Beaman, R.S The plants of Mount Kinabalu 3. Gymnosperm and Non-Orchid Monocotyledons. Natural History Publications (Borneo) in association with Royal Botanic Gardens, Kew. 220p. Beaman, T.E.; Wood, J.J.; Beaman, R.S. & Beaman, J.H Orchids of Borneo. Natural History Publications (Borneo) in association with Royal Botanic Gardens, Kew. 584p. Browne, F.G Forest trees of Sarawak and Brunei. Government Press, Sarawak. 369p. Burgess, P.F Timbers of Sabah. Sabah Forest Records N o 6. Forest Department, Sabah. 511p. Burkill, I.H Botanical collectors, collections and collecting places in the Malay Peninsula. The Gardens Bulletin, Straits Settlements 4: Burkill, I.H A dictionary of the economic products of the Malay Peninsula. Vols. 1 & 2. Governments of the Straits Settlements and Federated Malay States, London. Volume I (A-H) p and Volume II (I- Z) p. Burn-Murdoch, A.M Trees and timbers of the Malay Peninsula, Part 1. F.M.S. Government Press, Kuala Lumpur. 14p + 13pl. Burn-Murdoch, A.M Trees and timbers of the Malay Peninsula, Part 2. F.M.S. Government Press, Kuala Lumpur. 17p + 6pl. Clarke, C.C Nepenthes of Borneo. Natural History Publications, Kota Kinabalu and Science and Technology Unit, Sabah. 209p. Rodriguésia 66(4):

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28 The flora of Malaysia projects Latiff, A. & Turner, I.M. 2001c. Quarterly notes: new taxa and records of Malaysian vascular plants. Folia malaysiana 2: Latiff, A. & Turner, I.M. 2001d. Quarterly notes: new taxa and records of Malaysian vascular plants. Folia malaysiana 2: Latiff, A. & Turner, I.M. 2002a. Quarterly notes: new taxa and records of Malaysian vascular plants. Folia malaysiana 3: Latiff, A. & Turner, I.M. 2002b. Quarterly notes: new taxa and records of Malaysian vascular plants. Folia malaysiana 4: Latiff, A. & Turner, I.M Quarterly notes: new taxa and records of Malaysian vascular plants. Folia malaysiana 4: Masamune, G Enumeratio Phanerogamarum Bornearum. Taihoku Imperial University, Japan. 739p. Masamune, G Enumeratio Pteridophytarum Bornearum. Imperial University, Japan 124p. Merrill, E.D A bibliographic enumeration of Bornean plants. Journal of Straits Branch Royal Asiatic Society, Special Number. 637p. Newman, M.F.; Burgess, P.F. & Whitmore, T.C Manuals of dipterocarps for foresters: Borneo Island Light Hardwoods. Royal Botanic Gardens Edinburgh & CIFOR. Bogor. 275p. Ng, F.S.P. (ed.) Tree flora of Malaya. Malayan Forest Records 26. Vol. 3. Forest Department, West Malaysia. 339p. Ng, F.S.P. & Jacobs, M A guide to King s Materials for a flora of the Malayan Peninsula. The Gardens Bulletin Singapore 36: Ng, F.S.P. (ed.) Tree flora of Malaya. Malayan Forest Records 26. Vol. 4. Forest Department, West Malaysia. 549p. Ng, F.S.P.; Low, C.M. & Mat Asri, N.S Endemic trees of the Malay Peninsula. Research Pamphlet 106. Forest Research Institute Malaysia, Kuala Lumpur. 118p. Parris, B.S.; Beaman, R.S. & Beaman, J.H The Plants of Mount Kinabalu 2. Ferns. Royal Botanic Gardens, Kew. 165p. Parris, B.S. & Latiff, A Towards a Pteridophyte flora of Malaysia: A provisional checklist of taxa. Malayan Nature Journal 50: Parris, B.S.; Kiew, R.; Chung, R.C.K.; Saw, L.G. & Soepadmo, E. (eds.) Flora of Peninsular Malaysia. Series I: Ferns and Lycophytes, Vol. 1. Malayan Forest Records N o 48. Forest Research Institute Malaysia, Kepong, Selangor. 249p. Parris, B.S.; Kiew, R.; Chung, R.C.K. & Saw, L.G. (eds.) Flora of Peninsular Malaysia Series I: Ferns and Lycophytes, Vol. 2. Malayan Forest Records N o 48. Forest Research Institute Malaysia, Kepong, Selangor. 241p. Piggott, A.G Ferns of Malaya in colour. Tropical Press Sdn. Bhd. Kuala Lumpur, Malaysia. 458p. 959 Poulsen, A.D Etlingera of Borneo. Natural History Publications (Borneo) in association with Royal Botanic Garden Edinburgh, Kota Kinabalu. 263p. Primack, R.B Forester s guide to the Moraceae of Sarawak. Forest Department Sarawak, Kuching. 140p. Ridley, H.N Materials for a Flora of the Malayan Peninsula. 3 Parts. Methodist Publishing House, Singapore. Part 1, 233p.; Part 2, 235p.; Part 3, 197p. Ridley, H.N The flora of the Malay Peninsula. Vol L. Reeve & Co. Ltd, London. 1: 918, 2: 672, 3: 405, 4: 383, 5: 470. Ridley, H.N The ferns of the Malay Peninsula. Journal of the Malayan Branch, Royal Asiatic Society 4: Saw, L.G. & Chung, R.C.K Towards the flora of Malaysia. In: Chua, L.S.L.; Kirton, L.G. & Saw, L.G. (eds.). Proceedings of seminar and workshop on the status of biological diversity in Malaysia & threat assessment of plant species in Malaysia. Forest Research Institute Malaysia, Kepong. Pp Seidenfaden, G. & Wood, J.J The orchids of Peninsular Malaysia and Singapore. Olsen & Olsen, Fredensborg. 779p. Smythies, B.E Common Sarawak trees. Borneo Literature Bureau, Sarawak. 153p. Soepadmo, E Background to the Tree Flora of Sabah and Sarawak Project. In: Soepadmo, E. & Wong, K.M. (eds.). Tree Flora of Sabah and Sarawak. Vol. 1. Forest Research Institute Malaysia, Kuala Lumpur, Sabah Forestry Department, Sandakan & Sarawak Forestry Department, Kuching. Pp. XIII-XIX. Soepadmo, E. & Wong, K.M. (eds.) Tree flora of Sabah and Sarawak. Vol. 1. Forest Research Institute Malaysia, Kuala Lumpur, Sabah Forestry Department, Sandakan & Sarawak Forestry Department, Kuching. 513p. Soepadmo, E Botanical study of Malaysian medicinal plants an appraisal. In: Manaf Ali, A. et al. (eds.). Phytochemical and biopharmaceutins from the Malaysian rain forest. FRIM & PSSM, Kuala Lumpur. Pp Soepadmo, E.; Wong, K.M. & Saw, L.G. (eds.) Tree flora of Sabah and Sarawak. Vol. 2. Forest Research Institute Malaysia, Kuala Lumpur, Sabah Forestry Department, Sandakan & Sarawak Forestry Department, Kuching. 443p. Soepadmo, E. & Saw, L.G. (eds.) Tree flora of Sabah and Sarawak. Vol. 3. Forest Research Institute Malaysia, Kuala Lumpur, Sabah Forestry Department, Sandakan & Sarawak Forestry Department, Kuching. 511p. Soepadmo, E.; Saw, L.G. & Chung, R.C.K. (eds.) Tree Flora of Sabah and Sarawak, Vol. 4. Forest Research Institute Malaysia, Kuala Lumpur, Sabah Forestry Department, Sandakan & Sarawak Forestry Department, Kuching. 388p. Rodriguésia 66(4):

29 960 Saw, L.G. & Chung, R.C.K. Soepadmo, E.; Saw, L.G. & Chung, R.C.K. (eds.) Tree Flora of Sabah and Sarawak. Vol. 5. Forest Research Institute Malaysia, Kuala Lumpur, Sabah Forestry Department, Sandakan & Sarawak Forestry Department, Kuching. 528p. Soepadmo, E.; Saw, L.G. & Chung, R.C.K. (eds.) Tree Flora of Sabah and Sarawak. Vol. 6. Forest Research Institute Malaysia, Kepong. 335p.Soepadmo, E.; Saw, L.G.; Chung, R.C.K. & Kiew, R. (eds.) Tree Flora of Sabah and Sarawak. Vol. 7. Forest Research Institute Malaysia, Kepong. 450p. Soepadmo, E.; Saw, L.G.; Chung, R.C.K. & Kiew, R. (eds.) Tree Flora of Sabah and Sarawak. Vol. 8. Forest Research Institute Malaysia, Kepong. 248p. Symington, C.F Foresters manual of Dipterocarps. Malayan Forest Records N o 16. Syonan-Hakubutukan, Kuala Lumpur. 244p. Symington, C.F (reprint). Foresters manual of Dipterocarps. Malayan Forest Records N o 16. Penerbit Universiti Malaya, Kuala Lumpur. 244p. + plate Thiers, B Index Herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden s Virtual Herbarium. Available at < Access on 31 July Turner, I.M ( 1995 ). A catalogue of the vascular plants of Malaya. The Gardens Bulletin Singapore 47: Turner, I.M The taxonomy of Malaysian vascular plants: new taxa ( ) and endemic genera. Folia malaysiana 2: Van Steenis, G.G.G.J Annotated selected bibliography. Flora Malesiana Series I. Vol. 5. Noordhoff-Kolff N.V., Djakarta. 595p. Van Steenis-Kruseman, M.J Cyclopaedia of collectors. Flora Malesiana Series I. Vol. 1. Noordhoff- Kolff N.V., Djakarta. 639p. Whitmore, T.C. (ed.) Tree flora of Malaya. Malayan Forest Records 26. Vol. 1. Forest Department, West Malaysia. 471p. Whitmore, T.C. (ed.) Tree flora of Malaya. Malayan Forest Records 26. Vol. 2. Forest Department, West Malaysia. 444p. Whitmore, T.C.; Tantra, I.G.M. & Sutisna, U. 1990a. Tree flora of Indonesia. Check list of Kalimantan. Part I. Forest Research and Development Centre, Bogor. Pp Whitmore, T.C.; I.G.M. & Sutisna, U. 1990b. Tree flora of Indonesia. Check list of Kalimantan. Part II. 1. Forest Research and Development Centre, Bogor. Pp Whitmore, T.C.; I.G.M. & Sutisna, U. 1990c. Tree flora of Indonesia. Check list of Kalimantan. Part II. 2. Forest Research and Development Centre, Bogor. Pp Wong, K.M The herbarium and arboretum of the Forest Research Institute of Malaysia at Kepong - a historical perspective. The Gardens Bulletin Singapore 40: Wong, K.M. 1995a. A brief history of botanical collecting and documentation in Borneo. P. XXI-XLI In: Soepadmo, E. & Wong, K.M. (eds.). Tree flora of Sabah and Sarawak. Vol. 1. Forest Research Institute Malaysia, Kuala Lumpur, Sabah Forestry Department, Sandakan & Sarawak Forestry Department, Kuching. 513p. Wong, K.M. 1995b. The Bamboos of Peninsular Malaysia. Malayan Forest Records N o 41. Forest Research Institute Malaysia, Kepong. 199p. Wong, K.M Patterns of plant endemism and rarity in Borneo and the Malay Peninsula. In: Peng, C.-I. & Lowry II, P.P. (eds.). Rare, threatened, and endangered floras of Asia and the Pacific Rim. Academia Sinica Monograph Series N o 16: Wood, G.H.S. & Agama, J Check list of forest flora of North Borneo. North Borneo Forest Records N o 6. Sunshine Print, Sandakan. 42p Wood, G.H.S. & Meijer, W Dipterocarps of Sabah (North Borneo). Sabah Forest Records N o 5. Forest Department, Sandakan. 344p. Wood, J.J.; Beaman, R.S. & Beaman, J.H The plants of Mount Kinabalu 2. Orchids. Royal Botanic Gardens, Kew. 411p. Wood, J.J. & Cribb, P.J A checklist of orchids of Borneo. Royal Botanic Gardens, Kew. 421p. Wyatt-Smith, J Pocket check list of timber trees. Malayan Forest Records N o 17. Forest Department, Kuala Lumpur. 367p. Wyatt-Smith, J. 1953a. Manual of Malayan timber trees. Burseraceae. F.R.I. Research Pamphlet N o 1. Forest Research Institute, Kepong. 33p. Wyatt-Smith, J. 1953b. Manual of Malayan timber trees. Leguminosae. F.R.I. Research Pamphlet N o 2. Forest Research Institute, Kepong. 74p. Wyatt-Smith, J. 1953c. Manual of Malayan timber trees. Myristicaceae. F.R.I. Research Pamphlet N o 3. Forest Research Institute, Kepong. 25p. Wyatt-Smith, J. 1954a. Manual of Malayan timber trees. Sapotaceae. F.R.I. Research Pamphlet N o 4. Forest Research Institute, Kepong. 56p. Wyatt-Smith, J. 1954b. Manual of Malayan timber trees. Lauraceae. F.R.I. Research Pamphlet No. 5. Forest Research Institute, Kepong. 64p. Wyatt-Smith, J. 1954c. Manual of Malayan timber trees. Sapindaceae. F.R.I. Research Pamphlet No. 6. Forest Research Institute, Kepong. 23p. Artigo recebido em 11/06/2015. Aceito para publicação em 03/09/2015. Rodriguésia 66(4):

30 Rodriguésia 66(4): DOI: / The World Flora Online 2020 project: will Cameroon come up to the expectation? Jean Michel Onana 1 Abstract Biodiverse Cameroon has been highlighted as the top country in tropical Africa for plant species diversity per degree square, with a higher diversity than all other West African countries added together, and including two of the top documented centres of plant diversity in Tropical Africa. Despite its reduced taxonomic capacity, with only six active taxonomists a high level of botanical activity in the country has resulted in accomplishments such as the databasing of the YA Herbarium (over 60,000 records), which has an in-country collection coverage of almost 95% of the known plant species that are recorded for Cameroon. Other accomplishments are the Red Data Book of the Flowering Plants of Cameroon, several local checklists and published volumes of the Flore du Cameroun which covers 37% of the country s species. Currently the checklist of Cameroon records 7,850 taxa at species and infraspecific level. Resources are needed to support and heighten the profile of this small botanical community. Already thanks to strong collaboration between Cameroon and renowned botanical institutes of others countries, in particular France and United Kingdom, one hundred and thirteen plant families have been published and would help this country to complete the recording of its biodiversity towards contributing to the World Flora Online 2020 project. Key words: Africa, botanical research, checklist, flora, Republic of Cameroon. Resumo A República dos Camarões tem sido destacado como o principal país na África tropical para a diversidade de espécies de plantas por grau quadrado, com uma diversidade maior do que todos os outros países do Oeste Africano juntos, e incluindo dois dos maiores centros de diversidade de plantas documentadas na África Tropical. Apesar da sua reduzida capacidade taxonômica, com apenas seis taxonomistas ativos, um elevado nível de atividade botânica no país resultou na digitalizaçãodo herbário YA (com mais de registros), tendo em seu acervo quase que 95% das espécies de plantas conhecidas para os Camarões. Outras conquistas para a botânica do país são o Livro Vermelho das Angiospermas dos Camarões, várias listas de verificação locais e volumes publicados da Flore du Cameroun, que abrange 37% das espécies do país. Atualmente, a lista dos Camarões apresenta 7,850 táxons registrados tanto em nível específico como infraspecífico. Porém mais recursos financeiros são necessários para apoiar e elevar o perfil desta pequena comunidade botânica no país. Graças a uma forte colaboração entre os Camarões e renomados institutos de botânica de outros países, em especial da França e do Reino Unido, os tratamentos para cento e treze famílias de plantas já foram publicados e representam um importante passo para que este país possa completar o registro da sua biodiversidade, contribuindo assim para o projeto da Flora Mundial on-line Palavras-chave: África, pesquisa botânica, República dos Camarões. Introduction As a Party of the Conference of Parties, Cameroon has accepted the duty to implement decision X/17 of the consolidated update of the Global Strategy for Plant Conservation for a World Flora online by 2020 (UNEP/CBD/SBSTTA/16/ INF/38, 23 rd April 2012). However, the 5 th national report for the Convention of Biodiversity - CBD (République du Cameroun 2014) did not make any mention of this global project. This does not mean that there is no intention to continue with the efforts to complete the flora of Cameroon. According to Letouzey (1968), floristic research in Cameroon began in earnest when Gustav Mann investigated Mount Cameroon for the first 1 National Herbarium of Cameroon, PO. Box 1601, Yaoundé, Cameroon and Department of Plant Biology, Faculty of Sciences, University of Yaoundé I, PO. Box 812 Yaoundé, Cameroon. jmonana2002@yahoo.fr

31 962 Onana, J.M. time in December Later, several botanists travelled to Cameroon and collected material which was deposited in European Herbaria, particularly B, K, and P (Holmgren et al., 1990). The creation of the Herbarium in Yaoundé dates from 1948; initially specializing in forest trees as part of the Forest Research Section of Cameroon (SRFK (Kamerun) or SRFCam). This facility was later expanded to include specimens of all vascular plants and, in 1971, it became the National Herbarium of Cameroon (YA). The project Etude de la Flore du Cameroun has been running since 1950 with three main objectives: set up a collection of specimens of all vascular plant species of Cameroon; establish a map of the main phytochoria; and describe the material family by family in the series Flore du Cameroun (FDC). While much additional work has been carried out through different studies and projects recording and describing taxa from Cameroon, the work in the Cameroonian flora continues to move forward as our plant species become better known. Emerging patterns from our rich flora cause the taxonomists in Cameroon to ask the following questions: What is the present situation of the flora of Cameroon? Is there a working species list? How is the species list maintained and updated? What is the methodology that Cameroon will follow to complete its flora by 2020? How is the local scientific community going to be coordinated and stimulated towards improving the quality of the knowledge regarding Cameroon s flora? What are the biggest challenges and obstacles envisaged in order to complete Cameroon s flora by 2020? How do we envisage that the data/information from the Cameroon flora will be inserted into the World Flora 2020 project? What is the importance of the World flora project for Cameroon? Before answering these questions, an overview of the location and biophysical features of the country, together with the current situation of the plant collections, collection coverage, floristic richnes estimates, state of floristic publications and an update on Cameroonian threatened plants will be presented. Methodology The Republic of Cameroon, covering an area of 475,442 km 2, is situated in west-central Africa (Fig. 1) between latitude 1 and 13 N and longitudes 8 and 17 E, extending from the Atlantic coastline in the south to lake Chad in the extreme north, in the Sahel at the edge of the Sahara Desert. Four major biophysical features dominate the country Figure 1 Cameroon in Africa. Maps of Cameroon date 31 June From Wikimedia Commons. Author: MS Claudia. (< org/wiki/file:cameroon_(orthographic_projection). svg.>). Dowloaded 27/05/2015. in relation to the relief and geomorphology (Morin 1979; Tchawa 2006), climate (Moby 1979; Tsaléfac 2006), including the majority of all known African vegetation types (Letouzey 1979, 1985; Onana 2013), originating the expression Africa in miniature (Fig. 2). These are (from south to north): 1. The south Cameroon plateau At m altitude, occupies about a third of the country. The rainfall is generally around 1,500 mm per annum, with four seasons of unequal intensity divided in two wet seasons, a heavy rainy season from mid August to mid November and a light wet season from mid March to mid June; and two dry seasons, a more intense dry season from mid November to mid March and a weaker one from mid June to mid August. Semi-deciduous rain forest is the major vegetation type. 2. The coastal plain Is km wide, with altitude varying between sea level and 300 m, with rainfall around 4,000 mm (exceptionally high at Cape Debundscha at the foot of Mt Cameroon on the Atlantic Ocean side, one of the five wettest places in the world, Rodriguésia 66(4):

32 Cameroon in the World Flora 2020 where it can reach over 10,000 mm yearly). The climate in the coastal plains is known as type camerounien with two major seasons, a long rainy season and a short dry season lasting from December to February. Ombrophilous evergreen rainforest covers the whole area. 3. The Cameroon Highlands Are formed by an interrupted volcanic line from the sea to the extreme north of Cameroon in the Mandaras Mountains, with a curve on the Adamaoua plateau (1,000 1,200 m altitude). Cameroon contains Africa s highest mountains west of the Albertine Rift, namely Mount Cameroon (4,070 m), Mt Oku (3,011 m) and Mount Bamboutos (2,740 m). The highlands continue off shore towards the southwest into a line of islands, most notably Bioko Island (Equatorial Guinea), São Tomé Island, Príncipe Island (both Republic of São Tomé and Príncipe) and Pangalu or Annobon Island (Equatorial Guinea). The climate is a variant of the coastal plain climate, with increased Figure 2 Vegetation cover in Cameroon (source: Interactive Atlas of forestry resources of Cameroon, version 2.0. Cameroon Ministry of Forestry and Fauna/ World Resources Institute 2007) 963 rainfall caused by clouds being forced up from the southwest African monsoon winds in the period April-October, the main wet season. The vegetation comprises Submontane rain forest at around 1,000 to 2,000 m alt., Montane rainforest between 2,000 and 3,000 m alt. to Afrosubalpine grassland and prairies over 3,000 m alt. on two of the mountains: Mt Oku and Mt Cameroon. On the Adamaoua, the soudanian area of the Cameroon Mountains, the highest summits are Tchabal Mbabo (2,640 m altitude), Hosséré Vokré (2,049 m), Tchabal Nganha (1,923 m) and Tchabal Ngangdaba (1,960 m). Rainfall is lower, at around 1,500 mm per annum. Vegetation on the plateau is Open forest to Woody savanna. 4. The north plain plateau Altitude varies from 300 m to up to 1,442 m at the Mandaras Mountains, the northern portion of the Cameroonian mountains. The rainfall varies from 300 mm in the far north to 800 mm further south, with two seasons: a short rainy season from July-August to September-October and a long dry season from September-October to July-August. The tropical savanna vegetation includes Woody savanna to Herbaceous, open savanna and steppes in the extreme north. Results Collections Around 501 expeditions involving approximately 634 botanists were carried out in Cameroon from 1861 to the turn of this century. Since then, the collection effort has been project based, involving field work for taxonomic studies or biodiversity assessments for projects involving land development and designation of protected areas (Cheek et al. 2004, 2010, 2011). According to methodology suggested by Poncy & Labat (1994), Cameroon s density of plant collection (Fig. 3) referring to the Indices of Density of Collection (IDC) reached about 0.3 collection/km 2 (30 collections/100 km 2 ) in 2010 (Onana 2010). Overall, therefore, the floristics of Cameroon remains little known at the botanical level when considering that a minimum rate of 1 specimen/km 2 is the acceptable value for the inventory of a territory (Campbell & Hammond 1989). For Cameroon, Poncy & Labat (1994) calculated that, from a base of 10 collections/100 km 2 (0,1 collections/km 2 ) in 1981, with an increase of 1,38 collections/100 km 2 per annum in 7 years ( ), 65 years would be necessary to reach the target of 100 collections/100 km 2 (1 collection/ Rodriguésia 66(4):

33 964 Onana, J.M. Figure 3 Overview of botanical effort collection in Cameroon based on the distribution of 36,588 specimens at YA using half degree cells (source: Onana 2011). Figure 4 Overview of vascular plant species richness in Cameroon, based on the distribution of 36,588 specimens at YA using half degree cells (source: Onana 2011). km 2 ); implicating that the target would be reached in In 2015, we are still very far from reaching this target. By 2015, around 65,000 specimens were incorporated in the YA working collection, with duplicates of these deposited in various Herbaria (B, BM, FHI, G, K, P, MO, SCA, WAG and Z). Considering that material of only 376 species of the 7,850 known species is not currently deposited at YA, our National Herbarium has an excellent coverage of about 95% of the total number of species recorded for Cameroon. The database. The digitization of the label data has started in 2002 with the RIHA database on an Access platform with PostGrès (Chevillotte et al. 2006). The database is supported by the Institut de Recherche pour le Développement (IRD) and the Royal Botanic Gardens, Kew (RBG Kew). Since December 2010, all the 65,000 specimens of the working collection are included in the database. New collections are added regularly and updates continue for localities and scientific names. Floristic richness in Cameroon The floristic richness per experimental unit is unevenly distributed as it currently appears to follow the collection density (Fig. 4). According to Onana (2010b, 2011), the floristic richness in vascular plant species per taxa at the family, genus and species levels is as follows: 1. Families The number of families of vascular plants recorded in Cameroon varies according to the system followed: according to Brummitt (1992) and Flora of Cameroon vol. 3 there are 248 families (25 of Pteridophyta), of them 228 being native and 20 introduced families of seed plants. According to Heywood et al. (2007) there are 195 families of flowering plants; according to APG III (APG III, 2009; Chase & Reveal 2009) there are 182 families of flowering plants. For the non-flowering vascular plants, Roux (2009), in line with results of recent phylogenetic studies, lycophytes were separated from ferns recognizing 30 families (3 families for Lycopodiophyta and 27 of Pteridophyta). Using the latest phylogenetic studies (APG III) as a basis for our count, Cameroon has 212 families of vascular plants (Tracheophyta). All families known in Cameroon are represented in the collections of YA. 2. Genera According to Onana (2011) there are 1,884 plant genera in Cameroon (amongst them 87 non-flowering vascular plant genera) of which 1,641 are native and 243 are introduced. The only two genera not found at YA are Namatea (Sapindaceae) and Winklerella (Podostemaceae). Rodriguésia 66(4):

34 Cameroon in the World Flora Species From the 7,850 species reported for the Cameroonian territory 7,385 are native and 534 are introduced, of which 185 are naturalized while 349 are cultivated. Collection effort gaps need to be addressed in order to complete the records. According to Joppa et al. (2010), if the undescribed or yet unrecorded species comprise around 10 20% of those already described (Bebber et al. 2010, Küper et al. 2006; Onana 2011), the specific richness of vascular plant species can be estimated at between 8,700 and 9,500 species. Publications on the Cameroon flora The series Flore du Cameroun (FDC) comprises at the moment of fourty two published volumes. Three volumes not dedicated to taxonomic treatment of families present the history of the botanical collections in Cameroon (Letouzey 1968), a checklist of the vascular plant species of Cameroon (Onana 2011) and a checklist of endemic and rare plants of Cameroon (Onana 2013). In the thirty nine volumes others, installments by botanical family include descriptions of morphological characters, illustrations, keys for genera and species, geographical distribution and citation of studied material. The 39 volumes so far published by 38 taxonomists of c. 10 different countries of Europe and United States of America and one (Ntepe- Nyame) from Cameroon, comprise 112 families, 624 genera and 2,269 species. An addidional publication that dealt with 433 species of grasses (van der Zon, 1992) brings the total to 2,702 taxa (37% of the known taxa) published. The taxonomic checklist of vascular plants of Cameroon (Onana 2011) shows a total of 7,850 taxa (including subspecies and varieties). According to this figure, Cameroon is the fourth richest territory in vascular plant species in continental Africa, after South Africa (c. 23,400 species), Democratic Republic of Congo and Tanzania (c. 10,000 species each). The following families are currently being prepared for publication: Burseraceae, Gnetaceae, Lecythidaceae, Polygalaceae and Podocarpaceae,. This will increase the number of taxa published and improve the knowledge of the flora of Cameroon. At the moment genera of Annonaceae (Piptostigma), Orchidaceae (Angraecum) and Rubiaceae (Bertiera, Craterispermum, Psychotria, and Sabicea) are being studied by students developing doctoral thesis or as postdoctorate research work. 965 The bibliography dealing with the Cameroonian Flora also comprises around 600 papers published in peer-reviewed taxonomic journals describing new taxa or revising taxonomic groups in large families such as Rubiaceae (94 publications), Legumes (47), Acanthaceae (21), Annonaceae (18) and Euphorbiaceae (31). According to Polhill (1990 cited by Poncy & Labat 1994: 12) a taxonomist working midtime can complete 50 species descriptions in a year ; thus, in order to describe the known 5,500 species of the flora of Cameroon that remain to be described, from 2015, one taxonomist would be working for 116 years, while four botanists would take 29 years. This is surely slow progress and more should be done to train and employ taxonomists and to develop further collaborations worldwide to speed up the baseline study of the flora. In a more positive light, it is worth considering that the availability of digitized type as well as nontype specimens (Darbyshire et al. 2010) and the availability of taxonomic literature online have probably improved the productivity of scientists (Onana & Chevillotte 2015), indicating that the task may be concluded in less time (Costello et al. 2013). Local (vernacular) names Lists of vernacular names have been established for four national languages of the forest area, covering three administrative regions (Central, South and East - ewondo, bulu, baka or bibaya and bagueli languages) by Onana & Mezili (in prep.) and languages for the tropical savanna area of the North and Far North (fufulde, guizigua, hoho, madan, mafa, toupouri, massa, zoulgo) by Ntoupka (in prep.) The manuscripts for these lists remain unpublished, but the plan is to publish these data within the FDC series. Threatened flowering plant species of Cameroon. Studies were carried out to catalogue the threatened flowering plants species and their distribution and endemism. In the Red Data Book of flowering plant species of Cameroon (Onana & Cheek 2011), 815 flowering plant species are documented, including a map for each species evaluated as threatened using IUCN global assessments (IUCN 2001), most of them having been assessed for the first time. The limitations in the work are due to the impossibility to assess Rodriguésia 66(4):

35 966 Onana, J.M. families that lack a recent revision (Cheek in Onana & Cheek 2011). Since numerous species occur beyond Cameroon, the new assessments are also relevant to Nigeria, Gabon, and as far west as Guinea, Ivory Coast, to east as Congo, Rwanda and Burundi. The hotspots defined as areas of concentration of threatened species are shown on a map at the back cover (Fig. 5), where it can be seen that the Afromontane Archipelago Centre of endemism (White 1983) harbours the majority of threatened species, followed by the Lower Guinea Centre of endemism. The assessments are now in the process of validation via the Species Information Service (SIS) of IUCN. At the moment 100 of these assessments have been published in October 2014 on the IUCN website (< iucnredlist.org/>). Endemic and rare plant species The synopsis of endemic and rare species of vascular plant species of Cameroon (Onana 2013), where species represented by between one and ten collections are considered rare, documents 1,689 species (including infraspecific taxa), among them 585 known only in Cameroon and thus considered endemic sensu stricto. The synopsis records the habit, habitat, distribution, state of endemism and rarity and the IUCN (2001) status at the global level. Discussion Highlights regarding native species diversity in Cameroon are: Cameroon is believed to be the top country in tropical Africa for plant species diversity per degree square (Barthlott et al. 1996) with more than 5,000 species per degree square in parts of the southwest of the country; Cameroon alone has higher plant species diversity than the all the other countries in West Africa, from Senegal to Nigeria added (Cheek & Onana 2011); The top documented centres of plant diversity in Tropical Africa are Mount Cameroon (2,435 species) and Mt Kupe - Bakossi Mountains (2,412 species) (Cheek & Onana 2011) and Kribi-Campo area (2,297 species) (Tchouto 2004; Tchouto & Elad 2010). 1. Cameroon s working plant list The richness of the flora of Cameroon was evaluated by reference to the working collection in YA (Satabié 1999). The need for a list of accepted scientific names for the vascular plants of Cameroon is evident, because the information is contained in a large volume of taxonomic works, several recent regional floras, as well as electronic initiatives, such as the World Checklists of Selected Plant Families on-line (WCSP 2015) and the digitization of data in herbaria worldwide and at the YA Herbarium. The first species taxonomic checklist for Cameroon was published in 2011 (Onana 2011). The checklist comprises the groups of the vascular species: Angiosperms, Gymnosperms, Pteridophytes and Lycophytes. The taxonomic checklist comprises 7850 scientific names, followed by their IUCN status. The checklist serves as baseline for the Red data Book of flowering Plants of Cameroon (Onana & Cheek 2011). When crosschecked with the list of accepted names on-line (The Plant List 2013), it is found that 6,166 of these names are accepted, 936 are considered synonyms and the status of 748 names comprises unresolved names, names not currently found in The Plant List and different spellings. The necessary updates and corrections are being implemented. 2. How is the species list maintained and updated The Cameroon taxonomic checklist is supposed to be updated every five years. The methodology is to check regularly the taxonomic works in which material from Cameroon is cited. During the period April 2011 to August 2015, 36 new species have been recorded as follows: Afrothismia (2) ; Crateranthus (1), Cryptosepalum (1), Cyperus (1), Distylodon (1), Dracaena (1), Gilbertiodendron (4), Globulostylis (3), Gnetum (2); Hymenostegia (1); Kihansia (1), Leptonychia (3), Massularia (1), Memecylon (3), Napoleonaea (1), Neuropeltis (1), Polystachya (1), Psychotria (2), Rhipidoglossum (1), Sabicea (1) and Vepris (5). During the same period, few previously accepted in Globulostylis (1) and Vepris (1), have been synonymized. In total, about 34 names species can be added to the list, bringing the actual number of known species to 7,884. At present, it is important to push for publication of more revision of taxonomic group, including work regarding taxa already published in the FDC series, for example the work by Onana & Chevillotte (2015) complementing the volume on Rutaceae published in 1963; Stone (2015) Rodriguésia 66(4):

36 Cameroon in the World Flora Figure 5 Overview of hotspots of flowering plants in Cameroon (source: Onana & Cheek 2011, back cover ) Rodriguésia 66(4):

37 968 Onana, J.M. and Stone et al. (2008) for the Melastomataceae (volume published in 1983); van der Burgt (2010) for the Chrysobalanaceae (published in 1978); van der Burgt (2009), van der Burgt & Newbery (2006) for the Sapotaceae (published in 1964); and van der Burgt et al. (2012, 2015) for Leguminosae- Caesalpinioideae (published in 1970). New volumes of FDC dealing with families not yet treated, and the production of a more documented checklist including additional data such as habit, distribution and voucher specimens are all priorities to further the knowledge of Cameroonian plants. 3. Methodology to complete the Flora The World Flora 2020 project is not implemented in Cameroon as a project by itself. Floristic research is done in the framework of the project Etude de la Flore du Cameroun which aims to describe all the vascular plants of Cameroon and to publish in the FDC series. The project has no timetable and volumes are published as opportunities arise, on a voluntary basis, by taxonomists who are studying the flora of the country. 4. Coordinating and stimulating the local scientific community towards improving the quality of the knowledge regarding Cameroon s flora The Cameroonian scientific community for plant taxonomy is very small and the country lacks resources. Currently the local scientific community count only six active plant taxonomists among them three at YA Herbarium: Ghoghe Jean Paul (Annonaceae, Podostemaceae), Tchiengue (Guttiferae) and Onana Jean Michel (Burseraceae, Rutaceae); a further two at the University of Yaoundé I: Biye Elvira (Gnetaceae) and Sonké Bonaventure (Rubiaceae); and Sainge Moses (Thismiaceae) at the University of Buea. Floristic research in Cameroon is coordinated by the Head of the National Herbarium (YA) who is, in fact, the principal investigator of the project to study the country s flora. One of his duties is to stimulate the description of plants of Cameroon by exchange of material with foreign taxonomists who cannot travel to Cameroon, getting authorization of research and welcoming others for field work and helping students in the herbarium for their academic works. In addition, the taxonomists working as lecturers at the University of Yaoundé I describe species as part of their research, participating in the Flora of Cameroon project as associate researchers or fellows. 5. Biggest challenges and obstacles to completion of the Flora of Cameroon by 2020 In order to be able to complete the Flora of Cameroon by 2020, solutions for the following big obstacles need to be found: The lack of plant taxonomists (Surtcliffe & O Reilly 2010) is acute in Cameroon at present, with only four taxonomists regularly describing and publishing new taxa or revisions, and further two who have sporadically described new species in the five last years but are not working in floristic research. The lack of opportunity and stimulation for a carrier as plant taxonomist is the main reason why the three scientists who have completed their PhD in plant taxonomy (Kenfack 2008; Nguembou 2008; Taedoung 2014) left plant taxonomy for other fields such as forestry conservation. Taxonomic research is not amongst the country s priorities. In a developing country context where one of the big questions is food security, biodiversity research that leads to conservation is regarded as a possible obstacle for the development of agricultural activities, causing plant taxonomy research to be relegated to the lowest rank of priorities, with almost inexistent funding. It is an increasing challenge to convince taxonomists to agree to produce family accounts for Cameroon. There are no more taxonomists working on African tropical plants at the Muséum National d Histoire Naturelle in Paris (P Herbarium), the main partner for the production of floras in French. Other European and North American institutions are following the same direction, giving preference to other types of research such as molecular biology, as these papers can be published in journals with higher impact. The result is that they neglect the need to produce baseline data to address the knowledge of floras at local, country and regional levels. The lack of funding allocated to research and development also impacts upon the maintenance of collections, laboratories and general working conditions for taxonomists. 6. Data and information provision from Cameroon flora to the World Flora project Amongst the avenues that could be explored to ensure continuation of taxonomic research regarding the Flora of Cameroon, existing collaboration should be formalized by the signature of a Memorandum Rodriguésia 66(4):

38 Cameroon in the World Flora 2020 of Collaboration (MOC) between the Royal Botanic Gardens, Kew (RBG, Kew) and the National Herbarium of Cameroon (NHC). This important collaboration lead to the completion of a large number of conservation-lead projects, achieving the production of ten checklists, including seven regional checklists (Cable & Cheek 1998; Cheek et al. 2000, 2004, 2010, 2011; Harvey et al. 2004, 2010) with IUCN assessments at global level, a Red Data Book of flowering plants with IUCN assessments at global level (Onana & Cheek 2011), a taxonomic checklist (Onana 2011) and a checklist of endemic and rare species at the national level (Onana 2013). The purpose of signing a MOC would be to support the inclusion of NHC data amongst the contents of the World Flora 2020 project. The past record of RBG, Kew working together with NHC has helped the latter s involvement in collection-based international projects such as African Plant Initiative (API) and Global Plants Initiative (GPI) with 1002 items (images of types and endemic species specimens) from NHC available on-line through JSTOR Global Plants (< Similarly, French cooperation through the Institut de Recherche pour le Développement (IRD) and the Museum national d Histoire naturelle de Paris, with the database comprising herbarium label data that began in 2002 and is still ongoing. Data from 65,000 specimens of the working collection are already in the database. In the past, French cooperation has helped Cameroon to establish its portal for the Global Biodiversity Information Facility (GBIF), the Cameroon Biodiversity Information Facility (CamBIF) with a pilot project which permitted the insertion of c. 5,000 primary plant data records in the international network database of GBIF. The Clearing House Mechanism (CHM) of the Ministry of the Environment of Cameroon, the Protection of Nature and Sustainable Development (MINEPDED) has implemented the establishment of a centre for documentation on biodiversity. This is an opportunity to establish a MOU between the NHC, the CHM Cameroon and key partners such as the RBG, Kew to stimulate the production of data/ information which will be displayed by the CHM website and could then be transferred to the World Flora project. 7. The importance of the World Flora project From the national point of view, and departing from the principle that a flora is only useful after it 969 is complete (van Steenis cited by Poncy & Labat 1994), it is expected that the completion of the World Flora will: Help to provide tools (keys, photographs, descriptions, distribution data) for plant identification. Facilities such as The Plant List (2013) already help to update the taxonomic status of names, even though the list is static and many names there remain unresolved. Indicate the global distribution of species. This will help to improve the knowledge of endemism, restricted distributions, rarity and other distribution patterns, such as disjunctions. The phytogeography and biogeography of our vegetation will benefit from a better understanding regarding the floristic diversity and dynamics of our ecosystems. Provide a baseline framework for the compilation of plant information, including traditional knowledge and vernacular names. Stimulate the production of the flora at national level. The families in the FDC series have not been described by Cameroonian-based taxonomists but by researchers studying plant groups at the African or global level. It may facilitate the extraction of the data relevant for Cameroon in the long run. Provide guidance regarding names and distribution of potential invasive species. At the global level, a comprehensive global flora programme will encourage many countries to begin or continue producing and updating their Floras, and those may be expanded to include other items, such as traditional knowledge. The World Flora 2020 may raise the profile of floristic activities and direct the attention from our authorities to global activities and enable us to obtain funding for our currently undervalued projects. Conclusion Although the 2020 World Flora Online project is not explicitly mentioned in the research agenda of the Flora of Cameroon, different aspects of it are currently being implemented. The production of a checklist of vascular species is well advanced and backed by a database that allows the list to be updated and corrected. Cameroon s contribution, together with its European partners, such as IRD and RBG Kew, will hopefully accelerate the production of volumes of the Flore du Cameroun and, at the same time, provide needed information for the establishment of a complete World Flora that will hopefully include the endemic species from rich and diverse Cameroon. Rodriguésia 66(4):

39 970 Onana, J.M. Acknowledgements The author would like to thank Heads of the fomer Laboratoire de Phanérogamie, Museum national d Histoire naturelle de Paris and the Keepers of the Herbarium, Archives and Library of the Royal Botanic Gardens (RBG), Kew and particulary Dr Martin Cheek formerly Head of the wet tropic Africa team in RBG, Kew for their important commitment to the knowledge of the flora and vegetation of Cameroon. I m also grateful to Dr Daniela Zappi of RBG, Kew for the translation of the summary in Portuguese, and the critical comments on the first version of this paper, and to the two anonymous reviewers for their relevant comments which help to improve the first version of the manuscript. Heartfelt thanks go to all taxonomists who described plants from Cameroon over the years. References APG III (2009). An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161: Barthlott, W.; Lauer, W. & Placke A Global distribution of species diversity in vascular plants: towards a world map of phytodiversity. Erkunde band 50: Bebber, D.P.; Carine, M.A.; Wood, J.R.I.; Wortley, A.H.; Harris, D.J.; Prance, G.T.; Davidse, G.; Paige, J.; Pennington, T.G.; Robson, N.K.B. & Scotland, R.W Herbaria are a major frontier for species discovery. Proceedings of the National Academy of Sciences of the U.S.A. Available at < pnas.org/cgi/contens/short/ >. Access on 28 May Brummitt, R.K. (ed.) Vascular plant families and genera. Royal Botanic Gardens, Kew. 804 p. Cable, S. & Cheek, M The plants of Mount Cameroon. A conservation checklist. Royal Botanic Gardens, Kew. 198p. Campbell, D.G & Hammond H.D. (eds.) Floristic inventory of tropical countries. New York Botanical Garden, New York. 545p. Chase, M.W & Reveal, J.L A phylogenetic classification of the land plants to accompagny APG III. Botanical Journal of the Linnean Society 161: Cheek, M Limitations: future work needed. In: Onana, J.M. & Cheek, M. (eds.). Red Data Book of the flowering plants of Cameroon: IUCN global assessments. Royal Botanic Gardens, Kew. Pp Cheek M.; Harvey Y. & Onana J.-M The Plants of Dom, Bamenda highlands, Cameroon. A conservation checklist. Royal Botanic Gardens, Kew. 162p. Cheek M.; Harvey Y. & Onana J.-M The Plants of Mefou proposed national park, Yaoundé, Cameroon. A conservation checklist. Royal Botanic Gardens, Kew. 251p. Cheek, M. & Onana, J.M Red data plant species in Cameroon. A guide for secondary schools teachers, Royal Botanic Gardens, Kew. 17p. Cheek M.; Onana, J.M. & Pollard B.J The Plants of Mount Oku and the Ijim ridge, Cameroon. A conservation check-list. Royal Botanic Gardens, Kew. 211p. Cheek M.; Pollard, B.J.; Darbyshire, I.; Onana, J.M. & Wild, C The plants of Monts Kupe, Bakossi and Mwanengouba. A conservation check list. Royal Botanic Gardens, Kew. 504p. Chevillotte, H. & Florence, J. with the collaboration of Achoundong, G.; Chenin, E.; Ghogue, J.P.; Hoff M.; Labat, J.N.; Murail, J.-F. & Poncy, O RIHA, a database on plants biodiversity in Western and Central Africa: first step for a networking of African herbaria. In: Ghazanfar S.A. & Beentje H. (eds). Taxonomy and ecology of African plants, their conservation and sustainable use. Royal Botanic Gardens, Kew. Pp Costello, M.J.; May, R.M. & Stork, N.E Can we name Earth s species before they go Extinct? Science 339: DOI: /science Darbyshire, I.; Saltmarsh, A. & Malcolm, P Application of the ALUKA African plants resource to African Flora projects. In: Burgt, X. van der.; Maesen, J. van der. & Onana, J.-M. (eds), Systematics and Conservation of African Plants. Royal Botanic Gardens, Kew. Pp Harvey Y.; Pollard B.J.; Darbyshire I.; Onana J.-M. & Cheek M The plants of Bali Ngemba forest reserve. A Conservation check list. Royal Botanic Gardens, Kew. 154p. Harvey Y.; Tchiengue, B. & Cheek M The plants of Lebialem Highlands, Cameroon. A Conservation checklist. Royal Botanic Gardens, Kew. 170p. Haston, E.; Richardson, J.E.; Stevens, P.F.; Chase, M.W. & Harris, D.J The Linear Angiosperm Phylogeny Group (LAPG) III: a linear sequence of the families in APG III. Botanical Journal of the Linnean Society 161: Holmgren, P.K.; Holmgren, N.H. & Barnett, L.C Index Herbariorum. 8 th ed. New York Botanical Garden, New York. 693p. IUCN IUCN Red list categories and criteria: version 3.1. Second edition IUCN Species Survival Commission. IUCN, Gland, Switzerland and Cambridge. 32p. Joppa, L.N.; Roberts, D.L. & Pimm, S.L How many species of flowering plants are there? Rodriguésia 66(4):

40 Cameroon in the World Flora 2020 Proceedings of the Royal Society of London, serie B. Biological Sciences 278: DOI: /rspb Kenfack, D Systematic studies in Carapa (Meliaceae-Swietenioideae). Ph.D.Thesis. University of Missouri-Saint Louis, Saint Louis. 265p. Küper, W.; Sommer, J.H.; Lowett, J.C. & Bartholott, W Deficiency in African plant distribution data - missing pieces of the puzzle. Botanical Journal of the Linnean Society 150: Letouzey, R Les botanistes au Cameroun. In: Aubréville, A. (ed.). Flore du Cameroun 7. Muséum national d Histoire naturelle, Paris. 110p. Letouzey, R Végétation. In: Laclavère, G. (ed.). Atlas de la République Unie du Cameroun. Les Editions J.A., Paris. Pp Letouzey, R Carte Phytogéographique du Cameroun au 1: , accompagnée de la Notice en cinq fascicules: 1) S-S: Domaine sahélien et soudanien; 2) M-SM: Région montagnarde et étage submontagnard ; 3) SC: Domaine de la forêt dense humide semi-caducifoliée ; 4) TV: Domaine de la forêt dense humide toujours verte. Institut de la Carte Internationale de la Végétation, Toulouse. 240p. Morin, S. 1979a. Relief et hydrographie. In: Laclavère, G. (ed.). Atlas de la République Unie du Cameroun. Les Editions J.A., Paris. Pp Morin, S. 1979b. Géomorphologie. In: Laclavère, G. (ed.). Atlas de la République Unie du Cameroun. Les Editions J.A., Paris. Pp Ngembou K.C Phylogénie, distribution, écologie et révision taxonomique du genre Bertiera (Rubiaceae) en Afrique. Thèse de Doctorat en Sciences. Université Libre de Bruxelles, Bruxelles. 225p. Onana, J.M Etat de connaissance de la flore du Cameroun. In: van der Burgt, X.M.; van der Maesen, L.J.G. & Onana, J.M. (eds.). Systématique et Conservation des Plantes Africaines. Royal Botanic Gardens, Kew. Pp Onana, J.M The vascular plants of Cameroon. A taxonomic check list with IUCN assessements. Flore du Cameroun 39. IRAD-National Herbarium of Cameroon, Yaoundé. 195p. Onana, J.M Synopsis des espèces végétales vasculaires endémiques et rares du Cameroun. Check-liste pour la conservation et la gestion durable de la biodiversité. In: Onana J.-M. (ed.). Flore du Cameroun 40. Ministère de la Recherche Scientifique et de l Innovation, Yaoundé. 279p Onana, J.M Environnement biophysique, In: Onana, J.-M (ed.). Synopsis des espèces végétales vasculaires endémiques et rares du Cameroun. Checkliste pour la conservation et la gestion durable de la biodiversité. Flore du Cameroun Ministère de la Recherche Scientifique et de l Innovation, Yaoundé. Pp Onana, J.-M. & Cheek, M Red Data Book of the flowering plants of Cameroon: IUCN global assessments. Royal Botanic Gardens, Kew. 578p. Onana J.M. & Chevillotte H Taxonomie des Rutaceae-Toddalieae du Cameroun revisitée: découverte de quatre espèces nouvelles, validation d une combinaison nouvelle et véritable identité de deux autres espèces de Vepris Comm. ex A.Juss. Adansonia, sér. 3, 37: Poncy, O & Labat, J.-N. Etat actuel de l inventaire des flores tropicales. Conférence lors des Journées annuelles de la Société française de systématique en septembre 1994, Paris. Document non publié. République du Cameroun Cinquième Rapport National du Cameroun à la Convention de la Diversité Biologique - Ministère de l Environnement, de la Protection de la Nature et du Développement Durable. 100p. Roux, J.P Synopsis of Lycopodiophyta and Pteridophyta of Africa, Madagascar and neighbouring islands. Strelitza 23. South African Biodiversity Institute, Pretoria. 296p. Satabié, B Contribution à la connaissance de la biodiversité de la flore camerounaise. In: Timberlake, J. & Kativu, S. (eds.). African Plants: Biodiversity, Taxonomy and Uses. Royal Botanic Gardens, Kew. Pp Stone, R.D Taxonomic treatment of Memecylon L. section Felixiocylon R.D.Stone (Melastomataceae), with descriptions of four new species from Cameroon, Gabon, and Equatorial Guinea (Bioko). Adansonia, sér. 3, 37: Stone R.D.; Ghogue J.-P. & Cheek M Revised treatment of Memecylon sect. Afzeliana (Melastomataceae: Olisbeoideae), including three new species from Cameroon. Kew Bull 63: Surtcliffe, J. & O Reilly, C. (2010). Ecological skills: mind the gap(s). Kew Bulletin 65 : Taedoumg, E.H Etude biosystématique du genre Cratrispermum Benth.(Rubiaceae) en Afrique continentale. Thèse de Doctorat/PhD. Université de Yaoundé I, Yaoundé. 269p. The Plant List Version 1.1. Published on the Internet. Available at < org/>. Access on 6 June Tchawa, P Relief et hydrographie. In: Ben Yamed, D.; Houstin, N & Seignobos, C. (eds.). Cameroun. Atlas de l Afrique. Les Editions J.A., Paris. Pp Tchouto, M.P.G Plant diversity in a Central African rain forest: implication for conservation in Cameroon. PhD Thesis. Wageningen University, The Netherlands. 208p. Rodriguésia 66(4):

41 972 Onana, J.M. Tchouto, M.G.P. & Elad, M Liana composition and diversity in the Campo-Ma an rainforest in southern Cameroon. In: van der Burgt, X.M.; van der Maesen, J. & Onana, J.-M. (eds.). Systematics and conservation of African plants. Royal Botanic Gardens, Kew. Pp Tsaléfac, M Climat. In: Ben Yamed, D.; Houstin, N. & Seignobos, C. (eds.). Cameroun. Atlas de l Afrique. Les Editions J.A., Paris. Pp van der Burgt, X.M Lecomtedoxa plumosa (Sapotaceae), a new tree species from Korup National Park, Cameroon. Kew Bulletin 64: van der Burgt, X.M Two new taxa in Magnistipula (Chrysobalanaceae) from Korup National Park, Cameroon. Plant Ecology and Evolution 143: van der Burgt, X.M.; Eyakwe, M. & Motoh, J Gilbertiodendron newberyi (Leguminosae: Caesalpinioideae) a new tree species from Korup National Park, Cameroon. Kew Bulletin 67 : van der Burgt, X.M. & Newbery, D.M Glumea korupensis (Sapotaceae), a new tree from Korup National Park, Cameroun. Kew Bulletin 61: van der Burgt, X.M.; Mackinder, B.A.; Wieringa, J.J. & de la Estrella, M The Gilbertiodendron ogoouense species complex (Leguminosae: Caesalpinioideae), Central Africa. Kew Bulletin 70: 29. DOI: / S van der Zon, A.P.M Graminées du Cameroun 2. Wageningen. Agricultural University Papers 92: White, F The vegetation of Africa. In: UNESCO (ed.). A descriptive memoir to accompagny the U.N.E.S.C.O. / AETFAT / UNSO Vegetation Map of Africa. UNESCO, Paris. 356p. WCSP World checklist of selected plant families. Facilitated by the Royal Botanic Gardens, Kew. Published on the Internet. Available at < apps.kew.org/wcsp/>. Access 6 June Artigo recebido em 07/07/2015. Aceito para publicação em 03/09/2015. Rodriguésia 66(4):

42 Rodriguésia 66(4): DOI: / Flora of North America North of Mexico Nancy R. Morin 1, Luc Brouillet 2 & Geoffrey A. Levin 3 Abstract The Flora of North America north of Mexico treats all native and naturalized vascular plants and bryophytes in Canada, Greenland, St. Pierre et Miquelon, and the continental United States including the Florida Keys and Aleutian Islands (approximately 18 million square kilometers). It provides accepted names, literature citations, basionyms, synonyms, morphological descriptions, habitat, geographical distribution, conservation or weed status, and a discussion of taxonomic issues for approximately 20,000 species. Of the total 30 volumes anticipated, 18 have been published and one is in press, treating 2021 genera and 12,393 species. For the remaining volumes, 763 genera and 5,008 species have been submitted, and 82 of the 144 families have been submitted in full. Completion is anticipated by the end of The project is managed by the Flora of North America Association. Content from published volumes is available through efloras and JSTOR and has been provided to the World Flora informatics team. Key words: Flora, North America, United States, Canada, Greenland. Resumo A Flora da América do Norte ao Norte do México trata todas as espécies de plantas vasculares e briófitas ocorrente sno Canadá, Groenlândia, St. Pierre et Miquelon e na parte continental dos Estados Unidos, incluindo as Florida Keys e as Aleutian Islands (aproximadametne 18 milhões de kilômetros quadrados). Informações como nomes aceitos, citações bibliográficas, basiônimos, sinonímia, descrições morfológicas, habitat, distribuição geográfica, estado de conservação e informação sobre invasoras, bem como discussões relativas à taxonomia de aproximadamente espécies. A previsão final é de 30 volumes, sendo que, nos 18 já publicados e naquele que está no prelo, foram tratados 2021 gêneros e 12,393 espécies. Os restantes volumes contam com a submissão de 763 gêneros e espécies, sendo que 82 das 144 famílias estão completas. A finalização da flora está prevista para O projeto é gerenciado pela Flora of North America Association. O conteúdo dos volumes publicados está disponibilizado através de e-floras e do JSTOR, e foi repassado para o time de informática do projeto World Flora. Palavras-chave: Flora, Estados Unidos da América, Canadá, Groenlândia. Introduction The Flora of North America North of Mexico comprises treatments of vascular plants and bryophytes native or naturalized in Canada, Greenland, St Pierre et Miquelon, and the continental United States including the Florida Keys and Aleutian Islands (approximately 18 million square kilometers). Distribution information for Mexico, especially the northern tier of states, is provided, as well as generalized world-wide distribution. Treatments include accepted name, basionym, place of publication, synonyms, vernacular names, morphological descriptions, flowering time, habitat, elevation, geographical distribution, discussions, and references. Every genus and 1/6 of the species are illustrated by professional botanical artists working closely with authors and editors, assuring that the drawings reflect the author s concepts and complement the technical descriptions in a way that assists the general user. Small distribution maps are provided for all species and infraspecific taxa. 1 Flora of North America Association, P.O. Box 716, Point Arena, California 95468, USA. nancy.morin@nau.edu 2 Institut de recherché en biologie végétale, Université de Montréal, 4101 est, rue Sherbrooke, Montréal, Québec H1X 2B2, Canada. luc.brouillet@umontreal.ca 3 Illinois Natural History Survey, Prairie Research Institute, University of Illinois, 1816 S Oak St., Champaign, IL 61820, USA. levin1@illinois.edu

43 974 Morin, N.R.; Brouillet, L. & Levin, G.A. A total of 30 volumes will be published including an introductory volume and a final volume containing a cummulative index and bibliography. As of June, 2015, eighteen volumes had been published and one was in press (Tab. 1). Content of all published volumes is made available electronically through efloras and JSTOR Plant Science. Preparation of all remaining volumes is projected to be completed by the end of Table 1 Statistics from published volumes of Flora of North America north of Mexico. Volume Date published Contents Genera Species Volume Introductory chapters Volume Ferns and fern allies; Gymnosperms Volume Magnoliaceae to Casuarinaceae Volume Phytolaccaceae to Molluginaceae Volume Caryophyllaceae to Plumbaginaceae Volume 6 in press Cucurbitaceae to Droseraceae Volume Salicaceae to Brassicaceae Volume Paeoniaceae to Ericaceae Volume Picramniaceae to Rosaceae Volumes 19, 20, Asteraceae Volume Butomaceae to Marantaceae Volume Cyperaceae Volume Poaceae part I Volume Poaceae part II Volume Pontederiaceae to Orchidaceae Volume Bryophytes: Mosses part I Volume Bryophytes: Mosses part II Totals Brief history of Flora of North America north of Mexico project The scientific community has recognized the need for a continental scale flora of North America since the early 1800s. The first accounts of plants from the area were published in Europe by European botanists (e.g., in Robert Morison s Historiae [ ], Leonard Plukenet s Phytographia [ ], and John Ray s Historia Plantarum [ ]). Fredrick Pursh s Flora Americae Septentrionalis (1814) was the first flora of continental scope. Treatises on North American plants were written by resident botanists in the early 1800s, and the first attempt at production of a comprehensive flora of the continent was undertaken by John Torrey and Asa Gray the 1830s (see Reveal & Pringle 1993). Subsequently, botanists focused on producing regional floras, until 1905 when N.L. Britton began publishing the monographic series North American Flora at the New York Botanical Garden, with multiple authors. The first effort at a multiinstitutional, collaborative project started in 1966, stimulated by the completion of the Flora SSSR in 1963 (Komarov et al. 1963) and publication of the first volume of Flora Europaea in 1964 (Tutin et al. 1964). The project was to be headquartered at the Smithsonian Institution and funded ultimately through a congressional appropriation. A great deal of organizational work was done and information resources were developed but the project was suspended in 1972 because the funding plan Rodriguésia 66(4):

44 Flora of North America North of Mexico was not successful. Various other attempts were made to get the project restarted, and in 1982 a group of botanists met in St. Louis and decided to try again. A plan was developed, an editorial committee formed, and in 1987 the first funding was awarded by the Pew Charitable Trusts and the National Science Foundation. Until 1996, the Missouri Botanical Garden was the organizational center through which the project was funded, after which the Flora of North America Association was incorporated and funding, along with the work force, was distributed among Editorial Centers at various institutions. Underlying philosophy of Flora of North America north of Mexico The mission of the Flora of North America Association (FNAA) is to make the best scientific knowledge of the plants of North America north of Mexico publicly available in consistent and easy to use forms. To this end, FNAA coordinates the work of 900+ botanists to provide authoritative information on the names, circumscriptions, characteristics, and geographical and ecological distributions of the 20,000 species of plants native or naturalized in North America north of Mexico. The Flora of North America north of Mexico (FNANM) editorial committee developed policies and procedures for its completion based on recommendations made by a wide range of interested parties at a symposium titled Floristics for the 21 st Century (Morin et al. 1989). A guide for contributors was developed setting out style and format expectations, including that descriptions within a rank should be parallel to the extent possible and the descriptions should match the relevant entries in the identification key. The symposium participants recommended parallel descriptions so that FNANM would be more than a device to identify unknown plants; it should characterize the flora in a way useful to a wide range of audiences. Experts in each taxonomic group, when available, would be asked to write treatments of their groups in order to assure that the most authoritative, up-to-date information and taxonomic concepts were employed. Scientific reliability was deemed essential if the information was to inform management and conservation decisions. Regional reviewers (now numbering about 100) would review treatments to assure that each taxon was treated appropriately throughout 975 its range. The editorial committee concluded that decisions should be made by the people doing the work and that authors and editors would mostly work on a volunteer basis. Content A flora may be a checklist, an identification key to included taxa, a list of accepted species with distributions, a list of accepted names with brief descriptions, literature citations, synonyms, some with maps, some with illustrations, or various combinations of these components. Initially, the expectation was that the accounts in FNANM would be similar to those in Flora Europaea, with keys, short descriptions, minimum discussion, and no maps or illustrations, and that the entire series would comprise 14 volumes. After consideration, small distribution maps were added to help give users a quick visual impression of the geographical range of a taxon. The geographical area, continental North America north of Mexico, the Florida Keys, Aleutian Islands, St. Pierre et Miquellon, and Greenland, was selected because it forms a reasonable phytogeographical region. Mexico was not included because there were other efforts to treat that flora (regional floras, Flora Mesoamericana, Davidse et al. 1994), but distribution statements in FNANM include at least the northern tier of Mexican states when possible. Illustrations, initially for every genus and one-third of the species, were added to make the treatments more useful to non-botanists, and to show complex features difficult to portray with just words. The length of descriptions was expected to be 700 characters or less, but, for larger families and genera, the descriptions must be longer in order to make descriptions parallel and to assure that they contain features used in the key. Many of the treatments represent a botanist s lifetime of work, work that might otherwise never be published elsewhere. The editorial team has attempted to balance realistic constraints of time and resources (including a goal of completing the project within their lifetime, and physical limitations of number of pages that can be bound in one volume) with a wish to convey in the best possible way this wealth of knowledge. Taxonomic Arrangement When decisions were first being made about the content of Flora of North America volumes, the most comprehensive published modern Rodriguésia 66(4):

45 976 Morin, N.R.; Brouillet, L. & Levin, G.A. classification system for plants was by Arthur Cronquist (1981), and that work was adopted to set the circumscriptions of families and the order in which they would appear in the volumes. Beginning in 2007, Flora of North America adopted the system outlined by the Angiosperm Phylogeny Group (2003) for remaining volumes, to the extent practicable. Authors are encouraged to order genera and species according to their best understanding of relationships. If this is not possible, the taxa may be arranged alphabetically. Community and financial support In-kind support in the form of facilities and staff time has been essential for the success of the project. Editorial centers are or have been located at the The Arboretum at Flagstaff, Arizona (Volume 4), Botanical Research Institute of Texas, Fort Worth (Asteraceae, Volumes 19, 20, and 21), California Academy of Sciences, San Francisco (Volume 18 - now moving), Hunt Institute for Botanical Documentation, Pittsburgh (bibliography and Volumes 5, 6, and 14, 26, 30), Harvard University Herbaria (nomenclature), Illinois Natural History Survey, Champaign together with Canadian Museum of Nature (Volume 12), University of Kansas together with University of Michigan (Volumes 5, 8, 17), Missouri Botanical Garden (Volumes 1, 2, 3, 7, 10/11, 23, and the bryophyte volumes, 27, 28, 29), University of Montreal (Volumes 9, 13), University of Washington (Volume 15), and Utah State University (Poaceae, Volumes 24 and 25). Most editors, authors, and reviewers donate their time and expertise. Botanical artists and technical editors work on a contract basis and are coordinated by the Managing Editor and Scientific Director located at Missouri Botanical Garden. When funding has been available, a botanist has been employed on contract to write treatments for which no specialist is available. Botanical artists and technical editors, the Managing Editor, and half of the Scientific Director s time is paid by grants, contributions, art sponsorships, and earned income (royalties, etc.). Also funded are basic organizational costs such as insurance, financial audits, meetings, etc. The botanical community has supported work on FNANM by developing cooperative groups to agree on taxonomic approaches and engaging the efforts of specialists. For example, Volumes 24 and 25, which treat Poaceae, were prepared in collaboration with a large group of grass specialists working through Utah State University and funded in part by the U.S. Department of Agriculture. Volumes 19, 20, and 21, treating Asteraceae, were prepared under the guidance of a team of noted synantherologists coordinated at the Botanical Research Institute of Texas and funded by a grant from the U. S. National Science Foundation. Volumes 27, 28, and 29, Bryophytes, are being completed by a team of bryologists. The initial funding for the current Flora of North America project from the Pew Charitable Trusts was matched by a grant from the U. S. National Science Foundation, which made additional grants through A wide range of foundations made grants to the project from 1987 through The Chanticleer Foundation provided core funding The Andrew W. Mellon Foundation and an anonymous foundation also provided core funding in 2008 through Funding since 2012 has been received from the Philecology Foundation, the Franklinia Foundation, the David and Lucile Packard Foundation and other organizations and individuals. Information Technology The original Flora North America project was far ahead of its time in its vision of management and analysis of botanical data using computers. The 1967 proposal to NSF outlined goals of developing an information retrieval system and the ability to automatically update information continuously and retrieve it in newly combined forms. At the time, research indices like Index Kewensis were compiled by hand. The FNA organizers hoped they could provide the initial stimulus and technical foundation for development of comprehensive information retrieval systems. The current Flora of North America organizers decided to focus their efforts on acquiring the botanical treatments rather than developing information management tools. Nonetheless, the potential for using computers to assist with preparation of the flora was kept in mind. (Before the World Wide Web became publicly available in 1990, all manuscripts were duplicated and distributed using the U. S. Postal Service.) TROPICOS, a system developed by Missouri Botanical Garden botanists Robert Magill and Marshall Crosby in the early 1980s to manage plant nomenclature and bibliography, was adopted as the system in which FNA nomenclature Rodriguésia 66(4):

46 Flora of North America North of Mexico would be maintained. A bibliographic database for FNA was developed by Robert Kiger at the Hunt Institute for Botanical Documentation. Grants from the U. S. National Science Foundation supported development of innovative tools for collaborative work processes, envisioning the potential for multiple participants to work on the same manuscript and to monitor tasks and accomplishments. An initial goal of creating a massive relational database of characteristics has evolved into the use of semantic markup tools developed by James Macklin and Hong Cui (Cui 2010; Morris et al. 2013). Treatments parsed into their basic components are available through efloras (< and JSTOR (< Resource materials for authors and reviewers, links to published volumes, status of volumes in preparation, and other information about the FNANM project, are available on the FNA website: In the past year the FNA website received about 1,371,540 successful requests for 377,356 pages - on average 26,000 successful requests per week. Progress to date Flora of North America now has published 18 of the total 30 volumes and has one volume in press (Volumes 1-9, 19-28; 11,200 pages), containing 253 families, 2,125 genera, and 12,949 species (Flora of North America Editorial Committee, 1993+). These volumes also contain original illustrations of 7,200 species, subspecies, or varieties and maps of every species and infraspecific taxon. Volume 1, published in 1993, includes introductory chapters on the history of floristics and plant collecting in North America; climate and physiography; soils; history of the vegetation and paleoclimates, vegetation, and floras; vegetation, and phytogeography. Completing Flora of North America north of Mexico by 2020 The taxonomic volumes remaining to be completed: Volumes and 29, contain 144 families, 1103 genera, and 7365 species. Of these, 763 genera and 5008 species have been submitted, and 82 of the 144 families have been submitted in full. Volume 12, Vitaceae to Garryaceae, 29 families, 122 genera, and 760 species, is in final stages of editing and will be in press this fall. Volume 17, Polypremaceae to Orobanchaceae, containing 9 families, 94 genera, and 942 species, 977 will be in press by the end of 2015 or early For Volumes 10 and 11, being prepared together and containing largely Fabaceae and Onagraceae, all but those two families have been submitted in full and 91% of the genera and species are in hand and in process of review, revision, and editing. For the remaining five volumes containing treatments of vascular plants, more than half of the families and genera have been submitted and are being edited. All other treatments have authors assigned and are in preparation. The final Bryophyte volume, containing liverworts and hornworts, is also making good progress. Status of individual treatments in volumes to be published can be found at www. floranorthamerica.org. Methodology Each volume is assigned to an editorial center under the leadership of a lead editor (or co-lead editors). Each family is assigned to a taxon editor who works directly with authors, reviewers, and technical editors. Authors are selected based on their interest and expertise and with approval of the FNAA Board. Technical editors work on a contract basis; each one is assigned to a volume. Regional review is managed by regional coordinators, who send draft treatments to reviewers knowledgeable in their local flora and compile responses, which are returned to taxon editors. Authors provide material to botanical artists so they can prepare illustrations. Every genus and about one in six species are illustrated by these contract artists. Authors are asked to use terminology set out in (Kiger & Porter 2001). Names accepted in major regional floras are supplied to authors and are to be accounted for in the treatments. Robert Kiger, Hunt Institute for Documentation, edits all bibliographic references and compiles the cumulative bibliography. Kanchi Gandhi, Harvard University Herbaria, reviews nomenclature and citations. Some aspects of final production, such as indexing and page composition, are carried out by contract workers under the supervision of Heidi Schmidt, managing editor, and James Zarucchi, editorial director, who also oversee all manuscript flow. Challenges for completion include editorial capacity and time available to volunteer authors and reviewers. Botanists with broad floristic knowledge are becoming scarce, and there are plant groups for which no specialist is available. Fortunately, FNANM has been able to enlist the help of botanists who have recently completed their graduate degrees or are currently graduate students. Rodriguésia 66(4):

47 978 Morin, N.R.; Brouillet, L. & Levin, G.A. Resources for completing the Flora of North America north of Mexico 1. Regional Floras The Flora of North America project draws on a wealth of resources. A working checklist has been available from several sources, including the United States Department of Agriculture PLANTS database and The Natural Heritage Program through its Natureserve database. Canada has a working checklist through VASCAN, part of Canadensys. North America north of Mexico is a very large geographic area, politically and biogeographically diverse, so it is understandable that floristic work has focused on states, provinces, or regions. Regional floras recently completed for the United States include The Jepson Manual, covering all of California (Baldwin et al. 2012); the Intermountain Flora, covering all of the Great Basin (Cronquist et al. 1972); Flora of the Southern and MidAtlantic States (Weakley 2012); and Flora Novae Angliae (Haines 2011). Less recent but still important are Flora of the Great Plains (Great Plains Flora Association 1986); Manual of the Vascular Plants of Texas (Correll & Johnston 1970); and Manual of vascular plants of northeastern United States and adjacent Canada, second edition (Gleason & Cronquist 1991). For Canada, a basic reference is the Flora of Canada (Scoggan ); Budd s Flora of the Canadian Provinces (Budd et al. 1987); The vascular plants of British Columbia (Douglas et al ); Vascular plants of the continental Northwest Territories (Porsild & Cody 1980); Flore Laurentienne (Marie-Victorin, revised by Brouillet et al. 1995); Atlas of the vascular plants of the island of Newfoundland and of Saint Pierre et Miquelon (Rouleau and Lamoureux 1992); and The flora of New Brunswick (Hinds 2000). In preparation is a Flora of the vascular plants of the Canadian Arctic Archipelago, updating the Illustrated Flora of the Canadian Arcitc Archipelago (Porsild 1964). 2. Online specimen, occurrence, and nomenclatural databases Some regional or state specimen databases also functionally serve as checklists. They are a rich resource for authors and reviewers. As Flora of North America volumes are published, the databases are updated to reflect new classifications. Examples of such databases are: Country-wide Databases 2.1 Canadensys (< explorer/en/search>) The checklist covers all vascular plants reported in Canada, Greenland (Denmark), and Saint Pierre and Miquelon (France). The latter two regions are added because their floras are intimately related to that of Canada. Provincial distributions are provided to help users visualize the relationship among the floras of Canadian provinces and territories. VASCAN does not intend to replace regional or provincial lists but to act as a complement to them. The covered regions are, in alphabetical order: Alberta, British Columbia, Greenland, Labrador, Manitoba, New Brunswick, Newfoundland, Northwest Territories, Nova Scotia, Nunavut, Ontario, Prince Edward Island, Quebec, Saint Pierre and Miquelon, Saskatchewan, and Yukon. The distribution status of the plant is indicated per region. These can be grouped as present (native, introduced or ephemeral), previously reported but currently considered absent (excluded, extirpated), doubtful or not reported (absent). The latter status is not recorded in the database (null value). Excluded taxa are those considered not currently occurring in a region, due either to nonrecurring ephemeralness, misidentification, lack of supporting documentation, or when specimens are old and the taxon has not been observed again in more than 50 years. All distribution statuses are defined at (< about/#distribution>). The VASCAN website (< provides a distribution map for each taxon. 2.2 The PLANTS Database (< gov>) Provides standardized information about the vascular plants, mosses, liverworts, hornworts, and lichens of the U.S. and its territories. It includes names, plant symbols, checklists, distributional data, species abstracts, characteristics, images, crop information, automated tools, onward Web links, and references. It can be searched by taxonomy, morphological and habitat characteristics, geographical distribution, and weed, wetland, and conservation status. 3. Specimen Databases available online that also can generate checklists Most of these online databases are collaborations among regional networks of herbaria and provide links to the individual member organizations. Rodriguésia 66(4):

48 Flora of North America North of Mexico 3.1 Canadensys Explorer (< canadensys.net/explorer>) Serves 180,000 vascular plant specimen records from 13 Canadian herbaria. 3.2 Consortium of California Herbaria (< ucjeps.berkeley.edu/consortium>) The Consortium of California Herbaria provides data from over 2 million specimen records from over 30 institutions. 3.3 Consortium of Pacific Northwest Herbaria (< The Consortium of Pacific Northwest Herbaria hosts more than 2.4 million specimen records from 33 herbaria in Alaska, Idaho, Montana, Oregon, Washington, Wyoming, British Columbia, Yukon Territory, and Humboldt State University in northern California. 3.4 Intermountain Regional Herbarium Network (< The Intermountain Regional herbarium Network hosts collection data from herbaria in Nevada and Utah. (also served throught SEINET: (< 3.5 Southwest Environmental Information Network (< The Arizona Chapter of SEINET serves collections and observation data from 14 institutions in Arizona. SEINet allows searching across many major regional herbarium networks. 3.6 Flora of Texas Database (< utexas.edu/prc/tex.html>) Serves records of occurrences in Texas based on University of Texas and Lundell herbaria. 3.7 Rocky Mountain Regional Consortium Specimen Database (< Provides searching across 14 herbaria in Colorado, Idaho, Montana, and Wyoming (also served throught SEINET: (< org/seinet/index.php>)). 3.8 Consortium of Midwest Herbaria (< midwestherbaria.org/portal>) Hosts specimen data from 21 herbaria in Illinois, Indiana, Iowa, Michigan, Minnesota, Ohio, and Wisconsin. (also served through SEINET: (< Northern Great Plains Herbaria (< Hosts data from 19 herbaria in Illinois, Iowa, Kansas, Minnesota, Missouri, Nebraska, North Dakota, and South Dakota, (also served through SEINET: (< php>)) Consortium of Northeastern Herbaria (< Serves 776,600 records of vascular plants and bryophytes from 21 herbaria in Nova Scotia, Quebec; Connecticut, Massachusetts, New Hampshire, New Jersey, New York, Pennsylvania, Rhode Island, and Vermont Southeast Regional Network of Expertise and Collections (< Hosts records from 110 herbaria in Alabama, Arkansas, Delaware, Florida, Georgia, Kentucky, Louisiana, Maryland, Mississippi, North Carolina, South Carolina, Tennessee, Texas, Verginia, and West Virginia (also served through SEINET: (< Atlas of Florida vascular Plants (< Serves records from over 110,000 herbarium specimens from 8 herbaria in Florida. Conclusion Improving the knowledge of the Flora of North America north of Mexico Flora of North America north of Mexico has stimulated the completion and publication of research by authors of treatments and presents the best, most current understanding of the relationships, characteristics, and distribution of plants in the flora. It brings together in a standard, carefully reviewed form, the results of lifetimes of study and the findings of the most recent research. Aspects of phylogeny, biology, or ecology that need more study are mentioned in the discussion, which stimulates new research. Because the descriptions and discussion characterize the plants in more detail than more abbreviated floras or checklist, they provide a rich resource for research by ecologists, plant breeders, plant morphologists, physiologists, to name just a few. This resource will be much enhanced by the current effort to parse all descriptions, distributions, and habitats from published volumes. Rodriguésia 66(4):

49 980 Morin, N.R.; Brouillet, L. & Levin, G.A. Importance of Flora of North America for Environmental Research Flora of North America provides a common reference for the circumscription of plant taxa, correct application of names to those taxa, and descriptive information for each taxon. This is critically important for every kind of research from broad ecological studies to phenomics, genomics, and phylogenetic analysis. Although there are checklists of the plants of North America already available, none have the same level of scientific authority, and, since they lack descriptions, they are not sufficient to determine what is actually included within the named concept. There are few equivalents in terms of digital datasets in the animal or insect worlds, and none dealing with as many species. Importance of the World Flora It would be highly useful for further research, for conservation, and for resource management to have a common reference for the circumscription of plant taxa, correct application of names to those taxa, and descriptive information for each taxon, and worldwide distribution for all plants. Only elements of this, in various guises, are so far available. For Flora of North America north of Mexico, based on volumes published to date, 13% of the genera and 50% of the species are endemic to the flora area. FNANM has provided full descriptions and abbreviated worldwide distributions for 6500 species that also occur outside the flora area and for nearly 1900 genera that also occur outside the flora area. Of these, 315 of the genera and 1791 of the species in published volumes are introduced. Currently it is very difficult for authors to find the relevant data on the non-native taxa, as it must be for researchers outside North America to find data on the 10,000 species likely to be endemic in North America north of Mexico, since much of that information is contained in local floras or individual monographs. A World Flora would facilitate access to these data. Integrating Flora of North America north of Mexico data with the World Flora All published treatments are available in electronic form, and soon all will be fully parsed into individual components. The electronic files for volumes published have already been provided to the World Flora Online for incorporation. References Angiosperm Phylogeny Group An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141: Baldwin, B.G.; Goldman, D.; Keil, D.J.; Patterson, R.; Rosatti, T.J. & Wilken, D. (eds.) The Jepson manual: vascular plants of California, revised and expanded. University of California Press, Berkeley. 1568p. Britton, N.L. et al. (eds.) North American Flora. 47 vols. New York Botanical Garden, New York. Vols. 1-34, ; ser. 2, parts 1-13, Budd, A.C.; Looman, J. & Best, K.F Budd's flora of the Canadian Prairie Provinces, revised and enlarged. Publication Agriculture Canada Research Branch, Ottawa and Hull. 863p. Correll, D.S. & Johnston, M.C Manual of the vascular plants of Texas. Texas Research Foundation, Renner. 1881p. Cronquist, A An integrated system of classification of flowering plants. Columbia University Press, New York p. Cronquist, A.; Holmgren, A.H.; Holmgren, N.H.; Reveal, J.L.; Holmgren, P.K. & Barneby, R.C Intermountain Flora. Vascular plants of the Intermountain West, U.S.A. 8 vols. New York Botanical Garden, New York and London. 3865p. Cui, H Semantic annotation of morphological descriptions: an overall strategy. BMC Bioinformatics 11: 278. Davidse, G.; Sousa S.M. & Chater, A.O. et al. (eds.) Flora Mesoamericana. 4 vols. Universidad Nacional Autónoma de México, Instituto de Biología, Ciudad Universitaria, Mexico City. Missouri Botanical Garden Press, St. Louis. 2788p. Flora of North America Editorial Committee (eds.) Flora of North America North of Mexico. 19+ vols. Oxford University Press, New York and Oxford. 12,303p. Douglas, G.W.; Straley, G.B. & Meidinger, D.V The vascular plants of British Columbia. 4 vols. British Columbia Ministry of Forests, Special Report, Victoria. 800p. Great Plains Flora Association Flora of the Great Plains. University Press of Kansas, Lawrence. 1382p. Gleason, H.A. & Cronquist, A Manual of vascular plants of northeastern United States and Adjacent Canada. 2 nd ed. New York Botanical Garden, Bronx. 910p. Haines, A New England Wild Flower Society s Flora Novae Angliae: a manual for the identification of native and naturalized higher vascular plants of New England. Yale University Press, New Haven, Connecticut and London, United Kingdom. 1008p. Rodriguésia 66(4):

50 Flora of North America North of Mexico Hinds, H.R Flora of New Brunswick, second edition: a manual for identification of the vascular plants of New Brunswick. Department of Biology, University of New Brunswick, Fredericton. 699p. Kiger, R.W. & Porter, D.M Categorical glossary for the flora of North America project. Hunt Institute for Botanical Documentation, Carnegie Mellon University, Pittsburgh. 165p. Komarov, V. L. et al. (eds.) Flora of the U.S.S.R. (Flora SSSR). 30 vols. Translated from Russian. Israel Program for Scientific Translations, Jerusalem p. Marie-Victorin, Frère Flore Laurentienne. 3 rd ed. Annotated and updated by Luc Brouillet and Isabelle Goulet. University of Montreal Press, Montreal. 1083p. Morin, N.R.; Whetstone, R.D.; Wilken, D.H. & Tomlinson, K.L. (eds.) Floristics for the 21 st century. Proceedings of the workshop sponsored by the American Society of Plant Taxonomists, 4-7 May, 1988, Alexandria, Virginia. Monographs in Systematic Botany, Missouri Botanical Garden 28: Morison, R Plantarum historiae universalis oxoniensis. 2 vols. Theathro Sheldoniano, Oxford. 1498p. Morris R. A.; Dou, L.; Hanken, J.; Kelly, M.; Lowery, D.B. & Ludäscher, B. et al Semantic annotation of mutable data. PLoS ONE 8: e DOI: /journal.pone Plukenet, L Phytographia, sive, illustriorum & miniis cognitarum icones tabulis aeneis : summa diligentia elaboratae, quarum unaquaeq[ue] titulis descriptoriis ex notis suis propriis & characteristicis desumptis insignita, ab aliis ejusdem sortis facile descriminatur. Published by the author. 7 parts. 981 Porsild, A. E Illustrated flora of the Canadian Arctic Archipelago, 2 nd ed. Bulletin of the National Museum of Canada 146: Porsild, A.E. & Cody, W.J Vascular plants of continental northwest territories, Canada. National Museum of Natural Sciences, National Museums of Canada, Ottawa. 66 p. Pursh, F. [1813] Flora Americae Septentrionalis; or, a systematic arrangement and description of the plants of North America. 2 vols. White, Cochrane, and Co., London. 751p. Ray, J Historia plantarum : species hactenus editas aliasque insuper multas noviter inventas & descriptas complectens. Vol. 1: 1-983; vol. 2: ; vol. 3: 1-666, 1-135, Typis Mariæ Clark, prostant apud Henricum Faithorne [etc.], Londini. 983p. Reveal, J.L. & James S. Pringle Taxonomic botany and floristics. In: Flora of North America Editorial Committee (eds.). Flora of North America North of Mexico. 18 vols. Oxford University Press, New York and Oxford. Vol. 1, pp Rouleau, E. & Lamoureaux, G Atlas of the vascular plants of the Island of Newfoundland and of Saint-Pierre-et-Miquelon. Fleurbec, Saint-Henride-Lévis. 777p. Scoggan, H.J The flora of Canada. 4 parts. Publications in Botany 7. National Museum of Natural Sciences, Ottawa. 1711p. Tutin, T.G. et al. (eds.) Flora Europaea. 5 vols. Cambridge University Press, Cambridge. 2392p. Weakley, A Flora of the Southern and Mid- Atlantic States. Working Draft 20 November University of North Carolina Herbarium, Chapel Hill. 1225p. Artigo recebido em 01/07/2015. Aceito para publicação em 14/09/2015. Rodriguésia 66(4):

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52 Rodriguésia 66(4): DOI: / Hard Copy to Digital: Flora Neotropica and the World Flora Online William Wayt Thomas 1,2 & Melissa Tulig 1 Abstract One of the greatest challenges in achieving the goals of the World Flora Online (WFO) will be to make available the huge amount of botanical information that is not yet available digitally. The New York Botanical Garden is using the Flora Neotropica monograph series as a model for digitization. We describe our efforts at digitizing Flora Neotropica monographs and why digitization of hardcopy descriptions must be a priority for the WFO project. Key words: Electronic monographs, open access, Flora Neotropica, monographs. Resumo Um dos maiores desafios para alcançar as metas do projeto World Flora Online (WFO), será a disponibilizar a enorme quantidade de informações botânicas que ainda não estão disponíveis digitalmente. O New York Botanical Garden está utilizando a série de monografias da Flora Neotropica como um modelo para a digitalização. Nós aqui descrevemos nossos esforços na digitalização das monografias da Flora Neotropica e porque a digitalização das descrições impressas deve ser uma prioridade para o projeto WFO. Palavras-chave: Monografias eletrônicas, open access, Flora Neotropica, monografias. Introduction The World Flora Online (WFO) was developed as part of the United Nation s Global Strategy for Plant Conservation with the goal of providing an online flora of all known plants, One of the greatest challenges in achieving the goals of the WFO will be to make available the huge amount of botanical information that is not yet available digitally. Here, we describe our efforts at digitizing Flora Neotropica monographs, discuss why that series is our first priority, and why digitization of hardcopy descriptions must be a priority for the WFO project. World Flora Online Objective I of the United Nation s Global Strategy for Plant Conservation, states that by 2020 plant diversity should be expertly characterized and documented in a standardized format. In response, a consortium of global research institutions, endorsed by the international botanical community, will create a definitive, verified online resource documenting all known plants of the world. It is called the World Flora Online (WFO). This consortium of professionals will create openaccess one-stop searching of world flora with verified information, including new and previously published data, and coordinated with links to other plant database and catalog Web sites. The core information in the World Flora Online will be vetted by professional botanists and will include (*highest priority): A comprehensive list of names for accepted species*, with synonyms placed with each correct name. Published modern descriptions, as well as the protologue* (the original published description when the species was first described in scientific literature). Literature citations for each name*, with reference to the type specimen for each name. Images of each species, including digital images of herbarium specimens (including connections to the type images in JSTOR Global Plants) as well as photographs of living plants. Links to other portals with plant information (example: the herbarium specimen data for 1 The New York Botanical Garden, Bronx, NY , USA. 2 Author for correspondence: wthomas@nybg.org

53 984 Thomas, W.W. & Tulig, M. each species through the Global Biodiversity Information Facility (GBIF), among others). Geographic distribution of each species. Conservation status of each species. Plant Diversity of the Neotropics The Neotropics have functioned as an engine for generating plant diversity (Antonelli et al. 2015) but why do they deserve attention as a botanical priority, and as a test case for digitization? The Neotropics are bounded by the Tropics of Cancer and Capricorn. Although these boundaries are artificial, they approximate biological boundaries limiting the distribution of tropical species outside the tropics and vice versa. In Mexico, the Tropic of Cancer follows the break between the arid and semi-arid climates to the north and the humid and semi-humid climates to the south (Toledo et al. 1997). The Caribbean buffers the West Indies and maintains a tropical climate north into southernmost Florida. Farther north, however, the vegetation must withstand rare freezing; consequently, many species simply are not found north of the Everglades. To the south, the Tropic of Capricorn runs through the Atacama Desert of northern Chile, a significant barrier to plant life. To the east, the Tropic is found at the latitude of the city of São Paulo, just south of the southern limit of the cerrado. The Atlantic coastal forest continues south to the end of the Serra do Mar at about 30 south (Thomas 2005). The flora of tropical America includes many plant groups that are exclusively or primarily found in the New World, such as the Bromeliaceae (Smith & Downs 1974, 1977, 1979) and Gunneraceae (Mora-Osejo et al. 2011). In addition, there are many families for which genera or groups of genera are mostly American, such as the Lecythidaceae (Prance & Mori 1979; Mori & Prance 1990), Mimosa-Fabaceae (Barneby 1991), and Miconieae-Melastomataceae (Michelangeli et al onward). Although there are exceptions (Pennington et al. 2004), in many cases (see Smith et al. 2004), therefore, a revision of a Neotropical group will functionally be a monograph of that group - this is even more so if the small number of non-tropical outliers are also treated. So, how many species are there in the Neotropics, and what are our prospects of making their descriptions available through the World Flora Online? Estimates of species diversity are fraught with uncertainty, but do provide a useful yardstick and, for the Neotropics, new information means a new estimate is needed. Earlier estimates (Prance 1977; Raven 1988; Thorne 2002) of the number of Angiosperm species on earth were approximately 250,000, with about 36 percent (90,000) found in tropical America (Forero & Mori 1995; Maas & Westra 1998; Raven 1988; Thomas 1999). Govaerts (2001, 2003) estimated the World s species diversity of seed plants at a much higher 420,000 species. Since The Plant List (2013) is based on an actual count of species, it offers a more detailed estimate of the World s plant species. It includes 951,140 species names of Angiosperms. Of these 304,419 are accepted species names and another 216,375 unresolved. If percent of the unresolved names are found to be accepted, the estimate of the number plant species in the World rises to 350, ,000. With the estimate for the World s species having increased by almost 100,000, it is likely that the estimate for the Neotropics should be higher also - if the Neotropical flora comprises a third of the World s flora, it would total at least 120,000 species. Flora Neotropica monographs - A Model for Digitization Like many botanical journals, Flora Neotropica monographs were published in print form over many years. The idea of a Flora Neotropica project was first envisioned in 1957 by an advisory committee of UNESCO and, in 1964, the Organization for Flora Neotropica was created at a meeting held at the Institute of Botany of São Paulo at the invitation of the Institute s director, Dr. Alcides R. Teixeira. It also led to the creation of a commission to stimulate and assist in the preparation and publication of all the naturally occurring fungi and plants of tropical America in a monograph series entitled Flora Neotropica. Flora Neotropica was based at The New York Botanical Garden and, since its inception, all costs of the monograph series have been assumed by the Garden. The series is reserved for manuscripts that, because of their length, cannot be published in their entirety in other journals (Forero & Mori 1995; Thomas 2005). Since the first Flora Neotropica monograph appeared in 1965, an additional 113 volumes have been published. Of these, 87 were on vascular plants (16 were on fungi, and 11 on bryophytes). These 87 monographs treat 16,730 scientific names Rodriguésia 66(4):

54 Flora Neotropica and WFO and recognize 8008 species, of which 2087 (26%) were described by the authors of the monographs. These numbers demonstrate that monographers are those best able to recognize and circumscribe the species in the groups they study - they also perceive rare and cryptic species passed over by non-specialists and those which can be critical for conservation planning (Marhold et al. 2013). In order to increase contributions of monographs, especially from botanists in the Neotropics, monograph submissions are accepted in Portuguese, Spanish and French. Flora Neotropica monographs are published through The New York Botanical Garden Press and have always been available for sale. So that costs of printed monographs is minimized, in recent years the New York Botanical Garden Press has reduced original print runs and is making the monographs available through Print-on-Demand publication. Currently, Flora Neotropica is available only as hardcopy or with subscription access to JSTOR (< jstor.org>), making them generally unavailable to those not affiliated with a university or those from developing countries. Even for those with access, the PDFs available are only of images and do not contain any embedded text, making them unsearchable. Hard Copy to Digital: Flora Neotropica monographs The goals of WFO include providing published modern descriptions of each species, online. Most of the descriptions of the World s plant species have been published in print form. What proportion of the descriptions of the World s species already exist digitally as searchable text, and what proportion only exist as hardcopy, or as digital page images? The WFO Consortium has estimated that no more than 160,000 species have digital descriptions, meaning that there are an estimated 190,000 species for which descriptions must be transformed from hardcopy to digital searchable descriptions - a large task. With efforts like the Biodiversity Heritage Library (< access to much of the earlier literature, especially that out of copyright, has been made available to the public online. Many of these publications contain protologues (original descriptions), that are often the only available description for a given species. BHL will be an excellent resource for the WFO. Access and re-use of recent literature with treatments and 985 synonymy based on current classifications using modern techniques (DNA analyses, etc.), however, is generally restricted by copyright. Organizations such as Plazi argue that taxonomic descriptions do not fit the definition of copyright and can be marked up and re-used (Agosti and Egloff 2009), but there is still some hesitation with how copyright laws in different countries will be applied to this type of re-purposing of publications. Increasingly, semantically-enhanced, borndigital publications are competing with the printonly (or print plus paywall) approach. Open access publishing models promoted by publishers like Pensoft will revolutionize publishing (Miller et al. 2012), but there remains a large amount of literature still in copyright and mostly available online as PDFs behind subscription services or individually available by download from the publisher for a set price per article. In light of the WFO and because of its importance botanically, we are using Flora Neotropica monographs as a test case for converting a print plus paywall publication to marked up text and providing free access to these descriptions online. Even for those with access to digital copies of Flora Neotropica, the PDFs available are typically only page images and contain only some automatically generated, but not corrected or marked up text, making them unreliably searchable both individually and across all volumes. Each volume of Flora Neotropica was published with different licensing agreements with scientists and artists: database of generic- and species-level descriptions has therefore been generated from images of each volume. To convert each volume of Flora Neotropica to text, PDFs of the images are sent through ABBYY FineReader Professional OCR (optical character recognition) software. This software also highlights potentially misspelled words or incorrect character conversion to make it easier to quickly proofread. A botanical dictionary of plant morphology terms and taxonomic names was loaded into the software to minimize the number of potential misspellings. Since all volumes remained fairly standard in font and formatting, issues with conversion to text have been consistent across volumes and the software has been trained to recognize reccurring problems. Project staff manually edit any OCR transcription errors or formatting issues discovered during the process. Once the text is cleaned up, the information is Rodriguésia 66(4):

55 986 Thomas, W.W. & Tulig, M. parsed and mapped to the DarwinCore standard, adopted by the Biodiversity Information Standards (TDWG) association (< Once complete, DarwinCore-Archive (DwC-A) files will be produced and shared via the NYBG Integrated Publishing Toolkit (IPT) for harvesting by the World Flora Online Consortium portal, Global Biodiversity Information Facility (GBIF), the Encyclopedia of Life (EOL), the Digital Public Library of America (DPLA) or any interested party. They will also be available on the NYBG website, with links to cited NY specimen records in the Virtual Herbarium. Conclusion To reach the stated goals of the WFO by 2020, digitizing hardcopy plant descriptions is critically important and must be embraced by all members of the Consortium. We have started with Flora Neotropica because of the biological importance of the series and because we can clarify the issues of copyright and licensing agreements. Acknowledgments We thank the Alfred P. Sloan Foundation and Google, Inc., for their valuable support of this project. References Agnarsson, I. & Kuntner, M Taxonomy in a changing world: seeking solutions for a science in crisis. Systematic Biology 56: DOI: / Agosti, D. & Egloff, W Taxonomic information exchange and copyright: the plazi approach. BMC Research Notes 2: 53. DOI: / Antonelli, A.; Zizka, A.; Silvestro, D.; Scharn, R.; Cascales-Miñana, B. & Bacon, C.D An engine for global plant diversity: highest evolutionary turnover and emigration in the American tropics. Frontiers in Genetics 6: 130. DOI: / fgene Barneby, R Sensitivae Censitae - A description of the genus Mimosa Linnaeus (Mimosaceae) in the new world. Memoirs of the New York Botanical Garden 65: Forero, E. & Mori, S.A The organization for Flora Neotropica. Brittonia 47: Maas, P.J.N. & Westra, L.Y.T Familias de plantas neotropicales. Koeltz Scientific Books. Koenigstein, Germany. 315p. Marhold, K.; Stuessy, T.; Agababian, M.; Agosti, D.; Alford, M.H.; Crespo, A.M.; Crisci, J.V.; Dorr, L.J.; Ferencová, Z.; Frodin, D.; Geltman, D.V.; Kilian, N.; Linder, H.P.; Lohmann, L.G.; Oberprieler, C.; Penev, L.; Smith, G.; Thomas, W.; Tulig, M.; Turland, N. & Zhang, X.-C The future of botanical monography: report from an international workshop, March 2012, Smolenice, Slovak Republic. Taxon 62: Michelangeli, F.A.; Almeda, F.; Goldenberg, R.; Judd, W.S.; Becquer-Granados, E.R. & Tulig, M onward. PBI Miconieae: a complete web-based monograph of the tribe Miconieae (Melastomataceae). The New York Botanical Garden, Bronx, NY. Available at < nybg.org/melastomataceae>. Access on 3 September Miller, J.; Dikow, T.; Agosti, D.; Sautter, G.; Catapano T.; Penev, L.; Zhang, Z.-Q.; Pentcheff, D.; Pyle, R.; Blum, S.; Parr, C.; Freeland, C.; Garnett, T.; Ford, L.S.; Muller, B.; Smith, L.; Strader, G.; Georgiev, T. & Bénichou, L From Taxonomic Literature to Cybertaxonomic Content. BMC Biology 10: 87. DOI: / Mora-Osejo, L.E.; Pabón-Mora, N. & González, F Gunneraceae. Flora Neotropica 109: Mori, S.A. & Prance, G.T Lecythidaceae - part II, The Zygomorphic-flowered new world genera (Couroupita, Corythophora, Bertholletia, Couratari, Eschweilera, & Lecythis). Flora Neotropica 21: Pennington, R.T. & Dick, C.W The role of immigrants in the assembly of the South American rainforest tree flora. Philosophical Transactions of the Royal Society B: Biological Sciences 359: Pimm, S.L.; Clinton, N.J.; Joppa, L.N.; Roberts, D.L. & Russell, G.J How many endangered species remain to be discovered in Brazil? Natureza e Conservação 8: DOI: / natcon Prance, G.T Floristic inventory of the tropics: where do we stand? Annals of the Missouri Botanical Garden 64: Prance, G.T. & Mori, S.A Lecythidaceae - part I, the actinomorphic-flowered new world Lecythidaceae (Asteranthos, Gustavia, Grias, Allantoma, & Cariniana). Flora Neotropica 21: Raven, P. H Tropical floristics tomorrow. -Taxon 37: The Plant List Version 1.1. Published on the Internet. Available at < org/>. Access on 1 September Smith, L.B. & Downs, R.J Pitcairnioideae (Bromeliaceae). Flora Neotropica 14: Smith, L.B. & Downs, R.J Tillandsioideae (Bromeliaceae). Flora Neotropica 14: Rodriguésia 66(4):

56 Flora Neotropica and WFO Smith, L.B. & Downs, R.J Bromelioideae (Bromeliaceae). Flora Neotropica 14: Smith, N.; Mori, S.A.; Henderson, A.; Stevenson, D.W. & Heald, S Flowering plants of the Neotropics. Princeton University Press, Princeton. 594p. Thomas, W.W Conservation and monographic research on the flora of Tropical America. Biodiversity and Conservation 8: Thomas, W.W Flora Neotropica - Monographs as inventories. In: Species Plantarum 250 years: Proceedings of the Species plantarum symposium held in Uppsala August 22-24, Acta Universitatis Upsaliensis, Symbolae Botanicae Upsalienses 33: Thorne, R.F How many species of seed plants are there? Taxon 51: Artigo recebido em 08/09/2015. Aceito para publicação em 16/10/2015. Rodriguésia 66(4):

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58 Rodriguésia 66(4): DOI: / The Catalogue of Vascular Plants of the Southern Cone and the Flora of Argentina: their contribution to the World Flora Fernando O. Zuloaga 1,2 & Manuel J. Belgrano 1 Abstract The checklist of the vascular plants of the Southern Cone presents updated information of 19,787 taxa and 44,943 synonyms, distributed in 2,679 genera and 318 families. This checklist was prepared, and its permanently updated, using the database Documenta Florae Australis, with the contribution of numerous researchers and institutions, and constitutes the bases for the Flora of Argentina, currently under preparation. Here we evaluate the current knowledge of vascular plants in the Southern Cone and provide a summary of the ongoing Flora of Argentina, and the significance of these projects for the World Flora on line and to the botanical studies in the region. Key words: Endemism, Flora, South America, Southern Cone. Resumen A listagem das plantas vasculares do Cone Sul apresenta táxons com sinônimos, distribuídos em gêneros e 318 famílias. Preparada e continuamene atualizada utilizando o banco de dados "Documenta florae Australis", com a contribuição de vários pesquisadores e instituições, esta lista constitui a base para a Flora da Argentina, atualmente em preparação. Aqui avaliamos o conhecimento atual das plantas vasculares no Cone Sul e fornecemos um resumo sobre o andamento da Flora da Argentina, além da importância destes projetos para a Flora do Mundo Online e para os estudos botânicos na região. Palavras clave: Cono Sur, Endemismo, Flora, Sudamérica. Introduction The Catálogo de las Plantas Vasculares del Cono Sur presents a critical, commented list of the species and infraspecific taxa that occur in the extreme south of the Americas, comprising the totality of Argentina, southern Brazil (Paraná, Santa Catarina and Rio Grande do Sul states), Chile, Paraguay and Uruguay, including the majority of the subtropical and temperate to cold-temperate areas approximately south of 20ºS. The land area is equivalent to 4,708,617 km 2, representing 26% of the total land area of South America (Fig. 1). This project was preceeded by the Catálogo de Plantas Vasculares de la República Argentina (Zuloaga et al. 1994; Zuloaga & Morrone 1996, 1999), and started in 2001 at the Instituto de Botánica Darwinion (SI), Buenos Aires, Argentina, in collaboration with the Missouri Botanical Garden (MO), the Herbarium of the Universidad de Concepción (CONC), Chile, Herbarium of the Facultad de Agronomía (MVFA), Montevideo, Uruguay, the Herbarium of the Facultad de Ciencias Químicas (FCQ), Asunción, Paraguay and the Botany Departament of the Universidade Federal do Rio Grande do Sul (ICN), Porto Alegre, Brazil, involving numerous contributors from different botanical institutions. The resulting critical list of vascular plants from the Southern Cone, or Cono Sur in Spanish, was published in a printed version (Zuloaga et al. 2008), while the digital version is continuously updated and available for public searches through the Instituto Darwinion s webpage (< ar/proyectos/floraargentina/fa.htm>). During the year 2008, using as a base the existing Catálogo del Cono Sur and other 1 Instituto de Botánica Darwinion, Labardén 200, Casilla de Correo 22 (B1642HYD), San Isidro, Buenos Aires, Argentina. 2 Author for correspondence: fzuloaga@darwin.edu.ar

59 990 Zuloaga, F.O. & Belgrano, M.J. This contribution updates the floristic information found in the Catálogo de las plantas vasculares del Cono Sur (Zuloaga et al. 2008), particularly concerning data from Argentina, as the ongoing preparation of the Flora of Argentina, directed by A. M. Anton (CORD) & F. O. Zuloaga (SI), including analyses and floristic comments that will lead to future systematic works in the region, and to the integration of these projects into the World Flora online by Figure 1 Study area, the Southern Cone of South America (Argentina, Southern Brazil, Chile, Paraguay, and Uruguay). previous projects carried out in Argentina (Floras of Buenos Aires, Chaqueña, Entre Ríos, Jujuy, Patagonia, San Juan) and also the Proflora project (Hunziker ; Anton & Zuloaga ), the production and publication of a new Flora vascular de la República Argentina (from now on Flora of Argentina) begun. This work was planned to comprise 20 volumes and is currently underway, with an electronic version available at (< The web version incorporates additional information such as distribution maps, photographs of live plants in the field and complete list of examined specimens. To date, the published volumes of Flora of Argentina include Poaceae (Zuloaga et al. 2012), Asteraceae (parts 1, tribes Anthemideae to Gnaphalieae and 3, tribes Senecioneae and Vernonieae) (Zuloaga et al. 2014a, b), Brassicaceae (Anton & Zuloaga 2012a), Solanaceae (Zuloaga et al. 2013) and Verbenaceae (Anton & Zuloaga 2012b); the volumes regarding Basal Dicotyledoneae (published in September 2015) and Asteraceae (volume 7(2) is in press). Methodology The present work used the database Documenta Florae Australis (from now on referred as DFA), developed and maintained at the Instituto Darwinion during the last 20 years. Such relational database integrates scientific name of the accepted taxa with their synonyms, indicates original publication data, presence and distribution in each country of the South American Southern Cone (discriminating by regions, provinces or states, departments or municipalities), status (if the taxon is endemic, native or introduced), habit and lifecycle, including also bibliography and vouchers, line drawings and field photographs. A complete inventory of the vascular plants cited for the Southern Cone can be obtained from DFA. As soon as new taxonomic and systematic treatments are published, DFA data is updated and its records are increased daily by the contribution of numerous registered users. Also, more than 15 institutions are digitalising their collections and these are continuously uploaded into this system. Today, DFA has 66,784 scientific names recorded for the Southern Cone, of which 19,787 are accepted and the remaining 44,943 are synonyms. The number of herbarium specimens linked to the system is 442,155, the bibliography records amount to 10,005, while 12,554 names of taxonomists, collectors or botanical authorities, close to 9,000 line art illustrations and 35,000 pictures of plants in the field, amongst other items, are included in DFA. This database is open and available for consultation and represents a valuable resource for improving knowledge, management and conservation of the flora of Southern South America. Angiosperm families from the Southern Cone were organized according to the Angiosperm Phylogeny Group available at (< org/mobot/research/apweb/>), with a few punctual exceptions where the specialists preferred to maintain the traditional classification, such as the Chenopodiaceae, that remains separate from the Rodriguésia 66(4):

60 Vascular Plants of the Southern Cone 991 Amaranthaceae. For the Ferns and allies the system of R. M. Tryon & A. F. Tryon (1982) was followed, with subsequent modifications and updates (in the classification of families and genera). Distributional analysis of taxa followed the Biogeographical scheme proposed by Josse et al. (2003) (Fig. 2) with slight modifications. Results and Discussion A figure of 19,787 taxa, distribuited in 18,139 species, 2,679 genera and 318 families is currently recorded for the Southern Cone, 7,787 species are endemic to the region (Tab. 1). The Angiosperms comprise 95.40% of the vascular flora (70.68% Dicots and 24,72% Monocots) including the majority of the endemic species from the Southern Cone (7,630 species, or 85.08% of the total). The significant difference found between the present results and those published in the Catálogo del Cono Sur (Zuloaga et al. 2008: xxxviii), considering that a short period of time has elapsed between those milestones, points out the importance of keeping the floristic and taxonomic information available online and permanently updated. The total of families in 2008 was 308, with 2,586 genera and 17,693 species, of which 7,691 were endemic. The increase in family numbers was 10 while the genera increased by 93, with 446 additional species; of these, 96 more were endemic. There are several possible causes to explain this variation: description of new taxa, record of new occurrences for the area or adjustment of taxonomic circumscription at all levels, often as a result of new taxonomic and phylogenetic research. Therefore the species number accepted within a certain genus may decrease when new synonymies are proposed, as seen in genus Phyla L. (O Leary & Múlgura 2012) or it may increase through the description of new species, as was the case with Chusquea Kunth, through the description of Chusquea egluma Guerreiro & Rúgolo and Chusquea floribunda Guerreiro & Figure 2 Biogeographic regions of the Southern Cone of South America. Rúgolo in 2013 (Guerreiro & Rúgolo de Agrasar 2013). The number of genera may increase through segregation of a group of species to another genus, as seen in 2014 for Panicum validum Mez that was split from Panicum L. and placed within the new genus Osvaldoa J.R. Grande (Grande Allende 2014), causing an increase in the number of genera in the Poaceae. The number of families may also decrease, as it is the case of the Nolanaceae and Sclerophylacaceae, nowadays considered as part of the Solanaceae. Table 1 Southern Cone, summary of the taxa included in the catalogue. Group Families Genera Species % Species Endemics % Endemics Pteridophyta , ,91 Gymnospermae , ,67 Dicotyledoneae , ,76 Monocotyledoneae , ,32 Total Rodriguésia 66(4):

61 992 Zuloaga, F.O. & Belgrano, M.J. Table 2 Argentina, summary of the taxa recorded for the country. Group Families Genera Species % Species Endemics %Endemics Pteridophyta , ,10 Gymnospermae ,26 1 3,85 Dicotyledoneae , ,31 Monocotyledoneae , ,37 Total For the Flora of Argentina, there is a current record of 10,944 taxa distributed in 10,006 species, 2,090 genera and 283 families, with a total of 1,749 endemic species (Tab. 2) representing 17.48% of the country s flora. Plant diversity at family level The largest family in terms of species number in the Southern Cone is Asteraceae (2,523 species), followed by Poaceae (1,535), Fabaceae (1,365), Orchidaceae (1,042), Solanaceae (546), Cyperaceae (485), Euphorbiaceae (447), Malvaceae (408), Cactaceae (406) and Apocynaceae (333). These families comprise 50% of the species found in the area. Contrastingly, 202 families are represented by 15 or less species, of which 59 have a single species in the area. A complete list of the families verified for the Southern Cone is presented in Table 3, indicating the genus, species and endemic species number for each, together with the percentage of endemic species in relation to the total number of species in the family. Argentina has a similar representation at family level to what is found in the Southern Cone (55.16% of the species from the Southern Cone are present in Argentina); the top ten families are Asteraceae (1,502 species), Poaceae (1,170), Fabaceae (760), Solanaceae (322), Cyperaceae (314), Orchidaceae (281), Cactaceae (240), Malvaceae (233), Brassicaceae (229) and Euphorbiaceae (218); 202 families with 15 or less species among which 68 have a single representative in the country (Tab. 4). Plant diversity at genus level Among the 2,679 genera found in the Southern Cone, the largest one is Senecio L. (Asteraceae, 415 species), followed by Solanum L. (Solanaceae, 213), Baccharis L. (Asteraceae, 208), Adesmia DC. (Fabaceae, 199), Mimosa L. (Fabaceae, 169), Croton L. (Euphorbiaceae, 126), Oxalis L. (Oxalidaceae, 124), Eugenia L. (Myrtaceae, 122), Carex L. (Cyperaceae, 110), Paspalum L. (Poaceae, 109), Viola L. (Violaceae, 107) and Sisyrinchium L. (Iridaceae, 103). The remaining genera have 100 or less species recorded for the region. Particularly, for the Flora of Argentina, the richest genus in number of species is Senecio (with 260 species), followed by Solanum (126), Adesmia (102), Baccharis (101), Paspalum (77), Oxalis (71), Astragalus L. (69), Mimosa (53), Viola (53), Valeriana L. (48), Polygala L. (44) and Calceolaria L. (41). In the other side of the spectrum, 949 genera (45.40 % of the total) are represented by a single species in Argentina. Endemism A species or taxon is considered endemic in the present work when its distribution is known only for the area of this project. Thus, for the Southern Cone, there are eight endemic families (3.25% of the total number of families): Francoaceae, Gomortegaceae and Lactoridaceae, known only in Chile, Halophytaceae, exclusive from Argentina and Aextoxicaceae, Misodendraceae, Philesiaceae and Tribelaceae, shared between Argentina and Chile. The number of endemic genera for the region is 241 (9% of the total genera) and the family with the largest number of endemic genera is Asteraceae, with 46 endemic genera, followed by Amaryllidaceae (16), Brassicaceae (13), Apiaceae (11), Cactaceae (10), Solanaceae (9), Poaceae (9), Alliaceae (8) and Fabaceae (6). Table 5 details the endemic genera with species number and their presence in each country. The total of endemic species in the Southern Cone is 7,787, representing 42,93 % of the total plant species found in the region. Meanwhile, for the Flora of Argentina, the only endemic family is the monotypic Rodriguésia 66(4):

62 Vascular Plants of the Southern Cone 993 Table 3 Southern Cone, number of families, genera, species, and endemic species. Group Family Genera Species Endemics % Endemism Dicotyledoneae Acanthaceae ,6 Dicotyledoneae Aceraceae 1 2 Dicotyledoneae Achariaceae 1 1 Dicotyledoneae Achatocarpaceae ,0 Dicotyledoneae Adoxaceae 2 3 Dicotyledoneae Aextoxicaceae ,0 Monocotyledoneae Agapanthaceae 1 1 Monocotyledoneae Agavaceae 2 3 Dicotyledoneae Aizoaceae ,0 Monocotyledoneae Alismataceae ,7 Monocotyledoneae Alstroemeriaceae ,0 Dicotyledoneae Amaranthaceae ,7 Monocotyledoneae Amaryllidaceae ,0 Dicotyledoneae Anacampserotaceae ,0 Dicotyledoneae Anacardiaceae ,7 Pteridophyta Anemiaceae ,1 Dicotyledoneae Annonaceae ,4 Monocotyledoneae Anthericaceae ,5 Dicotyledoneae Apiaceae ,1 Dicotyledoneae Apocynaceae ,6 Monocotyledoneae Aponogetonaceae 1 1 Dicotyledoneae Aquifoliaceae ,7 Monocotyledoneae Araceae ,5 Dicotyledoneae Araliaceae ,4 Gymnospermae Araucariaceae ,0 Monocotyledoneae Arecaceae ,0 Dicotyledoneae Aristolochiaceae ,3 Monocotyledoneae Asparagaceae 1 5 Monocotyledoneae Asphodelaceae 1 1 Pteridophyta Aspleniaceae ,8 Monocotyledoneae Asteliaceae ,0 Dicotyledoneae Asteraceae ,2 Pteridophyta Athyriaceae 3 19 Dicotyledoneae Avicenniaceae 1 1 Pteridophyta Azollaceae 1 2 Dicotyledoneae Balanophoraceae 4 5 Dicotyledoneae Balsaminaceae 1 3 Dicotyledoneae Basellaceae 2 4 Dicotyledoneae Begoniaceae ,5 Dicotyledoneae Berberidaceae ,2 Dicotyledoneae Berberidopsidaceae ,0 Rodriguésia 66(4):

63 994 Zuloaga, F.O. & Belgrano, M.J. Group Family Genera Species Endemics % Endemism Dicotyledoneae Betulaceae 2 3 Dicotyledoneae Bignoniaceae ,3 Dicotyledoneae Bixaceae 1 1 Pteridophyta Blechnaceae ,6 Dicotyledoneae Bombacaceae 5 12 Dicotyledoneae Boraginaceae ,5 Dicotyledoneae Brassicaceae ,6 Monocotyledoneae Bromeliaceae ,7 Monocotyledoneae Burmanniaceae 6 9 Dicotyledoneae Burseraceae ,3 Dicotyledoneae Cabombaceae 1 2 Dicotyledoneae Cactaceae ,4 Dicotyledoneae Calceolariaceae ,3 Dicotyledoneae Callitrichaceae ,4 Dicotyledoneae Calyceraceae ,6 Dicotyledoneae Campanulaceae ,3 Dicotyledoneae Canellaceae ,0 Dicotyledoneae Cannabaceae 1 2 Monocotyledoneae Cannaceae ,0 Dicotyledoneae Capparaceae ,0 Dicotyledoneae Caprifoliaceae 2 2 Dicotyledoneae Cardiopteridaceae ,7 Dicotyledoneae Caricaceae ,7 Dicotyledoneae Caryocaraceae 1 1 Dicotyledoneae Caryophyllaceae ,7 Dicotyledoneae Cecropiaceae 3 4 Dicotyledoneae Celastraceae ,2 Dicotyledoneae Celtidaceae 2 5 Monocotyledoneae Centrolepidaceae ,0 Dicotyledoneae Ceratophyllaceae 1 2 Dicotyledoneae Cervantesiaceae ,3 Dicotyledoneae Chenopodiaceae ,0 Dicotyledoneae Chloranthaceae 1 1 Dicotyledoneae Chrysobalanaceae 4 6 Dicotyledoneae Cistaceae ,0 Dicotyledoneae Clethraceae ,0 Dicotyledoneae Clusiaceae 5 7 Dicotyledoneae Cochlospermaceae 1 2 Dicotyledoneae Combretaceae ,0 Monocotyledoneae Commelinaceae ,9 Dicotyledoneae Connaraceae 2 2 Dicotyledoneae Convolvulaceae ,9 Dicotyledoneae Coriariaceae ,0 Monocotyledoneae Corsiaceae ,0 Rodriguésia 66(4):

64 Vascular Plants of the Southern Cone 995 Group Family Genera Species Endemics % Endemism Monocotyledoneae Costaceae 1 3 Dicotyledoneae Crassulaceae ,4 Dicotyledoneae Cucurbitaceae ,3 Dicotyledoneae Cunoniaceae ,2 Gymnospermae Cupressaceae ,0 Pteridophyta Cyatheaceae 3 17 Monocotyledoneae Cyclanthaceae ,0 Monocotyledoneae Cymodoceaceae 1 1 Monocotyledoneae Cyperaceae ,4 Pteridophyta Dennstaedtiaceae ,1 Dicotyledoneae Desfontainiaceae 1 1 Pteridophyta Dicksoniaceae ,0 Pteridophyta Didymochlaenaceae 1 1 Dicotyledoneae Dilleniaceae 4 11 Monocotyledoneae Dioscoreaceae ,4 Dicotyledoneae Dipsacaceae 3 5 Dicotyledoneae Donatiaceae ,0 Dicotyledoneae Droseraceae ,7 Pteridophyta Dryopteridaceae ,7 Dicotyledoneae Ebenaceae ,3 Dicotyledoneae Elaeagnaceae 1 1 Dicotyledoneae Elaeocarpaceae ,0 Dicotyledoneae Elatinaceae ,0 Dicotyledoneae Empetraceae 1 1 Dicotyledoneae Epacridaceae ,0 Gymnospermae Ephedraceae ,7 Pteridophyta Equisetaceae 1 2 Dicotyledoneae Eremolepidaceae ,7 Dicotyledoneae Ericaceae ,8 Monocotyledoneae Eriocaulaceae ,0 Dicotyledoneae Erythroxylaceae ,1 Dicotyledoneae Escalloniaceae ,1 Dicotyledoneae Eucryphiaceae ,0 Dicotyledoneae Euphorbiaceae ,9 Dicotyledoneae Fabaceae ,7 Dicotyledoneae Francoaceae ,0 Dicotyledoneae Frankeniaceae ,0 Dicotyledoneae Fumariaceae 1 7 Dicotyledoneae Gentianaceae ,8 Dicotyledoneae Geraniaceae ,5 Dicotyledoneae Gesneriaceae ,5 Pteridophyta Gleicheniaceae ,2 Dicotyledoneae Gomortegaceae ,0 Rodriguésia 66(4):

65 996 Zuloaga, F.O. & Belgrano, M.J. Group Family Genera Species Endemics % Endemism Dicotyledoneae Goodeniaceae 2 2 Dicotyledoneae Griseliniaceae ,0 Dicotyledoneae Grossulariaceae ,3 Dicotyledoneae Gunneraceae ,6 Dicotyledoneae Halophytaceae ,0 Dicotyledoneae Haloragaceae ,3 Monocotyledoneae Heliconiaceae 1 6 Monocotyledoneae Hemerocallidaceae 1 1 Monocotyledoneae Herreriaceae ,3 Dicotyledoneae Hippuridaceae 1 1 Dicotyledoneae Humiriaceae 1 1 Monocotyledoneae Hyacinthaceae ,0 Dicotyledoneae Hydnoraceae 1 2 Dicotyledoneae Hydrangeaceae ,0 Monocotyledoneae Hydrocharitaceae 5 7 Dicotyledoneae Hydroleaceae 1 2 Pteridophyta Hymenophyllaceae ,8 Dicotyledoneae Hypericaceae ,0 Monocotyledoneae Hypoxidaceae ,0 Dicotyledoneae Icacinaceae ,0 Monocotyledoneae Iridaceae ,3 Pteridophyta Isoëtaceae ,0 Dicotyledoneae Juglandaceae 2 2 Monocotyledoneae Juncaceae ,6 Monocotyledoneae Juncaginaceae ,0 Dicotyledoneae Krameriaceae ,3 Dicotyledoneae Lacistemataceae 1 3 Dicotyledoneae Lactoridaceae ,0 Dicotyledoneae Lamiaceae ,5 Dicotyledoneae Lardizabalaceae ,0 Dicotyledoneae Lauraceae ,8 Monocotyledoneae Laxmanniaceae ,0 Dicotyledoneae Lecythidaceae 1 1 Dicotyledoneae Ledocarpaceae ,7 Monocotyledoneae Lemnaceae 5 11 Dicotyledoneae Lentibulariaceae ,0 Monocotyledoneae Limnocharitaceae 2 6 Dicotyledoneae Linaceae ,4 Dicotyledoneae Linderniaceae 2 3 Pteridophyta Lindsaeaceae 1 10 Dicotyledoneae Loasaceae ,0 Dicotyledoneae Loganiaceae ,4 Pteridophyta Lomariopsidaceae 2 2 Rodriguésia 66(4):

66 Vascular Plants of the Southern Cone 997 Group Family Genera Species Endemics % Endemism Dicotyledoneae Loranthaceae ,0 Monocotyledoneae Luzuriagaceae ,0 Pteridophyta Lycopodiaceae ,1 Pteridophyta Lygodiaceae 1 2 Dicotyledoneae Lythraceae ,0 Dicotyledoneae Malesherbiaceae ,8 Dicotyledoneae Malpighiaceae ,8 Dicotyledoneae Malvaceae ,4 Monocotyledoneae Marantaceae ,4 Pteridophyta Marattiaceae 3 5 Dicotyledoneae Marcgraviaceae 2 2 Pteridophyta Marsileaceae ,0 Dicotyledoneae Martyniaceae ,0 Monocotyledoneae Mayacaceae 1 2 Dicotyledoneae Melastomataceae ,1 Dicotyledoneae Meliaceae ,1 Dicotyledoneae Menispermaceae ,7 Dicotyledoneae Menyanthaceae 1 2 Dicotyledoneae Microteaceae 1 5 Dicotyledoneae Misodendraceae ,0 Dicotyledoneae Molluginaceae 2 3 Dicotyledoneae Monimiaceae ,8 Dicotyledoneae Montiaceae ,5 Dicotyledoneae Moraceae ,2 Dicotyledoneae Muntingiaceae 1 1 Dicotyledoneae Myoporaceae 1 1 Dicotyledoneae Myricaceae 1 2 Dicotyledoneae Myristicaceae 1 1 Dicotyledoneae Myrsinaceae ,7 Dicotyledoneae Myrtaceae ,4 Monocotyledoneae Najadaceae 1 3 Dicotyledoneae Nanodeaceae ,0 Pteridophyta Nephrolepidaceae 1 6 Dicotyledoneae Nolanaceae ,0 Dicotyledoneae Nothofagaceae ,0 Dicotyledoneae Nyctaginaceae ,9 Dicotyledoneae Nymphaeaceae 3 10 Dicotyledoneae Ochnaceae 2 7 Dicotyledoneae Olacaceae 1 2 Dicotyledoneae Oleaceae ,8 Pteridophyta Oleandraceae ,0 Dicotyledoneae Onagraceae ,6 Pteridophyta Ophioglossaceae ,7 Rodriguésia 66(4):

67 998 Zuloaga, F.O. & Belgrano, M.J. Group Family Genera Species Endemics % Endemism Dicotyledoneae Opiliaceae 1 2 Monocotyledoneae Orchidaceae ,2 Dicotyledoneae Orobanchaceae ,7 Pteridophyta Osmundaceae 2 3 Dicotyledoneae Oxalidaceae ,9 Dicotyledoneae Papaveraceae ,3 Pteridophyta Parkeriaceae 1 1 Dicotyledoneae Parnassiaceae 1 1 Dicotyledoneae Passifloraceae ,0 Dicotyledoneae Pentaphylacaceae 1 2 Monocotyledoneae Philesiaceae ,0 Dicotyledoneae Phrymaceae ,7 Dicotyledoneae Phyllanthaceae ,5 Dicotyledoneae Phytolaccaceae ,3 Dicotyledoneae Picrodendraceae 1 1 Gymnospermae Pinaceae 2 4 Dicotyledoneae Piperaceae ,0 Dicotyledoneae Plantaginaceae ,9 Dicotyledoneae Plumbaginaceae ,7 Monocotyledoneae Poaceae ,3 Gymnospermae Podocarpaceae ,5 Dicotyledoneae Podostemaceae ,1 Dicotyledoneae Polemoniaceae ,8 Dicotyledoneae Polygalaceae ,2 Dicotyledoneae Polygonaceae ,9 Pteridophyta Polypodiaceae ,7 Monocotyledoneae Pontederiaceae 3 10 Dicotyledoneae Portulacaceae ,8 Monocotyledoneae Potamogetonaceae ,7 Dicotyledoneae Primulaceae ,3 Dicotyledoneae Proteaceae ,9 Pteridophyta Psilotaceae 1 1 Pteridophyta Pteridaceae ,3 Dicotyledoneae Quiinaceae 1 1 Dicotyledoneae Quillajaceae ,0 Dicotyledoneae Rafflesiaceae ,0 Dicotyledoneae Ranunculaceae ,7 Dicotyledoneae Resedaceae 1 4 Monocotyledoneae Restionaceae ,0 Dicotyledoneae Rhamnaceae ,3 Dicotyledoneae Rhizophoraceae 1 1 Dicotyledoneae Rosaceae ,8 Dicotyledoneae Rubiaceae ,7 Rodriguésia 66(4):

68 Vascular Plants of the Southern Cone 999 Group Family Genera Species Endemics % Endemism Monocotyledoneae Ruppiaceae 1 3 Monocotyledoneae Ruscaceae 1 1 Dicotyledoneae Rutaceae ,3 Dicotyledoneae Sabiaceae 1 1 Pteridophyta Saccolomataceae 1 3 Dicotyledoneae Salicaceae ,8 Pteridophyta Salviniaceae 1 4 Dicotyledoneae Samolaceae ,0 Dicotyledoneae Santalaceae ,0 Dicotyledoneae Sapindaceae ,4 Dicotyledoneae Sapotaceae ,7 Dicotyledoneae Saxifragaceae ,1 Pteridophyta Schizaeaceae 2 4 Dicotyledoneae Schlegeliaceae ,0 Dicotyledoneae Schoepfiaceae ,7 Dicotyledoneae Scrophulariaceae ,3 Pteridophyta Selaginellaceae ,1 Dicotyledoneae Simaroubaceae ,1 Dicotyledoneae Siparunaceae 1 1 Monocotyledoneae Smilacaceae ,3 Dicotyledoneae Solanaceae ,8 Dicotyledoneae Sphenocleaceae 1 1 Dicotyledoneae Sterculiaceae ,2 Dicotyledoneae Stylidiaceae ,0 Dicotyledoneae Styracaceae 1 9 Dicotyledoneae Symplocaceae ,8 Dicotyledoneae Talinaceae ,0 Dicotyledoneae Tamaricaceae 1 1 Monocotyledoneae Tecophilaeaceae ,9 Pteridophyta Tectariaceae 1 3 Dicotyledoneae Tetrachondraceae ,0 Dicotyledoneae Theaceae 2 2 Pteridophyta Thelypteridaceae ,0 Dicotyledoneae Theophrastaceae 1 1 Dicotyledoneae Thesiaceae 1 1 Dicotyledoneae Thymelaeaceae ,5 Dicotyledoneae Tiliaceae ,7 Dicotyledoneae Tribelaceae ,0 Dicotyledoneae Trigoniaceae 1 2 Monocotyledoneae Triuridaceae 2 2 Dicotyledoneae Tropaeolaceae ,8 Dicotyledoneae Turneraceae ,4 Monocotyledoneae Typhaceae 1 4 Rodriguésia 66(4):

69 1000 Zuloaga, F.O. & Belgrano, M.J. Group Family Genera Species Endemics % Endemism Dicotyledoneae Ulmaceae 1 1 Dicotyledoneae Urticaceae ,5 Dicotyledoneae Valerianaceae ,1 Monocotyledoneae Velloziaceae ,7 Dicotyledoneae Verbenaceae ,3 Dicotyledoneae Violaceae ,7 Dicotyledoneae Viscaceae ,9 Dicotyledoneae Vitaceae 3 16 Pteridophyta Vittariaceae 5 8 Dicotyledoneae Vivianiaceae ,0 Dicotyledoneae Vochysiaceae ,7 Dicotyledoneae Winteraceae ,0 Pteridophyta Woodsiaceae 2 3 Dicotyledoneae Ximeniaceae 1 2 Monocotyledoneae Xyridaceae ,7 Monocotyledoneae Zannichelliaceae 1 1 Monocotyledoneae Zingiberaceae 1 1 Monocotyledoneae Zosteraceae ,0 Dicotyledoneae Zygophyllaceae ,0 Total Table 4 Argentina, number of families, genera, species, and endemic species. Group Family Genera Species Endemics %Endemics Dicotyledoneae Acanthaceae ,8 Dicotyledoneae Aceraceae 1 2 Dicotyledoneae Achatocarpaceae 1 2 Dicotyledoneae Adoxaceae 2 3 Dicotyledoneae Aextoxicaceae 1 1 Monocotyledoneae Agapanthaceae 1 1 Monocotyledoneae Agavaceae 2 3 Dicotyledoneae Aizoaceae ,0 Monocotyledoneae Alismataceae 4 15 Monocotyledoneae Alstroemeriaceae ,7 Dicotyledoneae Amaranthaceae ,3 Monocotyledoneae Amaryllidaceae ,5 Dicotyledoneae Anacampserotaceae ,0 Dicotyledoneae Anacardiaceae ,4 Pteridophyta Anemiaceae 1 7 Dicotyledoneae Annonaceae 2 5 Monocotyledoneae Anthericaceae ,0 Dicotyledoneae Apiaceae ,7 Rodriguésia 66(4):

70 Vascular Plants of the Southern Cone 1001 Group Family Genera Species Endemics %Endemics Dicotyledoneae Apocynaceae ,2 Dicotyledoneae Aquifoliaceae 1 6 Monocotyledoneae Araceae Dicotyledoneae Araliaceae 6 8 Gymnospermae Araucariaceae 1 2 Monocotyledoneae Arecaceae ,3 Dicotyledoneae Aristolochiaceae ,8 Monocotyledoneae Asparagaceae 1 5 Monocotyledoneae Asphodelaceae 1 1 Pteridophyta Aspleniaceae ,4 Monocotyledoneae Asteliaceae 1 1 Dicotyledoneae Asteraceae ,8 Pteridophyta Athyriaceae 3 8 Pteridophyta Azollaceae 1 2 Dicotyledoneae Balanophoraceae 3 4 Dicotyledoneae Balsaminaceae 1 2 Dicotyledoneae Basellaceae 2 4 Dicotyledoneae Begoniaceae ,5 Dicotyledoneae Berberidaceae ,0 Dicotyledoneae Betulaceae 2 3 Dicotyledoneae Bignoniaceae ,8 Dicotyledoneae Bixaceae 1 1 Pteridophyta Blechnaceae 1 22 Dicotyledoneae Bombacaceae 3 5 Dicotyledoneae Boraginaceae ,1 Dicotyledoneae Brassicaceae ,6 Monocotyledoneae Bromeliaceae ,1 Monocotyledoneae Burmanniaceae 3 5 Dicotyledoneae Cabombaceae 1 1 Dicotyledoneae Cactaceae ,8 Dicotyledoneae Calceolariaceae ,4 Dicotyledoneae Callitrichaceae 1 8 Dicotyledoneae Calyceraceae ,3 Dicotyledoneae Campanulaceae ,7 Dicotyledoneae Cannabaceae 1 2 Monocotyledoneae Cannaceae ,0 Dicotyledoneae Capparaceae Dicotyledoneae Caprifoliaceae 2 2 Dicotyledoneae Cardiopteridaceae 1 3 Dicotyledoneae Caricaceae 2 5 Dicotyledoneae Caryophyllaceae ,0 Dicotyledoneae Cecropiaceae 1 1 Dicotyledoneae Celastraceae ,8 Rodriguésia 66(4):

71 1002 Zuloaga, F.O. & Belgrano, M.J. Group Family Genera Species Endemics %Endemics Dicotyledoneae Celtidaceae 2 5 Monocotyledoneae Centrolepidaceae 1 1 Dicotyledoneae Ceratophyllaceae 1 2 Dicotyledoneae Cervantesiaceae 2 3 Dicotyledoneae Chenopodiaceae ,2 Dicotyledoneae Cistaceae 1 1 Dicotyledoneae Clethraceae 1 1 Dicotyledoneae Clusiaceae 2 2 Dicotyledoneae Cochlospermaceae 1 1 Dicotyledoneae Combretaceae 2 6 Monocotyledoneae Commelinaceae 8 25 Dicotyledoneae Convolvulaceae ,6 Dicotyledoneae Coriariaceae 1 1 Monocotyledoneae Corsiaceae 1 1 Monocotyledoneae Costaceae 1 1 Dicotyledoneae Crassulaceae ,1 Dicotyledoneae Cucurbitaceae ,8 Dicotyledoneae Cunoniaceae 3 4 Gymnospermae Cupressaceae 4 4 Pteridophyta Cyatheaceae 2 4 Monocotyledoneae Cyperaceae ,4 Pteridophyta Dennstaedtiaceae 3 7 Dicotyledoneae Desfontainiaceae 1 1 Pteridophyta Dicksoniaceae 2 2 Pteridophyta Didymochlaenaceae 1 1 Dicotyledoneae Dilleniaceae 1 1 Monocotyledoneae Dioscoreaceae ,6 Dicotyledoneae Dipsacaceae 3 4 Dicotyledoneae Donatiaceae 1 1 Dicotyledoneae Droseraceae 1 3 Pteridophyta Dryopteridaceae ,9 Dicotyledoneae Ebenaceae 1 1 Dicotyledoneae Elaeagnaceae 1 1 Dicotyledoneae Elaeocarpaceae 4 4 Dicotyledoneae Elatinaceae ,0 Dicotyledoneae Empetraceae 1 1 Dicotyledoneae Epacridaceae 1 1 Gymnospermae Ephedraceae ,0 Pteridophyta Equisetaceae 1 2 Dicotyledoneae Eremolepidaceae 1 1 Dicotyledoneae Ericaceae 5 16 Monocotyledoneae Eriocaulaceae ,5 Dicotyledoneae Erythroxylaceae 1 5 Rodriguésia 66(4):

72 Vascular Plants of the Southern Cone 1003 Group Family Genera Species Endemics %Endemics Dicotyledoneae Escalloniaceae ,3 Dicotyledoneae Eucryphiaceae 1 1 Dicotyledoneae Euphorbiaceae ,8 Dicotyledoneae Fabaceae ,1 Dicotyledoneae Frankeniaceae ,9 Dicotyledoneae Fumariaceae 1 6 Dicotyledoneae Gentianaceae ,1 Dicotyledoneae Geraniaceae ,4 Dicotyledoneae Gesneriaceae 5 11 Pteridophyta Gleicheniaceae 2 3 Dicotyledoneae Griseliniaceae 1 2 Dicotyledoneae Grossulariaceae 1 6 Dicotyledoneae Gunneraceae 1 6 Dicotyledoneae Halophytaceae ,0 Dicotyledoneae Haloragaceae 1 2 Monocotyledoneae Heliconiaceae 1 3 Monocotyledoneae Herreriaceae 2 4 Dicotyledoneae Hippuridaceae 1 1 Monocotyledoneae Hyacinthaceae 1 3 Dicotyledoneae Hydnoraceae 1 2 Dicotyledoneae Hydrangeaceae 1 1 Monocotyledoneae Hydrocharitaceae 5 6 Dicotyledoneae Hydroleaceae 1 2 Pteridophyta Hymenophyllaceae ,4 Dicotyledoneae Hypericaceae 1 14 Monocotyledoneae Hypoxidaceae ,0 Monocotyledoneae Iridaceae ,3 Pteridophyta Isoëtaceae ,1 Dicotyledoneae Juglandaceae 2 2 Monocotyledoneae Juncaceae ,6 Monocotyledoneae Juncaginaceae 2 5 Dicotyledoneae Krameriaceae 1 1 Dicotyledoneae Lacistemataceae 1 2 Dicotyledoneae Lamiaceae ,7 Dicotyledoneae Lardizabalaceae 1 1 Dicotyledoneae Lauraceae 5 14 Monocotyledoneae Laxmanniaceae ,0 Dicotyledoneae Ledocarpaceae ,0 Monocotyledoneae Lemnaceae 4 10 Dicotyledoneae Lentibulariaceae 2 16 Monocotyledoneae Limnocharitaceae 2 5 Dicotyledoneae Linaceae 2 10 Rodriguésia 66(4):

73 1004 Zuloaga, F.O. & Belgrano, M.J. Group Family Genera Species Endemics %Endemics Dicotyledoneae Linderniaceae 1 1 Pteridophyta Lindsaeaceae 1 2 Dicotyledoneae Loasaceae ,2 Dicotyledoneae Loganiaceae 2 10 Dicotyledoneae Loranthaceae 5 8 Monocotyledoneae Luzuriagaceae 1 2 Pteridophyta Lycopodiaceae ,5 Pteridophyta Lygodiaceae ,0 Dicotyledoneae Lythraceae 8 35 Dicotyledoneae Malesherbiaceae 1 3 Dicotyledoneae Malpighiaceae ,5 Dicotyledoneae Malvaceae ,7 Monocotyledoneae Marantaceae 4 6 Pteridophyta Marattiaceae 1 1 Pteridophyta Marsileaceae 3 5 Dicotyledoneae Martyniaceae ,0 Monocotyledoneae Mayacaceae 1 2 Dicotyledoneae Melastomataceae ,1 Dicotyledoneae Meliaceae ,1 Dicotyledoneae Menispermaceae 3 6 Dicotyledoneae Menyanthaceae 1 1 Dicotyledoneae Microteaceae 1 1 Dicotyledoneae Misodendraceae 1 7 Dicotyledoneae Molluginaceae 2 3 Dicotyledoneae Monimiaceae 2 2 Dicotyledoneae Montiaceae ,8 Dicotyledoneae Moraceae 6 14 Dicotyledoneae Muntingiaceae 1 1 Dicotyledoneae Myoporaceae 1 1 Dicotyledoneae Myricaceae 1 1 Dicotyledoneae Myrsinaceae 1 6 Dicotyledoneae Myrtaceae ,1 Monocotyledoneae Najadaceae 1 2 Dicotyledoneae Nanodeaceae 1 1 Pteridophyta Nephrolepidaceae 1 1 Dicotyledoneae Nothofagaceae 2 6 Dicotyledoneae Nyctaginaceae ,5 Dicotyledoneae Nymphaeaceae 2 6 Dicotyledoneae Ochnaceae 1 2 Dicotyledoneae Oleaceae ,7 Dicotyledoneae Onagraceae ,9 Pteridophyta Ophioglossaceae 2 9 Dicotyledoneae Opiliaceae 1 1 Rodriguésia 66(4):

74 Vascular Plants of the Southern Cone 1005 Group Family Genera Species Endemics %Endemics Monocotyledoneae Orchidaceae ,4 Dicotyledoneae Orobanchaceae ,6 Pteridophyta Osmundaceae 2 2 Dicotyledoneae Oxalidaceae ,7 Dicotyledoneae Papaveraceae 6 12 Pteridophyta Parkeriaceae 1 1 Dicotyledoneae Parnassiaceae 1 1 Dicotyledoneae Passifloraceae 1 20 Dicotyledoneae Pentaphylacaceae 1 1 Monocotyledoneae Philesiaceae 1 1 Dicotyledoneae Phrymaceae 2 6 Dicotyledoneae Phyllanthaceae 2 10 Dicotyledoneae Phytolaccaceae ,7 Dicotyledoneae Picrodendraceae 1 1 Gymnospermae Pinaceae 2 4 Dicotyledoneae Piperaceae ,9 Dicotyledoneae Plantaginaceae ,2 Dicotyledoneae Plumbaginaceae 3 4 Monocotyledoneae Poaceae ,1 Gymnospermae Podocarpaceae 4 6 Dicotyledoneae Podostemaceae ,1 Dicotyledoneae Polemoniaceae ,2 Dicotyledoneae Polygalaceae ,4 Dicotyledoneae Polygonaceae ,4 Pteridophyta Polypodiaceae ,3 Monocotyledoneae Pontederiaceae 3 10 Dicotyledoneae Portulacaceae ,3 Monocotyledoneae Potamogetonaceae 2 12 Dicotyledoneae Primulaceae 6 11 Dicotyledoneae Proteaceae 6 8 Pteridophyta Psilotaceae 1 1 Pteridophyta Pteridaceae ,0 Dicotyledoneae Quillajaceae 1 1 Dicotyledoneae Rafflesiaceae 1 2 Dicotyledoneae Ranunculaceae ,5 Dicotyledoneae Resedaceae 1 1 Dicotyledoneae Rhamnaceae ,4 Dicotyledoneae Rosaceae ,5 Dicotyledoneae Rubiaceae ,8 Monocotyledoneae Ruppiaceae 1 3 Monocotyledoneae Ruscaceae 1 1 Dicotyledoneae Rutaceae 8 17 Pteridophyta Saccolomataceae 1 1 Rodriguésia 66(4):

75 1006 Zuloaga, F.O. & Belgrano, M.J. Group Family Genera Species Endemics %Endemics Dicotyledoneae Salicaceae ,3 Pteridophyta Salviniaceae 1 4 Dicotyledoneae Samolaceae 1 3 Dicotyledoneae Santalaceae 1 1 Dicotyledoneae Sapindaceae ,3 Dicotyledoneae Sapotaceae 3 7 Dicotyledoneae Saxifragaceae 5 6 Pteridophyta Schizaeaceae 1 1 Dicotyledoneae Schoepfiaceae ,7 Dicotyledoneae Scrophulariaceae ,9 Pteridophyta Selaginellaceae 1 8 Dicotyledoneae Simaroubaceae 6 8 Monocotyledoneae Smilacaceae 1 5 Dicotyledoneae Solanaceae ,0 Dicotyledoneae Sphenocleaceae 1 1 Dicotyledoneae Sterculiaceae ,7 Dicotyledoneae Stylidiaceae 1 1 Dicotyledoneae Styracaceae 1 2 Dicotyledoneae Symplocaceae 1 2 Dicotyledoneae Talinaceae 1 3 Dicotyledoneae Tamaricaceae 1 1 Pteridophyta Tectariaceae 1 1 Dicotyledoneae Tetrachondraceae 1 1 Dicotyledoneae Theaceae 1 1 Pteridophyta Thelypteridaceae ,6 Dicotyledoneae Thymelaeaceae 3 3 Dicotyledoneae Tiliaceae 4 12 Dicotyledoneae Tribelaceae 1 1 Dicotyledoneae Trigoniaceae 1 1 Monocotyledoneae Triuridaceae 1 1 Dicotyledoneae Tropaeolaceae ,8 Dicotyledoneae Turneraceae 2 14 Monocotyledoneae Typhaceae 1 4 Dicotyledoneae Ulmaceae 1 1 Dicotyledoneae Urticaceae ,0 Dicotyledoneae Valerianaceae ,8 Monocotyledoneae Velloziaceae ,0 Dicotyledoneae Verbenaceae ,9 Dicotyledoneae Violaceae ,4 Dicotyledoneae Viscaceae 1 16 Dicotyledoneae Vitaceae 3 10 Pteridophyta Vittariaceae 2 3 Dicotyledoneae Vivianiaceae 2 3 Rodriguésia 66(4):

76 Vascular Plants of the Southern Cone 1007 Group Family Genera Species Endemics %Endemics Dicotyledoneae Vochysiaceae 1 1 Dicotyledoneae Winteraceae 1 1 Pteridophyta Woodsiaceae 2 3 Dicotyledoneae Ximeniaceae 1 1 Monocotyledoneae Xyridaceae 1 7 Monocotyledoneae Zannichelliaceae 1 1 Monocotyledoneae Zingiberaceae 1 1 Dicotyledoneae Zygophyllaceae ,3 Total Halophytaceae and the endemic genera are 44 (2.10% of the total of Argentinian genera), with the Asteraceae leading in number of endemic genera (12), followed by Brassicaceae (8), Cactaceae (4), Fabaceae and Verbenaceae (3) (Tab. 5); the total endemic species for Argentina is 1,749 (17,48% of the total number of species for this country). Endemism in the Biogeographic Regions Of the eight families endemic from the Southern Cone, the monotypic Lactoridaceae is exclusive from the Robison Crusoe Island (Más a Tierra), in the Juan Fernández Archipelago (Chile), included in the Temperate-humid region of the Pacific, with a flora related to the Subantarctic phytogeographic province, however counting with subtropical and other specific elements from the Pacific Islands. The Francoaceae, with two monotypic genera and the monotypic Gomortegaceae are exclusive from the mediterranean region of Central Chile. All four endemic families shared between Argentina and Chile are from the Subantartic forests, the Aextoxicaceae and Tribelaceae are monotypic and the Philesiaceae comprises two monotypic genera, while the Misodendraceae has a single genus and eight hemiparasitic species. The Halophytaceae, the only endemic family from Argentina, is exclusive from the Monte region, in arid pre-montane areas. In the South-Central dry Andes, which include the Puna, Prepuna and high-andean regions in northern Argentina and Chile, there are several endemic genera of Asteraceae such as Cabreraea Bonifacino, Dolichlasium Lag. and Famatinanthus Ariza & S.E. Freire, of Brassicaceae, as Lithodraba Boelcke, Parodiodoxa O.E. Schulz, Sarcodraba Gilg & Muschl. and Zuloagocardamum Salariato & Al-Shehbaz and of Montiaceae, as Schreiteria Carolin, amongst others. Genera that are characteristic but not endemic to the Southern Cone are Clinanthus Herb. and Hieronymiella Pax (Amaryllidaceae), Chiliotrichiopsis Cabrera, Cuatrecasasiella H. Rob., Urmenetea Phil. and Werneria Kunth (Asteraceae), Aschersoniodoxa Gilg & Muschl., Mancoa Wedd. and Polypsecadium O. E. Schulz (Brassicaceae), Pycnophyllum J. Rémy (Caryophyllaceae), Cardenanthus R.C. Forster and Mastigostyla I.M. Johnst. (Iridaceae), Nototriche Turcz. and Tarasa Phil. (Malvaceae), Hypseocharis J. Rémy (Oxalidaceae), Aa Juss. and Myrosmodes Rchb. f. (Orchidaceae), Lenzia Phil. (Montiaceae) and Anthochloa Nees & Meyen and Aciachne Benth. (Poaceae), among others. Some genera are common between the high Andes and the Patagonic region, such as Adesmia (Fabaceae), Azorella Lam. and Mulinum Pers. (Apiaceae), Chuquiraga Juss. and Nardophyllum Hook. & Arn. (Asteraceae), Fabiana Ruiz & Pav. (Solanaceae) and Junellia Moldenke (Verbenaceae). Among the genera that are endemic to the central Andes of Mendoza and San Juan (Argentina) are Haroldia Bonifacino, Huarpea Cabrera and Katinasia Bonifacino (Asteraceae). The North-Central Humid Andes or Yungas show few endemic genera, one of the examples being Austropeucedanum Mathias & Constance (Apiaceae), and this may be explained by the continuation of the Yungas into bordering areas of Bolivia. The characteristic or exclusive elements within this region include holartic families such as BetulaWceae (Alnus Mill.) and Juglandaceae (Juglans L.) and other families that reach northern Argentina, such as Myricaceae (Myrica L.), Pentaphylacaceae (Ternstroemia Mutis ex L. f.) and Velloziaceae (Barbaceniopsis L.B. Sm.). Rodriguésia 66(4):

77 1008 Zuloaga, F.O. & Belgrano, M.J. Table 5 Southern Cone, endemic genera. Family Genus NºSpp. Countries Aextoxicaceae Aextoxicon 1 Argentina Chile Alstroemeriaceae Leontochir 1 Chile Amaranthaceae Quaternella 1 S Brasil Amaryllidaceae Ancrumia 1 Chile Beauverdia 4 Argentina S Brasil Uruguay Eithea 1 S Brasil Famatina 2 Argentina Chile Gethyum 2 Chile Gilliesia 5 Argentina Chile Ipheion 4 Argentina Chile Uruguay Leucocoryne 45 Chile Miersia 4 Chile Phycella 6 Argentina Chile Placea 5 Chile Solaria 3 Argentina Chile Speea 1 Chile Traubia 1 Chile Tristagma 20 Argentina Chile Zoellnerallium 2 Argentina Chile Apiaceae Asteriscium 9 Argentina Chile Austropeucedanum 1 Argentina Bolax 2 Argentina Chile Diposis 3 Argentina Chile Uruguay Gymnophyton 6 Argentina Chile Homalocarpus 6 Chile Huanaca 4 Argentina Chile Laretia 1 Argentina Chile Notiosciadium 1 Argentina Uruguay Oligocladus 1 Argentina Pozoa 2 Argentina Chile Apocynaceae Diplolepis 14 Argentina Chile Elytropus 1 Chile Rhyssostelma 1 Uruguay Araceae Mangonia 2 S Brasil Uruguay Arecaceae Juania 1 Chile Jubaea 1 Chile Asteraceae Acrisione 1 Argentina Chile Ameghinoa 1 Argentina Asteropsis 1 S Brasil Uruguay Aylacophora 1 Argentina Brachyclados 3 Argentina Chile Rodriguésia 66(4):

78 Vascular Plants of the Southern Cone 1009 Family Genus NºSpp. Countries Burkartia 1 Argentina Cabreraea 1 Argentina Calopappus 1 Chile Centaurodendron 2 Chile Chiliophyllum 3 Argentina Chile Chiliotrichum 2 Argentina Chile Criscia 1 Argentina S Brasil Uruguay Dendroseris 11 Chile Dolichlasium 1 Argentina Doniophyton 2 Argentina Chile Duseniella 1 Argentina Eriachaenium 1 Argentina Famtinanthus 1 Argentina Gamochaetopsis 1 Argentina Chile Guynesomia 1 Chile Gypothamnium 1 Chile Haroldia 1 Argentina Heterothalamulopsis 1 S Brasil Huarpea 1 Argentina Ianthopappus 1 Argentina S Brasil Uruguay Katinasia 1 Argentina Kieslingia 1 Chile Lepidophyllum 1 Argentina Chile Leptinella 1 Argentina Chile Leptocarpha 1 Chile Leunisia 1 Chile Lycapsus 1 Chile Macrachaenium 1 Argentina Marticorenia 1 Chile Microgyne 1 Argentina S Brasil Uruguay Micropsis 5 Argentina Chile Uruguay Moscharia 2 Chile Oxyphyllum 1 Chile Panphalea 9 Argentina S Brasil Uruguay Pleocarphus 1 Chile Podanthus 2 Chile Sommerfeltia 2 Argentina S Brasil Uruguay Thamnoseris 1 Chile Triptilion 7 Argentina Chile Urmenetea 1 Argentina Chile Yunquea 1 Chile Bignoniaceae Campsidium 1 Argentina Chile Boraginaceae Ixorhea 1 Argentina Rodriguésia 66(4):

79 1010 Zuloaga, F.O. & Belgrano, M.J. Family Genus NºSpp. Countries Nesocaryum 1 Chile Selkirkia 1 Chile Brassicaceae Chilocardamum 4 Argentina Delpinophytum 1 Argentina Hollermayera 1 Chile Ivania 1 Chile Lithodraba 1 Argentina Onuris 5 Argentina Chile Parodiodoxa 1 Argentina Phlebolobium 1 Argentina Sarcodraba 4 Argentina Schizopetalon 10 Argentina Chile Trichotolinum 1 Argentina Xerodraba 7 Argentina Chile Zuloagocardamum 1 Argentina Bromeliaceae Fascicularia 1 Chile Ochagavia 4 Chile Cactaceae Acanthocalycium 5 Argentina Austrocactus 4 Argentina Chile Denmoza 1 Argentina Eriosyce 5 Chile Maihuenia 2 Argentina Chile Pterocactus 9 Argentina Chile Pyrrhocactus 27 Argentina Chile Setiechinopsis 1 Argentina Tephrocactus 7 Argentina Thelocephala 6 Argentina Chile Calyceraceae Boopis 13 Argentina Chile Gamocarpha 6 Argentina Chile Nastanthus 9 Argentina Chile Campanulaceae Cyphocarpus 3 Chile Caryophyllaceae Philippiella 1 Argentina Chile Reicheella 1 Chile Corsiaceae Arachnitis 1 Argentina Chile Cucurbitaceae Halosicyos 1 Argentina Cupressaceae Austrocedrus 1 Argentina Chile Fitzroya 1 Argentina Chile Pilgerodendron 1 Argentina Chile Dicksoniaceae Thyrsopteris 1 Chile Epacridaceae Lebetanthus 1 Argentina Chile Escalloniaceae Valdivia 1 Chile Euphorbiaceae Adenopeltis 1 Chile Avellanita 1 Chile Rodriguésia 66(4):

80 Vascular Plants of the Southern Cone 1011 Family Genus NºSpp. Countries Colliguaja 5 Argentina S Brasil Chile Fabaceae Anarthrophyllum 15 Argentina Chile Balsamocarpon 1 Argentina Chile Lophocarpinia 1 Argentina Paraguay Ramorinoa 1 Argentina Stenodrepanum 1 Argentina Zuccagnia 1 Argentina Francoaceae Francoa 1 Chile Tetilla 1 Chile Gesneriaceae Asteranthera 1 Argentina Chile Mitraria 1 Argentina Chile Sarmienta 1 Chile Gomortegaceae Gomortega 1 Chile Halophytaceae Halophytum 1 Argentina Hymenophyllaceae Hymenoglossum 1 Argentina Chile Serpyllopsis 1 Argentina Chile Iridaceae Solenomelus 2 Argentina Chile Tapeinia 1 Argentina Juncaginaceae Tetroncium 1 Argentina Chile Lactoridaceae Lactoris 1 Chile Lamiaceae Cuminia 1 Chile Kurzamra 1 Argentina Chile Lardizabalaceae Boquila 1 Argentina Chile Lardizabala 1 Argentina Chile Laxmanniaceae Trichopetalum 2 Argentina Chile Loasaceae Huidobria 2 Chile Scyphanthus 1 Chile Loranthaceae Desmaria 1 Chile Notanthera 1 Chile Malpighiaceae Cordobia 1 Argentina Paraguay Dinemagonum 1 Chile Dinemandra 1 Chile Gallardoa 1 Argentina Peregrina 1 Argentina S Brasil Paraguay Tricomaria 1 Argentina Malvaceae Bordasia 1 Paraguay Calyculogygas 1 S Brasil Uruguay Calyptraemalva 1 S Brasil Corynabutilon 7 Argentina Chile Lecanophora 7 Argentina Chile Neobaclea 1 Argentina Tropidococcus 1 S Brasil Marsileaceae Regnellidium 1 Argentina S Brasil Uruguay Rodriguésia 66(4):

81 1012 Zuloaga, F.O. & Belgrano, M.J. Family Genus NºSpp. Countries Misodendraceae Misodendrum 8 Argentina Chile Monimiaceae Laureliopsis 1 Argentina Chile Peumus 1 Chile Myrtaceae Amomyrtus 2 Argentina Chile Legrandia 1 Chile Tepualia 1 Argentina Chile Nanodeaceae Nanodea 1 Argentina Chile Nothofagaceae Nothofagus 4 Argentina Chile Orchidaceae Bipinnula 10 Argentina S Brasil Chile Uruguay Codonorchis 2 Argentina S Brasil Chile Gavilea 14 Argentina Chile Philesiaceae Lapageria 1 Chile Philesia 1 Argentina Chile Phytolaccaceae Anisomeria 2 Chile Ercilla 2 Chile Plantaginaceae Fonkia 1 Argentina Chile Melosperma 1 Argentina Chile Monttea 3 Argentina Chile Poaceae Eremium 1 Argentina Chile Leptophyllochloa 1 Argentina Chile Megalachne 1 Chile Neobouteloua 2 Argentina Nicoraepoa 7 Argentina chile Oplismenopsis 1 Argentina Uruguay Osvaldoa 1 Argentina Podophorus 1 Chile Relchela 1 Argentina Chile Podocarpaceae Lepidothamnus 1 Argentina Chile Saxegothaea 1 Argentina Chile Podostemaceae Heterotristicha 1 Uruguay Polypodiaceae Synammia 3 Chile Portulacaceae Amphipetalum 1 Paraguay Lenzia 1 Argentina Chile Schreiteria 1 Argentina Proteaceae Gevuina 1 Argentina Chile Ranunculaceae Barneoudia 3 Argentina Chile Callianthemoides 1 Argentina Chile Hamadryas 5 Argentina Chile Rhamnaceae Ochetophila 2 Argentina Chile Retanilla 4 Argentina Chile Trevoa 1 Chile Rosaceae Margyracaena 1 Chile Rubiaceae Oreopolus 1 Argentina Chile Rodriguésia 66(4):

82 Vascular Plants of the Southern Cone 1013 Family Genus NºSpp. Countries Rutaceae Pitavia 1 Chile Rutaceae Raulinoa 1 S Brasil Santalaceae Myoschilos 1 Argentina Chile Sapindaceae Bridgesia 1 Chile Guindilia 3 Argentina Chile Saxifragaceae Saxifragella 1 Argentina Chile Saxifragodes 1 Argentina Chile Solanaceae Benthamiella 12 Argentina Chile Combera 2 Argentina Chile Latua 1 Chile Pantacantha 1 Argentina Reyesia 4 Argentina Chile Salpiglossis 2 Argentina Chile Schizanthus 14 Argentina Chile Sclerophylax 14 Argentina Paraguay Uruguay Vestia 1 Chile Tecophilaeaceae Conanthera 5 Chile Tecophilaea 2 Chile Tribelaceae Tribeles 1 Argentina Chile Verbenaceae Dipyrena 1 Argentina Neosparton 4 Argentina Parodianthus 2 Argentina Vivianiaceae Araeoandra 1 Chile Cissarobryon 1 Chile Viviania 4 Argentina S Brasil Chile Zygophyllaceae Metharme 1 Chile Pintoa 1 Chile Plectrocarpa 2 Argentina The Atlantic forest region, in the eastern extreme of Argentina and adjacent areas of Paraguay and southern Brazil, has high richness of families, genera and species, but there are few endemisms, with no records of endemic families, and just 9 genera that grow exclusively in the Paraná related forests of the Southern Cone: Quaternella Pederesen (Amaranthaceae), Eithea Ravenna (Amaryllidaceae), Criscia Katinas and Heterothalamulopsis Deble, A.S. Oliveira & Marchiori (Asteraceae), Peregrina W.R. Anderson (Malpighiaceae), Calyptraemalva Krapov. and Tropidococcus Krapov. (Malvaceae), Amphipetalum Bacigalupo (Portulacaceae) and Raulinoa R.S. Cowan (Rutaceae). In Chile, the region of the Chilean-Peruvian Desert includes aproximately 15 endemic genera, chiefly for the Cactaceae (Eriosyce Phil. and Thelocephala Y. Ito), Asteraceae (Gypothamnium Phil., Kieslingia Faúndez, Saldivia & A.E. Martic. and Oxyphyllum Phil.), Alstroemeriaceae (Leontochir Phil.), Loasaceae (Huidobria Gay) and Zygophyllaceae (Metharme Phil. ex Engl. and Pintoa Gay), among others. A high number of endemic species also occurs in this region, including numerous species of Cryptantha G. Don and Heliotropium L. (Boraginaceae), Copiapoa Britton & Rose (Cactaceae) and Nolana L.f. (Solanaceae). The Chilean mediterranean region comprises an important number of endemic taxa, with Rodriguésia 66(4):

83 1014 Zuloaga, F.O. & Belgrano, M.J. over 50 genera that are exclusive to this region or spreading to contiguous areas, for example Jubaea Kunth (Arecaceae), many genera of Amaryllidaceae (Gethyum Phil., Leucocoryne Lindl. and Speea Loes.), Asteraceae (Calopappus Meyen, Guynesomia Bonifacino & Sancho, Leunisia Phil., Marticorenia Crisci, Moscharia Ruiz & Pav., Pleocarphus D. Don and Podanthus Lag.), Bromeliaceae (Fascicularia Mez and Ochagavia Phil.), Monimiaceae (Peumus Molina), Tecophilaeaceae (Conanthera Ruiz & Pav. and Tecophilaea Bertero ex Colla), Vivianiaceae (Araeoandra Lefor and Cissarobryon Kunze ex Poepp.), Francoaceae (Francoa Cav. and Tetilla DC.), Gomortegaceae (Gomortega Ruiz & Pav.) and Zygophyllaceae (Pintoa). The number of endemic species is also outstanding. For the Pampas region, there are 15 endemic genera, including representatives of Asteraceae (Asteropsis Less., Ianthopappus Roque & D.J.N. Hind, Microgyne Less., Panphalea Lag. and Sommerfeltia Less.), Araceae (Mangonia Schott), Poaceae (Oplismenopsis Parodi and Osvaldoa), Amaryllidaceae (Beauverdia Herter and Ipheion Raf.), Apiaceae (Notiosciadium Speg.) and Apocynaceae (Rhyssostelma Decne.). The genus Sclerophylax Miers (Solanaceae) comprises 14 species chiefly from the Pampas with some extension into the Monte region. The Monte includes many endemic genera, more than 20 exclusive from Argentina, such as Acanthocalycium Backeb., Denmoza Britton & Rose, Setiechinopsis (Backeb.) de Haas and Tephrocactus Lem. (all Cactaceae), Ramorinoa Speg., Stenodrepanum Harms and Zuccagnia Cav. (Fabaceae), Dipyrena Hook. and Parodianthus Tronc. (Verbenaceae), Ixorhea Fenzl (Boraginaceae), Halosicyos Mart.Crov. (Cucurbitaceae), Plectrocarpa Gillies ex Hook. & Arn. (Zygophyllaceae) and Tricomaria Hook. & Arn. (Malpighiaceae). Genera found in the Monte sometimes extend into mediterranean Chile, such as Kurzamra Kuntze (Lamiaceae), Guindilia Hook. & Arn. (Sapindaceae), Monttea Gay (Plantaginaceae) and Barneoudia Gay (Ranunculaceae). Even taking into account that as one approaches higher latitudes the number of taxa diminishes, the number of endemic genera in the Patagonian region increases, with many representatives of Asteraceae (Ameghinoa Speg., Aylacophora Cabrera, Burkartia Crisci, Duseniella K. Schum., Lepidophyllum Cass. and Doniophyton Wedd.), Brassicaceae (Chilocardamum O.E. Schulz, Delpinophytum Speg., Onuris Phil., Phlebolobium O.E. Schulz and Xerodraba Skottsb.), Cactaceae (Austrocactus Britton & Rose, Maihuenia (F.C.A. Weber) K. Schum. and Pterocactus K. Schum.), Calyceraceae (Boopis Juss., Gamocarpha DC. and Nastanthus Miers), Fabaceae (Anarthrophyllum Benth.), Verbenaceae (Neosparton Griseb.), Solanaceae (Benthamiella Speg. and Pantacantha Speg.) and Malpighiaceae (Gallardoa Hicken). Genera present in this region and also in central Chile include Triptilion Ruiz & Pav. (Asteraceae), Trichopetalum Lindl. (Laxmanniaceae), Melosperma Benth. (Plantaginaceae) and Retanilla (DC.) Brongn. (Rhamnaceae), Austrocactus and Maihuenia (Cactaceae). Amongst Patagonian genera also found in the Monte we have Lecanophora Speg. (Malvaceae) and Neosparton (Verbenaceae). Only three genera are endemic from the Chaco: Lophocarpinia Burkart (Fabaceae), Bordasia Krapov. (Malvaceae) and Cordobia Nied. (Malpighiaceae), as this biome crosses over to Bolivia and the Central-Western area of Brazil (not included under this project). Subantartic forests, within the temperatehumid Pacific region, include expressive numbers of endemic families and genera; however, the number of endemic species is not very high. The four endemic families from the region were indicated above; other families that have representatives in other places have here their unique presence in South America, as the Cupressaceae (with three endemic genera: Austrocedrus Florin & Boutelje, Fitzroya Hook. f. ex Lindl. and Pilgerodendron Florin) and Restionaceae (Apodasmia B.G. Briggs & L.A.S. Johnson). There are around 60 endemic genera in this region, among them the Juan Fernández Islands host 13, including an Arecaceae (Juania Drude), genera of Asteraceae (Centaurodendron Johow, Dendroseris D. Don, Yunquea Skottsb.), Boraginaceae (Selkirkia Hemsl.), Dicksoniaceae (Thyrsopteris Kunze), Lactoridaceae (Lactoris Phil.), Lamiaceae (Cuminia Colla), Myrtaceae (Nothomyrcia Kausel) and Poaceae (Megalachne Steud. and Podophorus Phil.). The continental zone has endemic genera that are exclusive from Chile or shared between Chile and Argentina of Asteraceae (Gamochaetopsis Anderb. & S.E. Freire, Leptinella Cass., Leptocarpha DC. and Macrachaenium Hook.f.), Epacridaceae (Lebetanthus Endl.), Escalloniaceae (Valdivia Rodriguésia 66(4):

84 Vascular Plants of the Southern Cone Gay ex J. Rémy), Euphorbiaceae (Avellanita Phil.), Gesneriaceae (Asteranthera Hanst. and Sarmienta Ruiz & Pav.), Iridaceae (Solenomelus Miers and Tapeinia Comm. ex Juss.), Lardizabalaceae (Boquila Decne. and Lardizabala Ruiz & Pav.), Monimiaceae (Laureliopsis Schodde), Podocarpaceae (Lepidothamnus Phil. and Saxegothaea Lindl.), among others. Families Araucariaceae (Araucaria Juss.), Podocarpaceae (Podocarpus L Her. ex Pers.), Thymelaeaceae (Ovidia Meisn. and Drapetes Lam.), Hymenophyllaceae (Hymenophyllum Sm.), Blechnaceae (Blechnum L.) and Winteraceae (Drimys J.R. Forst. & G. Forst.) are disjunct between this region and the Atlantic Forest. The analysis of the distribution of endemisms through the different biogeographic regions allows us to conclude that the number of endemic or exclusive families and genera is high in the Subantarctic forests, but there is a smaller number of endemic species in relation to other areas. Mediterranean Chile and deserts of Patagonia and Monte have a high number of endemic families, genera and species when compared with the totality of the analised flora, and such indices decrease as the latitude increases towards Patagonia, together with the general decrease in species richness of the vascular flora. The Chilean-Peruvian desert and adjacent high Andean and Puna areas in norhtern Argentina do not possess endemic families and the number of endemic genera is smaller, while 1015 the number of endemic species is outstanding. The amount of endemism diminishes markedly from west to east, with low numbers in the Chaco and Pampa, growing again, in terms of number of endemic species, in the Atlantic Forest from of eastern Paraguay, northeastern Argentina and southern Brazil. Introduced taxa A total of 19 introduced families are present in the Southern Cone, representing 6% of the total number of families in this region, namely: Aceraceae, Achariaceae, Agapanthaceae, Agavaceae, Aponogetonaceae, Asparagaceae, Asphodelaceae, Balsaminaceae, Cannabaceae, Caprifoliaceae, Dipsacaceae, Elaeagnaceae, Fumariaceae, Myoporaceae, Pinaceae, Resedaceae, Ruscaceae, Tamaricaceae and Zingiberaceae. The area counts with a total of 344 genera represented by introduced species (12.84%) and a total of 1,129 introduced species (6.2% of the total of species found in the region). The family with the highest number of introduced genera is Asteraceae, with 54 genera, followed by Poaceae (42 genera), Brassicaceae (33), Fabaceae (22), Apiaceae (20), Lamiaceae (14) and Rosaceae (10) (Tab. 6). The country where the highest record of introduced taxa was found is Argentina with 892, or 8.91% of the total number of species recorded for this country. Table 6 Southern Cone, introduced genera. Family Genus Nº Spp. Countries Acanthaceae Hypoestes 1 Paraguay Aceraceae Acer 2 Argentina Achariaceae Hydnocarpus 1 Paraguay Agapanthaceae Agapanthus 1 Argentina Agavaceae Agave 1 Argentina Yucca 2 Argentina Aizoaceae Aptenia 1 Argentina Cypselea 1 Argentina Galenia 1 Chile Sesuvium 1 Argentina S Brasil Chile Paraguay Uruguay Alismataceae Alisma 2 Argentina Chile Amaryllidaceae Allium 6 Argentina Amaryllis 1 S Brasil Chile Paraguay Brunsvigia 1 Argentina Rodriguésia 66(4):

85 1016 Zuloaga, F.O. & Belgrano, M.J. Family Genus Nº Spp. Countries Pyrolirion 1 Chile Anacardiaceae Mangifera 1 S Brasil Paraguay Apiaceae Ammi 2 Argentina S Brasil Chile Uruguay Anethum 1 Argentina S Brasil Chile Anthriscus 2 Argentina Chile Bupleurum 1 Argentina Caucalis 1 Argentina Conium 1 Argentina S Brasil Chile Coriandrum 1 Argentina S Brasil Chile Paraguay Foeniculum 1 Argentina S Brasil Chile Paraguay Uruguay Heracleum 1 Argentina Levisticum 1 Chile Oenanthe 2 Argentina Uruguay Pastinaca 1 Argentina Chile Uruguay Petroselinum 1 Argentina S Brasil Chile Pimpinella 1 Argentina S Brasil Scandix 1 Argentina Chile Seseli 1 Chile Sium 1 Chile Tordylium 1 Argentina Torilis 2 Argentina S Brasil Chile Uruguay Apocynaceae Calotropis 1 S Brasil Paraguay Catharanthus 1 Argentina S Brasil Paraguay Uruguay Gomphocarpus 1 S Brasil Plumeria 1 Argentina S Brasil Vinca 1 Argentina S Brasil Chile Uruguay Aponogetonaceae Aponogeton 1 Chile Araceae Alocasia 1 Paraguay Arum 1 Argentina Uruguay Colocasia 1 Argentina Paraguay Uruguay Zantedeschia 1 Argentina S Brasil Chile Uruguay Araliaceae Hedera 1 Argentina Arecaceae Phoenix 1 Argentina Trachycarpus 1 Argentina Asparagaceae Asparagus 4 Argentina Uruguay Asphodelaceae Asphodelus 1 Argentina Chile Asteraceae Achillea 2 Argentina S Brasil Chile Uruguay Anthemis 2 Argentina S Brasil Chile Paraguay Uruguay Arctium 1 Argentina S Brasil Chile Uruguay Arctotheca 1 Argentina Chile Arctotis 1 Argentina S Brasil Uruguay Argyranthemum 1 Argentina Chile Arnica 1 Argentina Chile Rodriguésia 66(4):

86 Vascular Plants of the Southern Cone 1017 Family Genus Nº Spp. Countries Bellis 1 Argentina Chile Calendula 3 Argentina Chile Uruguay Carduus 5 Argentina S Brasil Chile Uruguay Carthamus 2 Argentina S Brasil Chile Uruguay Chondrilla 1 Argentina Chrysanthemoides 1 Chile Chrysanthemum 2 Argentina Chile Uruguay Cichorium 1 Argentina S Brasil Chile Uruguay Cirsium 2 Argentina S Brasil Chile Paraguay Uruguay Cladanthus 1 Argentina S Brasil Chile Uruguay Coleostephus 1 Argentina S Brasil Chile Uruguay Crepis 4 Argentina S Brasil Chile Uruguay Cynara 1 Argentina S Brasil Chile Uruguay Dahlia 1 Argentina Delairea 1 Argentina Chile Uruguay Dyssodia 1 Argentina Emilia 1 Argentina S Brasil Paraguay Euryops 1 Argentina Gazania 1 Argentina Gynura 1 Argentina Hedypnois 1 Argentina Chile Uruguay Helianthus 4 Argentina Uruguay Heterotheca 1 Argentina Lactuca 4 Argentina S Brasil Chile Paraguay Uruguay Lapsana 1 Argentina S Brasil Chile Leontodon 4 Argentina Chile Leucanthemum 1 Argentina S Brasil Chile Uruguay Logfia 1 Chile Malacothrix 1 Argentina Chile Matricaria 2 Argentina S Brasil Chile Paraguay Uruguay Microseris 1 Chile Montanoa 1 Argentina Onopordum 2 Argentina S Brasil Chile Uruguay Picris 1 Argentina Chile Uruguay Ratibida 1 Argentina Rhaponticum 1 Argentina Scolymus 1 Argentina Chile Scorzonera 1 Argentina Silybum 1 Argentina S Brasil Chile Uruguay Sonchus 4 Argentina S Brasil Chile Paraguay Uruguay Struchium 1 S Brasil Tanacetum 3 Argentina S Brasil Chile Uruguay Tithonia 2 Argentina Rodriguésia 66(4):

87 1018 Zuloaga, F.O. & Belgrano, M.J. Family Genus Nº Spp. Countries Tolpis 1 Chile Tragopogon 3 Argentina Chile Tripleurospermum 1 Argentina Chile Uruguay Urospermum 1 Argentina Chile Uruguay Athyriaceae Deparia 1 Argentina S Brasil Balsaminaceae Impatiens 3 Argentina S Brasil Bignoniaceae Campsis 1 Argentina Podranea 1 Argentina S Brasil Boraginaceae Anchusa 1 Argentina Asperugo 1 Argentina Borago 1 Argentina Chile Uruguay Echium 1 Argentina Chile Lithospermum 2 Argentina Uruguay Lycopsis 1 Argentina Omphalodes 1 Chile Symphytum 1 Argentina Brassicaceae Aethionema 1 Argentina Alliaria 1 Argentina Alyssum 1 Argentina Chile Arabidopsis 1 Argentina Chile Uruguay Arabis 1 Argentina Barbarea 2 Argentina Chile Brassica 5 Argentina Chile Paraguay Uruguay Cakile 1 Argentina Uruguay Camelina 3 Argentina Chile Uruguay Capsella 1 Argentina S Brasil Chile Uruguay Chorispora 1 Argentina Chile Cochlearia 1 Argentina Diplotaxis 3 Argentina Chile Uruguay Eruca 1 Argentina Chile Erysimum 2 Argentina Hesperis 1 Argentina Chile Hirschfeldia 1 Argentina Chile Uruguay Hornungia 1 Argentina Chile Iberis 1 Argentina Isatis 1 Chile Lobularia 1 Argentina Chile Uruguay Lunaria 1 Argentina Malcolmia 1 Argentina Matthiola 1 Chile Nasturtium 2 Argentina S Brasil Chile Uruguay Neslia 1 Argentina Raphanus 3 Argentina S Brasil Chile Paraguay Uruguay Rodriguésia 66(4):

88 Vascular Plants of the Southern Cone 1019 Family Genus Nº Spp. Countries Rapistrum 1 Argentina S Brasil Chile Paraguay Uruguay Sinapis 2 Argentina Chile Paraguay Sisymbrium 7 Argentina S Brasil Chile Uruguay Strigosella 1 Argentina Teesdalia 1 Chile Thlaspi 1 Argentina Chile Cactaceae Cylindropuntia 1 Argentina Chile Nopalea 1 Paraguay Campanulaceae Campanula 1 Argentina Hippobroma 1 S Brasil Cannabaceae Humulus 2 Argentina Caprifoliaceae Lonicera 1 Argentina S Brasil Uruguay Symphoricarpos 1 Argentina Caryophyllaceae Agrostemma 1 Argentina Chile Uruguay Dianthus 2 Argentina Chile Herniaria 2 Argentina Chile Holosteum 1 Argentina Lychnis 1 S Brasil Chile Petrorhagia 3 Argentina Chile Saponaria 1 Argentina Chile Uruguay Scleranthus 1 Argentina Chile Uruguay Vaccaria 1 Argentina Uruguay Chenopodiaceae Bassia 2 Argentina Beta 1 Argentina Chile Blitum 1 Argentina Chenopodiastrum 1 Argentina Chile Uruguay Cycloloma 1 Argentina Salsola 2 Argentina Chile Uruguay Crassulaceae Kalanchoe 1 Argentina Cucurbitaceae Citrullus 2 Argentina S Brasil Paraguay Cucumis 1 Argentina S Brasil Paraguay Lagenaria 1 Argentina S Brasil Paraguay Uruguay Luffa 3 Argentina S Brasil Paraguay Cupressaceae Juniperus 1 Argentina Dipsacaceae Dipsacus 1 Argentina Uruguay Knautia 2 Argentina Chile Scabiosa 1 Argentina Chile Uruguay Dryopteridaceae Cyrtomium 1 Argentina Elaeagnaceae Elaeagnus 1 Argentina Euphorbiaceae Aleurites 1 Argentina Breynia 1 S Brasil Mercurialis 1 Argentina Chile Ricinus 1 Argentina S Brasil Chile Paraguay Uruguay Rodriguésia 66(4):

89 1020 Zuloaga, F.O. & Belgrano, M.J. Family Genus Nº Spp. Countries Vernicia 1 Argentina Fabaceae Adenocarpus 1 Uruguay Amorpha 1 Argentina Uruguay Cajanus 1 S Brasil Paraguay Colutea 1 Argentina Coronilla 1 Argentina Cytisus 2 Argentina Chile Dipogon 1 Chile Galega 1 Argentina Chile Uruguay Genista 1 Argentina Lablab 1 Uruguay Laburnum 1 Argentina Leucaena 1 Argentina Medicago 9 Argentina S Brasil Chile Uruguay Melilotus 4 Argentina S Brasil Chile Paraguay Uruguay Neonotonia 1 Argentina Paraguay Ononis 1 Uruguay Psoralea 1 Uruguay Robinia 1 Argentina Chile Uruguay Spartium 1 Argentina Chile Uruguay Sphaerophysa 1 Argentina Trigonella 1 Argentina Chile Ulex 1 Argentina S Brasil Chile Uruguay Fumariaceae Fumaria 7 Argentina Chile Uruguay Gentianaceae Blackstonia 1 Argentina Uruguay Gentianaceae Irlbachia 1 Paraguay Hypoxidaceae Molineria 1 Argentina Iridaceae Belamcanda 1 Paraguay Chasmanthe 1 Argentina Crocosmia 1 Argentina Chile Freesia 2 Argentina Gladiolus 2 Argentina Iris 2 Argentina Chile Uruguay Romulea 1 Chile Sparaxis 2 Argentina Chile Watsonia 1 Argentina Juglandaceae Carya 1 Argentina Lamiaceae Ballota 1 Argentina Uruguay Galeopsis 1 Chile Glechoma 1 Argentina Chile Lamium 3 Argentina Chile Uruguay Leonotis 1 Argentina S Brasil Paraguay Uruguay Leonurus 2 Argentina S Brasil Paraguay Uruguay Rodriguésia 66(4):

90 Vascular Plants of the Southern Cone 1021 Family Genus Nº Spp. Countries Lycopus 1 Chile Marrubium 1 Argentina S Brasil Chile Uruguay Melissa 1 Argentina Chile Uruguay Mentha 4 Argentina S Brasil Chile Paraguay Uruguay Mesosphaerum 1 S Brasil Molucella 1 Argentina Chile Nepeta 1 Argentina Prunella 1 Argentina S Brasil Chile Uruguay Lemnaceae Landoltia 1 Chile Linderniaceae Torenia 1 S Brasil Malvaceae Alcea 1 Argentina Gossypium 1 Chile Malva 7 Argentina S Brasil Chile Paraguay Uruguay Meliaceae Melia 1 Argentina S Brasil Chile Paraguay Uruguay Montiaceae Claytonia 1 Argentina Moraceae Broussonetia 1 Argentina Myoporaceae Myoporum 1 Argentina Chile Uruguay Myrtaceae Eucalyptus 2 Argentina Nymphaeaceae Nelumbo 1 Paraguay Oleaceae Fraxinus 3 Argentina S Brasil Uruguay Ligustrum 2 Argentina Uruguay Syringa 1 Argentina Papaveraceae Chelidonium 1 Argentina Eschscholzia 1 Argentina Chile Glaucium 1 Argentina Papaver 5 Argentina Chile Uruguay Phrymaceae Mazus 1 Argentina S Brasil Pinaceae Pinus 3 Argentina Chile Pseudotsuga 1 Argentina Plantaginaceae Antirrhinum 2 Argentina Uruguay Cymbalaria 1 Argentina Chile Uruguay Digitalis 1 Argentina S Brasil Chile Uruguay Kickxia 1 Argentina Chile Uruguay Linaria 3 Argentina S Brasil Chile Uruguay Maurandya 3 Argentina S Brasil Poaceae Agropyron 1 Chile Aira 4 Argentina S Brasil Chile Uruguay Apera 1 Argentina Chile Arrhenatherum 1 Argentina Chile Arundo 1 Argentina S Brasil Chile Austrostipa 1 Argentina Avena 6 Argentina S Brasil Chile Uruguay Brachypodium 2 Argentina Chile Uruguay Rodriguésia 66(4):

91 1022 Zuloaga, F.O. & Belgrano, M.J. Family Genus Nº Spp. Countries Briza 3 Argentina S Brasil Chile Uruguay Catapodium 1 Argentina S Brasil Chile Uruguay Chrysopogon 1 Argentina Paraguay Coix 1 Argentina S Brasil Paraguay Corynephorus 2 Argentina Chile Cymbopogon 1 Argentina Chile Cynosurus 2 Argentina Chile Uruguay Dactylis 1 Argentina S Brasil Chile Uruguay Dactyloctenium 1 Argentina S Brasil Paraguay Uruguay Dichanthium 2 Argentina Paraguay Ehrharta 1 Argentina Elytrigia 1 Argentina Chile Gaudinia 1 Argentina Uruguay Hackelochloa 1 Argentina Paraguay Hainardia 1 Argentina Chile Uruguay Hemarthria 1 Argentina S Brasil Paraguay Uruguay Holcus 1 Argentina S Brasil Chile Uruguay Lachnagrostis 1 Argentina Chile Lagurus 1 Argentina S Brasil Chile Uruguay Lamarckia 1 Argentina Chile Leymus 1 Argentina Chile Megathyrsus 1 Argentina S Brasil Paraguay Uruguay Melinis 2 Argentina S Brasil Paraguay Uruguay Miscanthus 1 Chile Uruguay Moorochloa 1 Argentina Parapholis 2 Argentina Chile Uruguay Piptatherum 1 Argentina Chile Rottboellia 1 Argentina Schedonorus 1 Argentina Schismus 2 Argentina Chile Sclerochloa 1 Argentina Taeniatherum 1 Chile Themeda 1 Argentina Paraguay Thinopyrum 1 Argentina Polygonaceae Antigonon 1 Argentina Paraguay Emex 1 Argentina Chile Uruguay Eriogonum 1 Argentina Fallopia 1 Chile Primulaceae Centunculus 1 Argentina S Brasil Chile Paraguay Uruguay Proteaceae Grevillea 1 Argentina Ranunculaceae Aquilegia 1 Argentina Chile Ceratocephalus 1 Argentina Resedaceae Reseda 4 Argentina Chile Rodriguésia 66(4):

92 Vascular Plants of the Southern Cone 1023 Family Genus Nº Spp. Countries Rhamnaceae Hovenia 1 Argentina S Brasil Paraguay Rosaceae Cotoneaster 1 Argentina Crataegus 1 Argentina Duchesnea 1 Argentina S Brasil Chile Uruguay Eriobotrya 1 Argentina Malus 2 Argentina Pyracantha 1 Argentina Rosa 5 Argentina Chile Sanguisorba 1 Argentina Chile Sorbus 1 Argentina Spiraea 2 Argentina S Brasil Chile Rubiaceae Rubia 1 Argentina Chile Sherardia 1 Argentina Chile Uruguay Ruscaceae Cordyline 1 Argentina S Brasil Paraguay Uruguay Rutaceae Citrus 1 Argentina Paraguay Poncirus 1 Argentina Ruta 1 Argentina Chile Scrophulariaceae Scrophularia 1 Chile Verbascum 3 Argentina S Brasil Chile Paraguay Uruguay Simaroubaceae Ailanthus 1 Argentina Chile Uruguay Solanaceae Brugmansia 1 Argentina S Brasil Paraguay Tamaricaceae Tamarix 1 Argentina Theaceae Camellia 1 Argentina Urticaceae Soleirolia 1 Chile Valerianaceae Centranthus 1 Argentina Vitaceae Parthenocissus 2 Argentina Vitis 1 Argentina Zingiberaceae Hedychium 1 Argentina S Brasil Paraguay Conclusion The information compiled by these projects has proved the importance and richness of the flora of the temperate and cold-temperate ecosystems studied, the unique elements that integrate them and their relevance in terms of biodiversity. Therefore, continued projects to catalogue and study the vascular plants of meridional South America play an important part in the contribution towards the World Flora and, in a more immediate objective, in their input of data both to GBIF (Global Biodiversity Information Facility) and to the Sistema Nacional de Datos Biológicos of Argentina. It is important to highlight as a positive aspect of these projects the cooperation that was established at national and international level with similar projects in neighbouring countries of South America, such as the Flora de Chile (edited by C. Marticorena & R. Rodríguez at the Universidad de Concepción), Flora of Paraguay (edited by L. Ramella & P. Perret, at the Geneva Botánic Garden) and the Lista do Brasil (coordinated by R. Campostrini Forzza, at the Jardim Botanico do Rio de Janeiro), which include shared objectives and work methodology. As a synthesis, the development of projects to study plant diversity, uniting the work of botanists with tools such as databases, remote access to digitised herbarium specimens and bibliography are extremely beneficial because they lead to: Increment in plant knowledge and its availability for conservation purposes, providing a Rodriguésia 66(4):

93 1024 Zuloaga, F.O. & Belgrano, M.J. source of trusted and standardized data regarding Latin American botanical collections. Reinforcement of the collaboration between botanical institutes in South America. Strenghtening of the institutional activities, facilitating the informatization and digitisation processes of their collections and considerably increasing their visibility. Possibility to train young people in botanical collection management and informatization processing, producing high quality data. Facilitate taxonomic work by providing nonrestricted access to digitised botanical collections together with additional information comprising descriptions, photographs, illustrations, etc. Looking towards the future, the biggest challenge we face is to obtain a sustainable support, from both the economic and the human points of view. Floristic works are at a disadvantage when compared with other studies that are often punctual and have more immediate results in terms of time, and also by the preference given to researchers who publish their findings in high impact journals. This situation has brought to a diminishing number of botanists being dedicated to floristic and monographic works, likewise to the ongoing important tasks of collecting and identifying plants in general. Acknowledgements We wish to thank the editors of Rodriguésia and the external reviewers for their valuable contribution to improving this article. The project was funded by the National Council for Scientific and Technical Research of Argentina (CONICET, PIP ) and the National Geographic Society, grant References Anton, A.M. & Zuloaga, F.O. (eds.) Flora Fanerogámica Argentina. Fascículos Programa ProFlora (CONICET). Anton, A.M. & Zuloaga, F.O. (eds.). 2012a. Brassicaceae. Flora Argentina. Estudio Sigma, Buenos Aires. Vol. 8, pp Anton, A.M. & Zuloaga, F.O. (eds.). 2012b. Verbenaceae. Flora Argentina. Estudio Sigma, Buenos Aires. Vol. 14, pp Grande Allende, J.R Novitates Agrostologicae, IV. Additional segregates from Panicum incertae sedis. Phytoneuron 22: 1-6. Guerreiro, C. & Rúgolo de Agrasar, Z.E Two new species of Chusquea (Poaceae, Bambuseae) from northwestern Argentina. Systematic Botany 38: Hunziker, A.T. (ed.) Flora Fanerogámica Argentina. Fascículos Programa ProFlora (CONICET). O Leary, N. & Múlgura, M.E A taxonomic revision of the genus Phyla L. Annals of the Missouri Botanical Garden 98: Josse, C.; Navarro, G.; Comer, P.; Evans, R.; Faber- Langendoen, D.; Fellows, M.; Kittel, G.; Menard, S.; Pyne, M.; Reid, M.; Schulz, K.; Snow, K. & Teague, J Ecological systems of Latin America and the Caribbean: a working classification of terrestrial systems. NatureServe, Arlington. 47p. Tryon, R.M. & Tryon, A.F Ferns and allied plants, with special reference to Tropical America. Springer- Verlag, New York, Heidelberg, Berlin. 857p. Zuloaga, F.O.; Belgrano, M.J. & Anton, A.M (eds.). Solanaceae. Flora Argentina. Estudio Sigma, Buenos Aires. Vol. 13, pp Zuloaga, F.O.; Belgrano, M.J. & Anton, A.M. 2014a (eds.). Asteraceae, Anthemideae-Gnaphalieae. Flora Argentina. Estudio Sigma, Buenos Aires. Vol. 7, pars 1, pp Zuloaga, F. O.; Belgrano, M. J. & Anton, A. M. 2014b (eds.). Asteraceae, Senecioneae-Vernonieae. Flora Argentina. Estudio Sigma, Buenos Aires. Vol. 7, pars 3, pp Zuloaga, F.O. & Morrone, O. (eds.) Catálogo de las plantas vasculares de la República Argentina. I. Pteridophyta, Gymnospermae & Angiospermae (Monocotyledoneae). Monographs in Systematic Botany from the Missouri Botanical Garden 60: Zuloaga, F.O. & Morrone, O. (eds.) Catálogo de las plantas vasculares de la República Argentina. II. Angiospermae (Dicotyledoneae). Monographs in Systematic Botany from the Missouri Botanical Garden 64: Zuloaga, F.O.; Morrone, O. & Belgrano, M.J. (eds.) Catálogo de las plantas vasculares del Cono Sur (Argentina, Sur de Brasil, Chile, Paraguay y Uruguay). Monographs in Systematic Botany from the Missouri Botanical Garden 107: i-xcvi Zuloaga, F.O.; Nicora, E.G.; Rúgolo de Agrasar, Z.E.; Morrone, O.; Pensiero, J.F. & Cialdella, A.M Catálogo de la familia Poaceae en la República Argentina. Monographs in Systematic Botany from the Missouri Botanical Garden 47: Zuloaga, F.O.; Rúgolo, Z.E. & Anton, A.M. (eds.) Flora Argentina. Gráficamente Ediciones, Córdoba. 3(1) (Poaceae, Aristidoideae-Panicoideae): 1-588; 3(2) (Poaceae, Pooideae): Zuloaga, F.O.; Belgrano, M.J. & Anton, A.M. (eds.) Annonaceae (...). Flora Argentina. Estudio Sigma, Buenos Aires. Vol. 15, pp Artigo recebido em 07/07/2015. Aceito para publicação em 13/08/2015. Rodriguésia 66(4):

94 Rodriguésia 66(4): DOI: / Flora of Bolivia - where do we stand? RI. Meneses 1,2,7,8, S. Beck 1,3, E. García 1,3, M. Mercado 4, A. Araujo 5 & M. Serrano 6 Abstract The botanical exploration of Bolivia during the last two centuries did not leave a botanical legacy in the country. Only towards the end of the 20 th century Bolivia saw the start of the biology careers at its universities and the development of its own herbaria. Nowadays there are important herbaria in La Paz, Santa Cruz, Cochabamba and Sucre with collections ranging between 40,000 and 350,000 specimens. In 2014 a catalogue of the vascular flora of Bolivia was published under the auspices of the Missouri Botanical Garden, recording 15,345 species, of which 12,165 are native and 2,343 are endemic, while 694 are cultivated, 267 adventitious and 221 are naturalized. Endemic species of vascular plants add up to 2,343 species. The 286 families listed follow the APG III classification system. There are about 150 botanists in Bolivia interested in studying the country s rich flora. During a workshop organized in 2013 to promote a Flora of Bolivia, the participants established jointly a preliminary format for the taxonomic treatments. The Flora of Bolivia is planned to be an electronic, open access publication with international participation. The World Flora represents a challenge that must be tackled by circumscribing, verifying and recording all species known within our territory, and it is expected that it will have positive repercussions from and towards the ongoing Flora of Bolivia, in a similar way as the long running series of the Flora Neotropica has provided a wider picture that can be adapted and modified to fit our particular country. Key words: Flora, Bolivia, Chaco, Amazonia, endemics. Resumo A exploração botânica da Bolívia durante os últimos dois séculos não deixou um legado botânico no país. Apenas no final do século 20 o país começou a formar biólogos nas suas universidades e a desenvolver os seus acervos muselógicos. Atualmente existem herbários importantes em La Paz, Santa Cruz, Cochabamba e Sucre, com coleções acumulando entre 40 e 350 mil espécimes. O Catálogo da Flora Vascular da Bolívia, publicado em 2014 sob os auspícios do Missouri Botanical Garden, registrou espécies de plantas, das quais são nativas, enquanto 694 são cultivadas, 267 adventícias e 221 naturalizadas. Um total de espécies de plantas vasculares endêmicas foi registrado para o país. As 286 famílias listadas seguem o sistema de classificação APG III. Existem cerca de 150 botânicos ativos na Bolívia, cujo interesse é estudar a rica flora do país. Uma oficina foi organizada em 2013 com intuito de promover a Flora da Bolívia, durante a qual um formato preliminar para os tratamentos taxonômicos foi estabelecido pelos participantes. O intuito é de apresentar a Flora da Bolívia como uma publicação eletrônica e de acesso aberto ao público contando com participantes internacionais. Enquanto representa um grande desafio para a comunidade botânica mundial, a Flora do Mundo objetiva circunscrever, verificar e registrar todas as espécies conhecidas em nosso território, e espera-se que esse ambicioso projeto tenha repercussão positiva em relação à Flora da Bolívia, da mesma forma na qual a longa série Flora Neotropica tem gerado um esquema mais amplo que pode ser adaptado e modificado para servir ao nosso país em particular. Palavras-chave: Flora, Bolívia, Chaco, Amazônia, endêmica. 1 Herbario Nacional de Bolivia, Campus Universitario, Cota Cota, calle 27, Casilla 10077, La Paz, Bolivia. 2 Museo Nacional de Historia Natural. Cota Cota, calle 26, Casilla 8706, La Paz, Bolivia. 3 Instituto de Ecología, Universidad Mayor de San Andrés, La Paz, Bolivia. 4 Herbario Nacional Forestal Martín Cárdenas - Centro de Biodiversidad y Genética, Universidad Mayor de San Simón, Cochabamba. 5 Herbario Regional del Oriente, Universidad Autónoma Gabriel René Moreno, Santa Cruz. 6 Herbario del Sur, Universidad Mayor Real y Pontificia de San Francisco Xavier, Chuquisaca. 7 Correo Central Casilla 10077, Campus Universitario Cota Cota, calle 27, La Paz, Bolivia. 8 Author for correspondence: rosaiselameneses11@gmail.com

95 1026 Meneses, R. et al. Introduction Bolivia is among one of the most biodiverse countries in the world, home to around 17,000 species of vascular plants (Beck 1998). Highlighted by Lack (2012) as the only South American country that did not attempt to compile a complete floristic account until the end of the past century, as the in-country study of the Bolivian flora has started particularly late (Meneses et al. 2013). As an example, the Biology Course at the Universidad Mayor de San Andrés, La Paz, started only in 1972, while the Herbario Nacional de Bolivia (LPB) was created in 1984 and is only 31 years old. Other herbaria in the country are in a similar situation, contrasting with herbaria in neighbouring countries, such as the Instituto de Botánica Darwinion (SI) and the Museo de La Plata (LP), in Argentina, and the Jardim Botânico do Rio de Janeiro (RB) and Museu Nacional (R) in Brazil, all with over a hundred years of age and continuously developing collection and research activities, as well as a long tradition of forming taxonomists. These countries have created diverse initiatives to prepare local and regional floras and vegetation surveys (Hoehne ; Reitz 1965; Correa ; Klein 1978; Zuloaga & Morrone 1996,1999; Wanderley et al. 2001). A Flora is fundamental to enable a country to document its natural patrimony, be it as a guide for sustainable use of its plant resources or to inforce conservation measures to protect and manage plant diversity, being a data source for future basic or applied studies. Brief history of Bolivian botany The year of 1978 has marked the start of systematic floristic surveys in La Paz (LPB) through international collaboration between Bolivia, Germany and the United States, resulting in the collection of over 350,000 specimens of vascular plants (ferns, gymnosperms and angiosperms) and a smaller number of non vascular plants. These collections partially document the floristic richness of the country, while a group of more regional herbaria in Cochabamba, Santa Cruz and Sucre comprise respectively 60,000, 160,000 and 40,000 specimens (Tab. 1), complementing the overall picture. It is important to understand that the Herbario Nacional de Bolivia (LPB) was created as part of an agreement between the Instituto de Ecología de la Universidad Mayor de San Andrés and the Museo Nacional de Historia Natural, therefore it aims to incorporate valuable ecologic information to the floristic inventories, comprising auto and synecology data regarding species, as well as etnobotanical data in the form of use references. Such integration allows for a wider appreciation of the floristic diversity and also of the different vegetation types found in Bolivia, resulting in many publications and in recent comments on biogeography and evolution (Fernández et al. 2015; Maldonado et al. 2015; Gallegos et al. 2014; Saavedra et al. 2014; Apaza- Quevedo et al. 2013; Cuesta et al. 2012; Beck et al. 2010; Zenteno-Ruiz et al. 2009; Fuentes Claros 2006; Moraes et al. 2006). The recently published Catalogue of Vascular Plants (Jørgensen et al. 2014) represents an important milestone that will constitute the base for the preparation of the Flora of Bolivia. This publication took 15 years of intermittent work to be completed, and was carried under a broad collaboration from different institutions worldwide and published under the auspices of the Missouri Botanical Garden (MO), with the contribution of 221 specialists and 53 reviewers. The general plan for the Catalogue followed the system created for the Peru and Ecuador catalogues (Brako & Zarucchi 1993; Jørgensen & León-Yánez 1999). In Bolivia, 44 botanists from the Herbario Nacional de Bolivia (LPB), Herbario Nacional Forestal Martín Cárdenas (BOLV), Herbario Regional del Oriente (USZ) and the Herbario del Sur de Bolivia (HSB) were responsable for treatments at family or generic level. Table 1 Number of specimens deposited in major Bolivian Herbaria Department Herbaria Acronym Specimen La Paz Herbario Nacional de Bolivia LPB 350,000 Cochabamba Herbario Nacional Forestal Martín Cárdenas BOLV 60,000 Santa Cruz Herbario Regional del Oriente USZ 160,000 Chuquisaca Herbario del Sur de Bolivia HSB 40,000 Rodriguésia 66(4):

96 Flora of Bolivia - where do we stand? Preparation of the Flora of Bolivia As a basis for the preparation of the Flora of Bolivia, the published catalogue (Jørgensen et al. 2014) contains a synopsis of the vascular flora of Bolivia as it is known today, providing synonymy for species, bibliographic references for Bolivia, life-forms, origin, regional distribution, vegetation zones, altitudinal distribution and by geopolitical division, infraspecific taxa and common names when available. It is possible to access the electronic version in the webpage of the Missouri Botanical Garden (TROPICOS 2015). The data is managed as part of the TROPICOS, therefore it can be updated, added to and corrected in a dynamic way. The Bolivian catalogue documents 286 families arranged following the APG III (2009) classification system, comprising 15,345 species of which 12,165 are native species, while 694 are cultivated, 267 adventitious and 221 naturalized (Tab. 2), and 112 further putative species records not yet confirmed for the country. The number of endemic species currently recorded for Bolivia is 2,343, slightly less than 16% of the total species number. The present count departed from an initial total of 47,974 names recorded for Bolivia, amalgamating information from the catalogue of ferns and flowering plants of Bolivia (Foster 1958) and the Guía de árboles de Bolivia (Killeen et al. 1993) with records from Tropicos (2015) and The International Plant Names Index (IPNI 2015) either published from or with distribution referred to Bolivia. The major groups (Tab. 3) are 14,157 angiosperms, 1,145 ferns and 43 gymnosperms, most of the last introduced and/or cultivated. Orchidaceae, Asteraceae and Fabaceae 1027 contribute over 1,000 species and six of the largest genera (over 100 species), where Solanum alone comprises 192 species (Tab. 4, 5). Considering the 9 major geopolitical subdivisions, or departments (Fig. 1) of Bolivia, the four southwestern ones, La Paz, Oruro, Potosi and partly Cochabamba include the majority of the Andean highlands, while Chuquisaca, the eastern part of Cochabamba and La Paz are also home to the interandean valleys, Tarija, Chuquisaca and Santa Cruz are home to the Chaco (related to the Brazilian Pantanal), while Santa Cruz spans from Chaco in the south, Savanna in the far east, and transitional rainforest reaching the mostly Amazonian departments of Pando and Beni (CEDIB 2015). At present, La Paz features the highest number of species, while the entirely Amazonian department of Pando, with meagre 2,000 species records (Tab. 6), is likely to be undercollected. The same is true in the case of the vast Beni department (Tab. 5, Fig. 2) and the Andean highlands in Oruro and Potosi (Fig. 2), just to exemplify the areas showing sparse collecting activities in the map prepared from databased herbarium records. Even considering the harsh climatic conditions at high altitudes, Table 2 Vascular plants of Bolivia summarized according to category Category Nº species Accepted 15,345 Native 12,165 Endemic 2,343 Cultivated 694 Adventicious 267 Naturalized 221 *Several species are recorded in different categories, therefore the total number is higher than the accepted species number. Categorization according to Jørgensen et al Figure 1 Map of Bolivia highlighting the departaments. Rodriguésia 66(4):

97 1028 Meneses, R. et al. Andean altiplano highlands feature 1,565 species, while the Andean slopes 9,253 species, more than in the lowlands with 8,453 species (Jørgensen et al. 2014). A breakdown of species numbers by altitudinal quota (Tab. 7) shows the decrease in species number with altitude above two thousand meters and that, at the upper limit (above 5,000 m), 33 species have been recorded. Bolivia currently counts with approximately 150 botanists, mostly Biology graduates from the state universities of the Departments of La Paz, Cochabamba and Santa Cruz, but also agronomists and foresters from the universities in Beni, Chuquisaca and Tarija. Figure 2 Vascular specimen collections according to the LPB herbarium database, July Table 3 Vascular plant groups in Bolivia Groups Families Genera Species Ferns ,145 Gymnosperms 8 * Angiosperms 245 2,636 14,157 Total 286 2,782 15,345 * including 4 introduced families Table 4 Ten top families of vascular plants in Bolivia Families Nº species Orchidaceae 1,263 Asteraceae 1,256 Fabaceae 1,114 Poaceae 962 Malvaceae 445 Rubiaceae 429 Cyperaceae 348 Solanaceae 341 Melastomataceae 325 Euphorbiaceae 322 Towards a Flora of Bolivia The publication of the Bolivian catalogue edited by Jørgensen et al. (2014) has motivated the country s herbaria to pursue the organization of knowledge regarding botanical richness and diversity. These tasks are gaining momentum as it is understood that a Flora is important and most necessary as the baseline for basic and applied plant studies. A workshop Towards a Flora of Bolivia was organized in April 2013, with the participation of 40 botanists from Bolivia and other countries. The curators and representatives of the herbaria of Cochabamba (BOLV), Sucre (HSB), La Paz (LPB), Santa Cruz (USZ), Cobija, Oruro and Potosí were present alongside botanists that have supported the study of the Bolivian Flora for many years: the coordinator of the Madidi Project from the Missouri Botanical Garden (MBG), the author and editor of the Flora de Amboró from the New York Botanical Garden (NY), the coordinator of several Darwin Initiative projects from the Royal Botanic Gardens, Kew (K) and Oxford University and the Director of the Berlin Botanical Garden (BGBM). The results from the workshop were published by Meneses et al. (2013) to increase their visibility. Some of the conclusions from this meeting are summarized below. By consensus of all participants the Flora of Bolivia must be a publication of the highest scientific rigour and involving collaboration and communication with specialists worldwide. The task of preparing a flora is arduous for the botanists; the existence of a Flora of Bolivia will create the opportunity to develop and gather the scientific knowledge regarding the species, Rodriguésia 66(4):

98 Flora of Bolivia - where do we stand? Table 5 Ten top genera of vascular plants in Bolivia Genera N o species Solanum 192 Miconia 141 Epidendrum 128 Elaphoglossum 127 Peperomia 123 Tillandsia 104 Senecio 99 Thelypteris 96 Paspalum 95 Piper 93 Table 6 Species number and area of the 9 departments of Bolivia Departament Species number Approx. area (km 2 ) La Paz 9, ,000 Santa Cruz 7, ,000 Cochabamba 5,888 52,000 Beni 3, ,00 Chuquisaca 2,577 46,000 Tarija 2,554 35,000 Pando 2,001 60,000 Potosí ,000 Oruro ,000 thus facilitating future species identification, knowledge dissemination towards conservation and sustainable management of the plant resources. The workshop also provided an opportunity for the participants to establish the format for the taxonomic treatments, that was prepared in collaboration and accepted as a joint decision. The treatments of the Flora will include the accepted scientific name and selected synonyms according to the International Code for Nomenclature of Algae, Fungi and Plants (McNeill et al. 2012). The morphologic descriptions will be based in Bolivian specimens and include the most relevant characters compiled in a relatively short text ( words). Additional data will include habitat Altitude (m a.s.l.) Species number , ,000 4,972 1,000 1,500 4,316 1,500-2,000 3,902 2,000 2,500 3,536 2,500 3,000 3,331 3,000 3,500 2,650 3,500 4,000 1,855 4,000 4, ,500 5, > information, autoecologic information regarding the taxa (phenology, pollinators when known, etc.), the common names and known uses. Identification keys will include representative morphologic characters of vegetative and reproductive parts. Complementary ecologic information may be given in order to subsidise the species identification. A representative selection of up to 20 specimens will be used to fundament the specific or infraspecific taxonomic circumscription used in the flora. Distribution maps will contain georeferenced points based on specimens found in the available national and international databases containing Bolivian records after being examined by authors of the treatments. Illustrations will be provided to depict at least each genus, and may be either photographs or line-drawings. The classification will be based on up to date phylogenetic knowledge (APG III 2009) to order and plant family level. Generic classification will preferably consider monophyletic groups, bearing in mind that this level of information is not yet available for all plant groups. The Flora of Bolivia has been planned as an electronic, open access publication. The first fascicles will be prepared with groups comprising relatively few species, for which there are already Bolivian and foreign specialists engaged in studies, as for example families Amaranthaceae, Arecaceae, Lentibulariaceae, and selected groups of Orchidaceae and Poaceae. The participation of Bolivian botanists is strongly encouraged, however Table 7 Altitudinal distribution of the vascular plants of Bolivia Rodriguésia 66(4):

99 1030 Meneses, R. et al. we are aware that they are very few when taking into account the size of the task ahead. Bolivia has a positive track record with the Flora Neotropica Organization that helped the country, back in 1984, to found the Herbario Nacional de Bolivia (LPB), uniting collections, human capacity and infrastructure. The annual Flora Neotropica meeting in the Andes supported local herbaria and created new regional ones, enhancing the profile of botanical research in the country. Conclusion As all other tropical countries that are part of the World Flora, Bolivia needs to engage in the preparation of its national flora in order to contribute towards the final account at a global level. We do hope that the international scientific community and the national authorities are able to support the preparation of our Flora alongside the World Flora Project (Jackson & Miller 2015). Bolivian taxonomists could also contribute directly towards the World Flora, especially in particular groups such as the high-andean plants from the humid slopes of the Andes, highland wetlands and of the mostly endemic groups with a centre of diversity in the region, such as the Chloranthaceae, Cunoniaceae and Hypseocharitaceae. The contribution to the World Flora is seen as a positive challenge by the Bolivian botanical community. As the verification and recording of species started by the Bolivian Catalogue (Jørgensen et al. 2014) continues to be improved and the Bolivian taxa are better circumscribed, we seek to unite our knowledge across the borders into neighbouring countries. Our relatively young botanical institutions and community are keen to participate in the elaboration of widely accessible documents for all stakeholders interested in contributing towards the survival of humankind that is today threatened by the increasing loss of habitats, biodiversity and compromising of the Earth s ecologic processes. References Apaza-Quevedo, A.; Schleuning, M.; Hensen, I.; Saavedra, F. & Durka, W Forest fragmentation and edge effects on the genetic structure of Clusia sphaerocarpa and C. lechleri (Clusiaceae) in tropical montane forests. Journal of Tropical Ecology 29: APG III The Angiosperm Phylogeny Group III. (Brigitta Bremer, Kåre Bremer, Mark W. Chase, Michael F. Fay, James L. Reveal, Douglas E. Soltis, Pamela S. Soltis y Peter F. Stevens, además colaboraron Arne A. Anderberg, Michael J. Moore, Richard G. Olmstead, Paula J. Rudall, Kenneth J. Sytsma, David C. Tank, Kenneth Wurdack, Jenny Q.-Y. Xiang y Sue Zmarzty). An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants. Botanical Journal of the Linnean Society 161: APG III An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161: Beck, S.G Floristic inventory of Bolivia - An indispensable contribution to sustainable development. In: Barthlott, W. et al. (eds.). Biodiversity: a challenge for development research and policy. Springer, Heidelberg. Pp ISBN Beck, S.G.; Domic, A.; García, C.; Meneses, R.I.; Yager, K. & Halloy, S El Parque Nacional Sajama y sus plantas. Herbario Nacional de Bolivia. La Paz. 250p. Brako, L. & Zarucchi, J.L Catalogue of the flowering plants and Gymnosperms of Peru. Monographs in Systematic Botany from the Missouri Botanical Garden 45: i-xl, CEDIB Ecorregiones de Bolivia. Available at < ecorregiones/>. Access on 23 September Correa, M.N. (ed.). Flora patagónica. INTA, Buenos Aires. Pp Cuesta, F.; Muriel, P.; Beck, S.G.; Meneses, R.I.; Halloy, S.; Salgado, S.; Ortíz, E. & Becerra, M.T. (eds.) Biodiversidad y cambio climático en los Andes tropicales. Coformacion de un red de investigación para monitorear sus impactos y delinear acciones de adaptación. Red GLORIA- Andes, Lima-Quito. 180p. Fernández, M.; Navarro, L.M.; Apaza-Quevedo, A.; Gallegos, S.C.; Marques, A.; Zambrana-Torrelio, C.; Wolf, F.; Hamilton H.; Aguilar-Kirigin, A.J.; Aguirre, L.F.; Alvear, M.; Aparicio, J.; Apaza- Vargas, L.; Arellano, G.; Armijo, E.; Ascarrunz, N.; Barrera, S.; Beck, S.G.; Cabrera-Condarco, H.; Campos-Villanueva, C.; Cayola, L.; Flores-Saldana, N.P.; Fuentes, A.F.; García-Lino, M.C.; Gómez, M.I.; Higueras, Y.S.; Kessler, M.; Ledezma, J.C.; Limachi, J.M.; López, R.P.; Loza, M.I.; Macía, M.J.; Meneses, R.I.; Miranda, T.B.; Miranda Calle, A.B.; Molina-Rodriguez, R.F.; Moraes R.M.; Moya- Diaz, M.I.; Ocampo, M.; Perotto-Baldivieso, H.L.; Plata, O.; Reichle S.; Rivero, K.; Seidel, R.; Soria, L.; Terán, M.F.; Toledo, M.; Zenteno-Ruiz, F.S. & Pereira, H.M Challenges and opportunities for the BolivianBiodiversity Observation Network. Available at < >. Access on 1 October Rodriguésia 66(4):

100 Flora of Bolivia - where do we stand? Foster, R.C A catalogue of the ferns and flowering plants of Bolivia. Contribution of the Gray Herbarium 184: Fuentes Claros, A.F [2006]. Una introducción a la vegetación de la región del Madidi. Ecología de Bolivia 40: Gallegos, S.C.; Hensen, I. & Schleuning, M Secondary dispersal by ants promotes forest regeneration after deforestation. Journal of Ecology 102: Hoehne, F.C Flora Brasilica. Secretaria de Agricultura do Estado de São Paulo, São Paulo. IPNI The International Plant Names Index. Available at < Access 1 July Jørgensen, P.M. & León-Yánez, S. (eds.) Catalogue of the vascular plants of Ecuador. Monographs in Systematic Botany from the Missouri Botanical Garden 75: i-viii, Jørgensen, P.M.; Nee, M.H. & Beck, S.G. (eds.) Catálogo de plantas vasculares de Bolivia, Monographs in Systematic Botany from the Missouri Botanical Garden 127: i-viii, Killeen, T.; Garcia, E. & Beck, S.G. (eds.) Guía de árboles de Bolivia. Herbario Nacional de Bolivia, Missouri Botanical Garden, La Paz. 958p. Klein, R.M Mapa fitogeográfico do estado de Santa Catarina. In: Reitz, R. (ed.). Flora ilustrada catarinense. Herbário Barbosa Rodrigues, Itajaí. 24p. Lack, W Flora projects - a never ending synthesis. In: Lack, W. & Grotz K. (eds.), Flora s treasures - recording the green world. Botanisches Museum Berlin, Dahlem. Pp Maldonado, C.; Molina, C.I.; Zizka, A.; Persson, C.; Taylor, C.M.; Albán, J.; Chilquillo, E.; Rønsted, N. & Antonelli, A Estimating species diversity and distribution in the era of Big Data: to what extent can we trust public databases? Global Ecology and Biogeography 24: McNeill, J.; Barrie, F.R.; Buck, W.R.; Demoulin, V.; Greuter, W.; Hawksworth, D.L.; Herendeen, P.S.; Knapp, S.; Marhold, K.; Prado, J.; Prud homme Van Reine, W.F.; Smith, G.F.; Wiersema, J.H. & Turland, N.J. (eds.) International Code of Nomenclature for algae, fungi and plants (Melbourne Code) adopted by the Eighteenth International Botanical Congress Melbourne, 1031 Australia, July Regnum Vegetabile 154. Koeltz Scientific Books, Germany. 140p. Meneses R.I.; Borsch T.; Ortuño T. & Fuentes, A Hacia una flora de Bolivia. Revista de la Sociedad Boliviana de Botánica 7: Moraes R.M.; Øllgaard, B.; Kvist, L.P.; Borchsenius Kristensen, F. & Balslev, H. (eds.) Botánica económica de los Andes centrales. Universidad Mayor de San Andrés, La Paz. Pp Reitz, R Plano de coleção. In: Reitz, R. (ed.). Flora ilustrada catarinense. Herbário Barbosa Rodrigues, Itajaí. 71p. Saavedra, F.; Hensen, I.; Beck, S.G.; Böhning Gaese, K.; Lippok, D.; Töpfer, T. & Schleuning, M Functional importance of avian seed dispersers changes in response to human induced forest edges in tropical seed dispersal networks. Oecologia 176: DOI /s x. Tropicos Bolivia catalogue/catálogo de las plantas vasculares de Bolivia. Missouri Botanical Garden. Available at < Access on 1 July Wanderley, M.G.L.; Shepherd, G.J. & Giulietti, A.M Flora fanerogâmica do Estado de São Paulo. FAPESP/ HUCITEC, São Paulo. Pp Wyse Jackson, P. & Miller, J.S Developing a World Flora Online - a 2020 challenge to the world s botanists from the international community.rodriguésia 66: Zenteno-Ruiz, F.S.; Beck, S.G.; López, R.P. & Gallegos, S Caracterización de la vegetación y de la diversidad florística de la cuenca alta del río Bermejo, Tarija: implicaciones para su conservación. In: Beck, S.G. et al. (eds.). Biodiversidad y Ecología para la Conservación en Bolivia - Simposio de los 30 años del Instituto de Ecología. Instituto de Ecología, Universidad Mayor de San Andrés, La Paz. Pp Zuloaga, F.O. & Morrone, O. (eds.) Catálogo de las plantas vasculares de la República Argentina. I. Pteridophyta, Gymnospermae & Angiospermae (Monocotyledoneae). Monographs in Systematic Botany from the Missouri Botanical Garden 60: Zuloaga, F.O. & Morrone, O. (eds.) Catálogo de las plantas vasculares de la República Argentina. II. Angiospermae (Dicotyledoneae). Monographs in Systematic Botany from the Missouri Botanical Garden 64: Artigo recebido em 03/09/2015. Aceito para publicação em 25/09/2015. Rodriguésia 66(4):

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102 Rodriguésia 66(4): DOI: / Diversity of Brazilian Fungi Leonor C. Maia, Aníbal A. de Carvalho Júnior 1, Laise de H. Cavalcanti, Adriana de M. Gugliotta, Elisandro R. Drechsler-Santos, André L.M. de A. Santiago, Marcela E. da S. Cáceres, Tatiana B. Gibertoni, André Aptroot, Admir J. Giachini, Adriene M. da S. Soares, Allyne C.G. Silva, Altielys C. Magnago, Bruno T. Goto, Carla R.S. de Lira, Carlos A.S. Montoya, Carmen L.A. Pires-Zottarelli, Danielle K.A. da Silva, Dartanhã J. Soares, Diogo H.C. Rezende, Edna D.M.N. Luz, Emerson L. Gumboski, Felipe Wartchow, Fernanda Karstedt, Fernando M. Freire, Flávia P. Coutinho, Georgea S. N. de Melo, Helen M. P. Sotão, Iuri G. Baseia, Jadergudson Pereira, Jadson J.S. de Oliveira, João F. Souza, José L. Bezerra, Lídia S. Araujo Neta, Ludwig H. Pfenning, Luís F.P. Gusmão, Maria A. Neves, Marina Capelari, Melissa C.W. Jaeger, Melissa P. Pulgarín, Nelson Menolli Junior, Priscila S. de Medeiros, Raquel C.S. Friedrich, Renata dos S. Chikowski, Ricardo M. Pires, Roger F. Melo, Rosa M.B. da Silveira, Salomé Urrea-Valencia, Vagner G. Cortez & Valéria F. da Silva. Abstract Knowledge about the Brazilian fungal diversity was, until 2010, recorded in few taxonomy and ecology publications, as well as in a handful of species lists. With the publication of the Catálogo de Plantas e Fungos do Brasil and the continued availability of an online list, it has been possible to aggregate this dispersed knowledge. The version presented here adds 2,111 species names to the 3,608 listed in A total of 5,719 species of fungi distributed in 1,246 genera, 102 orders and 13 phyla represents a considerable increase over the last five years, when only 924 genera and 78 orders were registered. Basidiomycota (2,741 species in 22 orders) and Ascomycota (1,881 species in 41 orders) predominate over other groups. The Atlantic Rainforest has the largest number of records, with 3,017 species, followed by Amazon Rainforest (1,050), Caatinga (999), Cerrado (638) and Pampa and Pantanal with 84 and 35 species, respectively. The Northeast region has the greatest richness (2,617 species), followed by Southeast (2,252), South (1,995), North (1,301) and Central-West (488 species). Regarding the States of the Federation, São Paulo with 1,846 species, Pernambuco with 1,611 and Rio Grande do Sul with 1,377 species are the most diverse. Key words: Taxonomy, mycology, brazilian regions. Resumo Até 2010, o conhecimento sobre a diversidade de fungos do Brasil estava registrado em publicações esparsas de taxonomia e ecologia e em algumas poucas listas de espécies. Com a publicação do Catálogo de Plantas e Fungos do Brasil, e a disponibilização da lista online, tem sido possível agregar o conhecimento disperso. A versão ora apresentada acrescenta nomes de espécies aos listados em São citadas espécies de fungos distribuídas em gêneros, 102 ordens e 13 divisões, consistindo em considerável aumento em relação a 2010, quando estavam registrados 924 gêneros e 78 ordens. Predominam os Basidiomycota (2.741 espécies, em 22 ordens) e Ascomycota (1.881 espécies, em 41 ordens). A Mata Atlântica possui a maior quantidade de registros, com espécies, seguido pela Amazonia (1.050), Caatinga (999), Cerrado (638) e Pampa e Pantanal com 84 e 35 espécies, respectivamente. A região Nordeste tem a maior riqueza (2.617 especies), seguida pelo Sudeste (2.252), Sul (1.995), Norte (1.301) e Centro Oeste (488 espécies). Em relação aos Estados da Federação, São Paulo (1.846 espécies), Pernambuco (1.611) e Rio Grande do Sul (1.377) são os mais diversos. Palavras-chave: Taxonomia, micologia, regiões brasileiras. The full authorship of this article can be found in Appendix 1. 1 Author for correspondence: anibal@jbrj.gov.br

103 1034 Maia, L.C. et al. Introdução The publication of the Catálogo de Plantas e Fungos do Brasil (Forzza et al. 2010) was possible due to the voluntary participation of experts from various areas of Botany and Mycology. The catalogue constitutes a milestone in the systematization and dissemination of data on the diversity registered in Brazil. Besides the plants, organisms belonging to the kingdom Fungi (Ascomycota, Basidiomycota, Blastocladiomycota, Cryptomycota, Chytridiomycota, Entomophtoromycota, Glomeromycota and Zygomycota) and others, such as Hyphochytridiomycota, Labirinthulomycota, Myxomycota, Oomycota and Plasmodiophoromycota, currently classified in other kingdoms but traditionally studied by mycologists, have been treated and cited as Fungi sensu lato (Maia & Carvalho Jr 2010). The history of Brazilian mycology was described by Fidalgo (1968), who demonstrated that in the early stages of this discipline, the main collectors of the country s fungi were foreigners who sent material to be identified abroad. Among these, the best known are Montagne, Hennings, Bresadola and Rick, the latter considered the Father of Brazilian Mycology, due to his great contribution in this field. Arriving in Brazil in 1903, Rick initially asked for help from his foreign colleagues, sending specimens to herbaria outside, but from 1929 the collected material was studied and deposited in Brazilian herbaria, especially at the Anchieta herbarium (PACA), which has a fungal collection with around 13,000 specimens (Fungi Rickiani). After 1930, the situation changed and some isolated initiatives for knowledge of the country s fungi were published by researchers such as Viégas that, after 1930, released a series of papers on plant pathogenic fungi (Viégas 1939, 1940, 1943, 1944, 1945, 1946). Viégas better known work is the Index of Fungi of South America (Viégas 1961). Other important contributions came from Chaves Batista and colleagues, who published in the 1960s and 1970s more than 700 papers listed in Silva & Minter (1995) that are mostly available online ( fungibrasil.net/). After that, a list of Myxomycetes in São Paulo was provided by Hochgesand & Gottsberger (1996). Prior to the Catálogo de Plantas e Fungos do Brasil (Forzza et al. 2010), but already within this century, Góes Neto & Cavalcanti (2002) and Cavalcanti et al. (2006) presented respectively lists of Myxomycetes in the states of Bahia and Piauí; Cavalcanti (2002), Maimoni-Rodella (2002) and Putzke (2002) listed Myxomycetes occurring in different regions of the country; Curvo (2006) provided a list of fungi from Mato Grosso; Mendes et al. (1998, 2010) presented a list of species of fungi on plants in Brazil; Maia et al. (2002) showed the diversity of fungi in Pernambuco; Hennen et al. (2005) provided a catalogue of Brazilian rusts (Pucciniales); Gusmão & Maia (2006) showed the diversity of fungi in the semi-arid; Milanez et al. (2007) published a list of zoosporic fungi of Brazil; Maia et al. (2007) discussed the representation of fungi in Brazilian herbaria. Several lists on Agaricomycetes were also published: species in Brazil (Putzke 1994), in the Amazon Rainforest (Silva & Gibertoni 2009), in the Atlantic Rainforest (Baltazar & Gibertoni 2009), in the semiarid (Drechsler-Santos et al. 2009) and in the Cerrado (Gibertoni & Drechsler-Santos 2010). These publications considered specific groups (eg, plant parasitic) or focused on the diversity of fungi in some regions or states of the federation. Since Maia & Carvalho Jr. (2010), several articles were published, new listings were made available and more contributors continued to systematize information in the on-line database of the Brazilian List, which was open to improvement. Among these publications, included or not in the database, we found some related to: Pucciniales (Carvalho Jr. & Hennen 2010, 2012; Salazar-Yepes & Carvalho Jr. 2010a, b, 2012, 2014); conidial fungi, with new genera, species and records for Brasil (Almeida et al. 2011, 2014; Barbosa & Gusmão 2011; Izabel et al. 2011; Cruz et al. 2012; Almeida et al. 2013; Barbosa et al. 2013; Fiuza et al. 2014; Monteiro & Gusmão 2013; Monteiro et al. 2014a, b); Ingoldian fungi (Fiuza & Gusmão 2013a, 2013b; Fiuza et al. 2015), zoosporic fungi (Nascimento et al. 2012; Jesus et al. 2013), new species and records of Myxomycetes (Alves et al. 2010; Bezerra et al. 2010, 2014; Bezerra & Cavalcanti 2010; Silva & Cavalcanti 2010; Costa et al. 2011, 2014; Damasceno et al. 2011; Silva & Cavalcanti 2012; Parente & Cavalcanti 2013; Cavalcanti et al. 2014, 2015), new species and records of Zygomycota (Santiago et al. 2011a, b, 2013, 2014; Souza et al. 2012, 2014); new orders, families, genera, species, recombinations and review in Glomeromycota (Goto et al. 2011, 2012, 2013; Mello et al. 2012, 2013; Oehl et al. 2011a, b; Pontes et al. 2013), including phylogenetic studies of Gigasporales (Silva et al. 2013). In Ascomycota: new genus Rodriguésia 66(4):

104 Diversity of Brazilian Fungi and species of histeriaceous fungi (Almeida et al. 2014), generic reviews of: Ascobolus and Saccobolus (Melo et al. 2014), Diorygma (Feuerstein et al. 2014), Polymeridium (Aptroot & Cáceres 2014c); new genera: Anabahusakala (Carmo et al. 2014); Helicodochium (Monteiro et al. 2014a); Ypsilomyces (Almeida & Gusmão 2014); Ellisembiopsis (Izabel et al. 2013); Anacraspedodidymum (Silva et al. 2014); new records, genera and species of liquenized fungi (Alves et al. 2014; Aptroot et al. 2013; Aptroot & Cáceres 2014a, b; Cáceres et al. 2012, 2013; Feuerstein et al. 2014; Lima et al. 2013a, b; Menezes et al. 2013). In Basidiomycota: new species and records of Agaricomycetes (Abrahão et al. 2012; Baltazar et al. 2012; Cabral et al. 2012; Coimbra & Gibertoni 2015; Coimbra et al. 2012, 2013; Cortez et al. 2011; Drechsler-Santos et al. 2012a, b, c, 2013; Gibertoni et al. 2012; Gomes-Silva et al. 2012, 2013, 2014; Sá et al. 2014; Soares et al. 2014; Trierveiler- Pereira et al. 2011; Wartchow et al. 2013a, b), as well as approaches to some genera: Coltricia (Baltazar et al. 2010), Phellinus and Inonotus (Baltazar & Gibertoni 2010), Trichaptum (Gibertoni et al. 2011), Diplomitoporus (Baltazar et al. 2013), Pleurotus (Menolli Jr. et al. 2014), Henningsia (Gibertoni & Ryvarden 2014), among others. After 2010, some checklists were also published, including: Glomeromycota of the Caatinga (Maia et al. 2010) and semiarid region (Goto et al. 2010); Agaricomycetes of the Cerrado (Gibertoni & Drechsler-Santos 2010; Abrahão et al. 2012); fungi of the Serra da Jibóia, Bahia (Barbosa et al. 2014); coprophilous fungi (Calaça et al. 2014) and a book on macrofungi of the semiarid region (Neves et al. 2013). In recent years the taxonomic placement of fungi has undergone major changes, especially after the work of Hibbett et al. (2007), with more than 60 colleagues, who proposed a phylogenetic classification for the kingdom. Throught the use of this new approach, the classification of many groups of fungi changed at different taxonomic levels. These analyses accelerated the discovery of new genera and species, creating new families, rearrangement of orders, classes and phyla in an attempt to group taxa phylogenetically to produce a more consistent classification. In this context, the listing process also involves vastly updating the nomenclature of species and moving them to their current position, which involves extra work. Taking as example the Glomeromycota, a phylum proposed by Schüssler et al. (2001) to include only arbuscular mycorrhizal fungi, which were previously included in Zygomycota In 2001 this group was divided into four orders and seven families, comprising about 150 species; today the phylum is represented by three classes, five orders, 15 families and 38 genera, with more than 250 species, of which 157 occur in Brasil. Among these, more than 20 were originally described from material collected in Brazil (Goto et al. 2011, 2012, 2013; Mello et al. 2012, 2013; Marinho et al. 2014). Another significant change occurred in Zygomycota, disregarded for being polyphyletic (Hibbett et al. 2007). The species formerly placed in this group are currently classified in Entomophthoromycota (Humber 2012) and four subphyla of uncertain position: Mucoromycotina, Zoopagomycotina, Kickxellomycotina and Mortierellomycotina (Hibbet et al. 2007; Hoffmann et al. 2011). However, considering that the subphylum category was not included in the present work, part of the species of the group were kept in the previously considered phylum Zygomycota, while the rest is included in Entomophthoromycota. This information shows how quickly the classifications have changed and how the new findings contribute to our work, considering the wealth of fungal species yet to be unveiled. Methodology The method used in this study was the same adopted in Forzza et al. (2010) and Maia & Carvalho Jr. (2010), and the work was improved by invited experts who used the on-line database available at the web page of Jardim Botânico do Rio de Janeiro. Each expert had the task of including new records and modifying data from previous records, in the light of the latest knowledge to their speciality group. The coordinators of the group of fungi were tasked with reviewing and complementing data which eventually have not been filled in by the specialists. The dataset finalized in March 2015, was used to prepare the present analyses, and can be found at the Brazilian List of Fungi sensu lato (see supplementary material < dx.doi.org/ /m9.figshare > - DOI: / ). Only the checked and accepted names of fungi with the respective authors were used as base for inclusion of data in the present article. Due to the work required, this review was not complete although most of the data have been included. Experts that contributed to this data collection, listed in alphabetical order of the first author: Aptroot, A. (Ascomycota-Acarosporales, Baeomycetales, Lichinales, Mycocaliciales); Aptroot, A., Cáceres, Rodriguésia 66(4):

105 1036 Maia, L.C. et al. M. (Ascomycota-Arthoniales, Candelariales, Ostropales, Peltigerales, Pertusariales, Pyrenulales, Teloschistales, Trypetheliales, Verrucariales); Aptroot, A., Gumboski, E.L., Cáceres, M. (Ascomycota-Lecanorales); Baseia, I.G., Cortez, V.G. (Basidiomycota-Geastrales, Hysterangiales, Phallales); Bezerra, J.L. (Ascomycota- Helotiales, Pezizales); Bezerra, J.L., Soares, D.J. (Ascomycota-Meliolales, Rhytismatales); Bezerra, J.L., Soares, D.J., Aptroot, A., Coutinho, F., Melo, R.F. (Ascomycota-Dothideales); Bezerra, J.L., Soares, D.J., Pfenning, L. (Ascomycota- Phyllachorales); Bezerra, J.L., Soares, D.J., Pfenning, L., Drechsler-Santos, E.R., Palácio, M., Freire, F.M., Friedrich, R.C.S., Gusmão, L.F.P. (Hypocreales); Cáceres, M. (Ascomycota-Incertae sedis); Capelari, M., Cortez, V.G., Neves, M.A., Baseia, I.G., Wartchow, F., Menolli Júnior, N., Karstedt, F., Oliveira, J.J.S., Urrea-Valencia, S. (Basidiomycota Agaricales); Carvalho Jr., A. A. de (Basidiomycota Urocystidiales); Carvalho Jr., A. A. de, Sotão, H.M.P. (Basidiomycota-Pucciniales), Cavalcanti, L.H. (Myxomycota-Ceratiomyxales, Echinosteliales, Liceales, Physarales, Stemonitales, Trichiales); Bezerra, J.L., Coutinho, F. (Ascomycota- Asterinales, Microthyriales); Drechsler-Santos, E.R., Melo, G.S.N., Palácio, M., Gomes-Silva, A.C. (Basidiomycota Gloeophyllales); Giachini, A., Silveira, R.M.B., Drechsler-Santos, E.R. (Basidiomycota-Gomphales); Gibertoni, T.B., Gomes-Silva, A.C., Chikowski, R.S., Lira, C.R.S., Soares, A.M.S., Melo, G.S.N., Araújo Neta, L., Gugliotta, A.M., Medeiros, P.S.,Silva, V.F., Silveira, R.M.B., Drechsler-Santos, E.R., Montoya, C.A.S. (Basidiomycota Hymenochaetales) Gibertoni, T.B., Neves, M.A., Wartchow, F., Chikowski, R.S., Silveira, R.M.B. (Basidiomycota-Cantharellales); Goto, B.T., Maia, L.C. (Glomeromycota- Paraglomerales), Goto, B.T., Maia, L.C., Silva, D.K.A. (Glomeromycota-Archaeosporales, Diversisporales, Glomerales); Gugliotta, A.M., Gibertoni, T.B., Drechsler-Santos, E.R., Silveira, R.M.B., Chikowski, R.S., Pires, R.M., Montoya, C.A.S., Souza, J.F., Palácio, M., Rezende, D.H.C. (Basidiomycota-Polyporales); Gumboski, E. L. (Ascomycota-Lecanorales); Gusmão, L.F.P., Melo, R. (Ascomycota-Sordariales); Gusmão, L.F.P., Pfenning, L. (Incertae sedis); Neves, M.A., Magnago, A.C. (Basidiomycota- Boletales); Neves, M.A., Gibertoni, T.B., Jaeger, M.C.W., Melo, G.S.N., Gomes-Silva, A.C., Araújo Neta, L., Wartchow, F., Chikowski, R.S., Silveira, R.M.B. (Basidiomycota-Russulales); Pfenning, L., Gusmão, L.F.P. (Ascomycota- Eurotiales); Pereira, J. (Ascomycota-Xylariales); Pires-Zottarelli, C.L.A. (Hyphochytriomycota- Hyphochytriales; Labyrinthulomycota- Thraustochytriales; Oomycota-Albuginales, Haptoglossales, Incertae sedis, Leptomitales, Myzocytiopsidales, Olpidiopsidales, Pythiales, Rhipidiales, Rozellopsidales, Saprolegniales; Plasmodiophoromycota-Plasmodiophorales; Blastocladiomycota-Blastocladiales; Chytridiomycota-Chytridiales, Incertae sedis, Lobulomycetales, Monoblepharidales, Rhizophlyctidales, Spizellomycetales; Cryptomycota); Pires-Zottarelli, C.L.A., Luz, E.D. (Oomycota-Peronosporales); Santiago, A.L.C.. (Entomophthoromycota-Entomophthorales; Zygomycota Dimargaritales, Endogonales, Mortierellales, Mucorales, Zoopagales); Silva, D.K.A., Silva, G.A., Maia, L.C. (Glomeromycota- Gigasporales); Silveira, R.M.B. (Basidiomycota- Atheliales, Auriculariales, Corticiales, Thelephorales); Soares, D.J., Coutinho, F., Melo, R.F. (Ascomycota-Capnodiales, Diaporthales, Erysiphales, Doassansiales); Soares, D.J., Coutinho, F., Melo, R.F., Gusmão, L.F.P. (Ascomycota- Pleosporales). Considering that information on distribution of the occurrence and research in the various biomes and Brazilian ecosystems is still insufficient, it was impossible to determine with certainty which species were endemic and threatened taxa; therefore this aspect was not addressed in this study. Results & Discussion From Maia & Carvalho Jr. (2010), there has been gradual increase in the number of fungal species registered, that culminated with the complete dataset found at the Brazilian List of Fungi sensu lato (see supplementary material < dx.doi.org/ /m9.figshare > - DOI: / ). There is a greater increase in the contribution of species records between the years 2010 and In 2010, 3,608 species were recorded and until of 2015, more 2,111 species, reaching a final count of 5,719 species names were added to this new version. Species of Basidiomycota (2,741 species) and Ascomycota (1,881 species) predominated in the current list (Tab. 1) as was expected given that these are the two major groups of fungi, respectively with more than 30 and 60 Rodriguésia 66(4):

106 Diversity of Brazilian Fungi Table 1 Number of fungal species, by phylum, registered in the Brazilian List of Plants and Fungi in 2010 and Phylum Number of fungal species Basidiomycota Ascomycota Incertae sedis Myxomycota Oomycota Glomeromycota Chytridiomycota Zygomycota Blastocladiomycota Cryptomycota 0 5 Entomophthoromycota 0 5 Hyphochytriomycota 4 5 Labyrinthulomycota 4 4 Plasmodiophoromycota 4 4 thousand species already described (Kirk et al. 2008). For other phyla, the number of species described worldwide is very low, ranging from less than 30 (Hyphochytridiomycota) to about 1,100 (Chytridiomycota, Myxomycota, Oomycota and Zygomycota, for example). In the range of 50 to 250 species are the Labirinthulomycota (48), Plasmodiophoromycota (47) and Glomeromycota (250). The representativeness of these phyla in Brazil is not fully known, neither is Ascomycota and Basidiomycota, but almost the totality of what was recorded in the country for these smaller groups is now listed. Of the 250 species of Glomeromycota in the world, for example, 157 are cited for Brazil, which represents approximately 66% of the total, but for some groups this ratio is much lower. Regarding the Myxomycetes, the ratio is less than 30% for the total of 900 species currently accepted, although Brazil is the second country in the Neotropics and the first in South America in number of records (considering as yet that the species recorded in Pantanal and Pampa are still not included in the present dataset); for Chytridiomycota there is about 10% of the registered in the world, while 1037 for Zygomycota the proportion is less than 5%, indicating the need for more experts and work dedicated to these groups. The Ascomycota teleomorphs were initially more addressed in studies of leaf fungi by Batista et al. (Silva & Minter 1995; Batista 2015), and are now being investigated further by experts in plant pathology. As a highlight of the latest studies with teleomorphs are those performed with pathogenic, coprophilous and lichenized fungi. However, much of what has been recorded, especially in the Central-West and Southeast regions, has not yet been recorded in the present list. Ascomycota anamorphic were also a well studied group among leaf fungi and continue to be investigated under several aspects: taxonomic, ecological, phytopathological and biotechnological. A large portion of the taxa is as yet without a defined taxonomic position and they appear in insertae sedis groups in the present list. In Brazil, few experts are dedicated to the Ascomycota, and, considering that this is the largest group of fungi, it is understandable the need for increasing specialized staff training. The Basidiomycota have more experts and, as many form macroscopic, often showy structures, are better studied and known. This group also includes the rusts, important parasites of plants, which have attracted the attention of a few researchers. For other phyla, the number of specialists is even smaller or nonexistent in Brazil and it is clear the need for training human resources in taxonomy in order to investigate and record the occurrence of the country s fungi. Some groups are barely known in Brazil as the Blastocladiomycota with 17 species recorded, Cryptomycota, Entomophthoromycota, Hyphochytriomycota, Labyrinthulomycota and Plasmodiophoromycota with five or fewer species. The 2015 list includes 13 phyla (plus a taxonomic group placed as insertae sedis), 102 orders (another insertae sedis group), 1,246 genera and 5,719 species of fungi. These numbers represent a considerable increase when compared to 2010, when 78 orders and 924 genera were recorded (Maia & Carvalho Jr. 2010). Of the 102 orders, 41 belong to Ascomycota and 22 to Basidiomycota. In Ascomycota, the six orders with higher number of species are Xylariales (275 species), Asterinales (248) Lecanorales (219), Ostropales (162) Microthyriales (120) and Capnodiales (99) (Tab. 2). In Basidiomycota, the orders with higher species numbers are Agaricales (927 species), Pucciniales (750) Polyporales (453), Hymenochaetales (166), Russulales (137) and Boletales (92) (Tab. 2). Rodriguésia 66(4):

107 1038 Maia, L.C. et al. Figure 1 Increase in the number of species registered in the Brazilian phythogeographical domains from 2010 to Table 2 Increase in the number of species recorded in the six more representative Orders of Ascomycota and Basidiomycota in the Brazilian List of Plants and Fungi comparing 2010 with Phylum/Ordem Number of fungal species Ascomycota Xylariales Asterinales Lecanorales Ostropales Microthyriales Capnodiales 4 99 Basidiomycota Agaricales Pucciniales Polyporales Hymenochaetales Russulales Boletales Most of the fungi mentioned in the current list are saprobes, but parasites and symbionts also stand out. Among the plant parasitic species, rusts (Pucciniales) are the second order better represented among the Basidiomycota and among the Ascomycota the second order with more records is Asterinales. Microthyriales and Capnodiales, with a significant number of species already listed, also add many plant pathogenic species. Other groups, Figure 2 Number of fungal species recorded in each Brazilian region in 2010 and Table 3 Distribution of the 21 richest genera of fungi in number of species recorded in the first version of the Brazilian List of Plants and Fungi (Maia & Carvalho Jr. 2010) and the current number (2015). The richest Generaof fungi Number of fungal species Puccinia Marasmius Xylaria Cladonia Uromyces Lepiota Uredo Asterina 0 73 Aecidium Scolecopeltidium 0 65 Asteridiella 1 53 Phellinus Physarum Glomus Ravenelia Pythium Pluteus 6 39 Lembosia 0 38 Penicillium Entoloma 9 37 Prospodium Total Rodriguésia 66(4):

108 Diversity of Brazilian Fungi such as Oomycota and Plasmodiophoromycota are also represented by several species that attack plants. Equally important is to record symbionts, consisting of lichens, or lichenized fungi (most Ascomycota, mainly represented by Lecanorales) and mycorrhizal species (all orders of Glomeromycota, several Basidiomycota and a few Ascomycota and Zygomycota, specifically Endogonales). The distribution of the 21 richest genera of fungi in number of species recorded in 2010 and the comparison with the current number is presented in Table 3. There was no significant increase in the number of species in some genera such as Puccinia, Uromyces and Uredo because the vast majority of records of these taxa in the country was already included in the former edition of the list (Maia & Carvalho 2010). Conversely, previously not recorded Asterina and Cladonia passed respectively to 73 and 113 recorded species, and Marasmius increased from two species to 128 species records in the country. This increase is due to the participation of more experts who accepted the invitation to contribute to the list between 2010 and 2015, to the study of some taxa such as Marasmius, a genus revised during this period, to the discovery of new species to science, to new collections being carry out in areas previously poorly studied (as the lichens in Roraima, for example), and due to an increase in the species records already known for Brazil, with expansion of the known geographical distribution. Thus, the inclusion of genera and species that had not yet been recorded in 2010 led to changes in classification of the richest genera, resulting in a new configuration. The Atlantic Rainforest, increasing in 1,353 species since 2010 and currently with 3,017 species recorded, remains the better known and most investigated biome. On the other hand, the Amazon Rainforest increased in 531 species since 2010, currently with 1,050 species, surpassing the number of species records from the Caatinga, which has 999 taxa. The Cerrado, currently with 638 records has increased 347species since Pampa currently with 84 and Pantanal with 35 species of fungi also reversed the positions compared to 2010 (Fig. 1). The Northeast region remains on the lead in terms of number of records, totalling 2,617 species, followed by the Southeast (2,252), South (1,995), North (1,301) and Central-West (488), each of these regions were increased by , 841, 675, 558 and 192 respectively since 2010 (Fig. 2). Regarding the distribution of the fungi in the Brazilian states, there was a significant contribution of the number of species compared to 2010 (Tab. 4) and highlight is given to the number of records for São Paulo and Pernambuco, followed by Rio Grande do Sul, Bahia, Amazonas, Paraná and Santa Catarina. Table 4 Number of fungal species, lato sensu and stricto sensu, by State (in descending order of records) in the first version of the Brazilian List of Plants and Fungi (Maia & Carvalho Jr. 2010), and the current number (2015). States Number of fungal species São Paulo Pernambuco Rio Grande do Sul Bahia Amazonas Paraná Santa Catarina Rio de Janeiro Minas Gerais Pará Paraíba Alagoas Roraima Sergipe Ceará Rio Grande do Norte Piauí Roraima Mato Grosso Maranhão Goiás Distrito Federal Amapá Mato Grosso do Sul Acre Espírito Santo Tocantins 5 25 Rodriguésia 66(4):

109 1040 Maia, L.C. et al. With smaller representation of fungi are, in descending order: Goiás, Amapá, Mato Grosso do Sul, Acre, Tocantins and Espírito Santo. The records for Tocantins are from recent collections. Older samplings are possible registered for the state of Goiás, of which Tocantins was a part until These states of the Central-West region need to be more investigated to survey their mycota. The Federal District also has few records of fungi (about 160), although mycologists at the University of Brasilia have been working in the region in recent decades. This indicates that much of what is studied is not available in the virtual herbarium and that we need a greater effort of mycologists to join the community that worked in the presently list, including the fungi identified during their taxonomic studies. Table 4 shows that, compared to 2010, there was a significant contribution of the number of fungal species registered in all federal units. Conclusions The great effort made by mycologists in the last five years resulted in increased completeness of the Fungal list in Despite the hard work of mycologists, the reality is that, as already highlighted by Maia & Carvalho Jr. (2010), the information available derives from samples collected in regions where there are more active groups of mycologists. For certain Brazilian states such as Tocantins, data are practically non-existent. Relevant contribution was recently given by Marcela Cáceres group, who investigated lichenized fungi in Rondônia, adding 104 new records for the state, of which 75 are new to science (Aptroot & Cáceres 2014a, 2014b; Cáceres et al. 2014a, 2014b, 2014c). This example highlights the potential for mycological studies and demonstrates that much field and laboratory work must still be carried out to obtain a closer estimate of the number of species of Brazilian fungi. Despite the significant increase in the number of occurrence records of fungi for Brazil in the last five years, the number of cited species does not reflect the totality of what potentially exists in the country. Several publications, among which state lists (Maia et al. 2002; Meijer 2008); regional lists (Gusmão & Maia 2006) and general lists (Putzke 1994; Mendes et al. 1998, 2010.), where approximately 7,000 taxa are mentioned could not be fully incorporated at the moment, among other reasons due to limitations of staff and time. Numerous fungal records in herbaria and culture collections are not yet recorded within this list, and these collections are important sources of information that should be included in subsequent editions. Considering the latest estimates that quote 5.1 million of fungal species worldwide (Blackwell 2011) and that, by 2008, slightly less than 100,000 species were described (Kirk 2008), the current knowledge approaches 2% of the estimated diversity of the group. It appears that, despite the efforts and the information that may be included in future projects, there is still a long way to go in terms of developing a basic knowledge of the diversity of the Brazilian mycota. The database of fungi from the 2015 list is still preliminary, but it represents a rare opportunity to increase the effective systematization of data that were scattered in various publications and to provide information on the fungi that occur in the country. To increase knowledge about the occurrence and distribution of this important group of organisms we need the help of a vast number of mycologists, to intensify inventories, taxonomic studies and training of specialized human resources to cover the insufficiently surveyed areas, which are numerous, and disseminate more effectively the mycological knowledge. Acknowledgements We thank our botanical colleagues, for the opportunity to participate, that allowed the inclusion of fungi in this project. Paula Leitman and Fabiana Luiza Ranzato Filardi, for their patience and dedicated assistance. Special thanks are due to CNPq, which has funded research that underlie the data analysed here through various projects (Protax, Reflora, Sisbiota, Universal, INCT-Herbário Virtual and others). References Abrahão, M.C.; Gugliotta, A.M. & Bononi, V.L.R Xylophilous Agaricomycetes (Basidiomycota) of the Brazilian Cerrado. Check List 8: Almeida, D.A.C.; Izabel, T. dos S.S.& Gusmão, L.F.P Fungos conidiais do Bioma Caatinga. I. Novos registros para o Continente americano, Neotrópico, América do Sul e Brasil. Rodriguesia 62: Almeida, D.A.C.; Cruz, A.C.R.; Marques, M.F.O. & Gusmão, L.F.P Conidial fungi from semiarid Caatinga biome of Brazil. New and interesting Zanclospora species. Mycosphere Online - Journal of Fungal Biology 4: Rodriguésia 66(4):

110 Diversity of Brazilian Fungi Almeida, D.A.C. & Gusmão, L.F.P Ypsilomyces, a new thallic genus of conidial fungi from the semi-arid Caatinga biome of Brazil. Mycotaxon 129: , Almeida, D.A.C.; Gusmão, L.F.P. & Miller, A.N A new genus and three new species of hysteriaceous ascomycetes from the semiarid region of Brazil. Phytotaxa 176: Alves, M.H.; Costa, A.A.A. & Cavalcanti, L.H Myxomycetes, state of Ceará, northeastern Brazil. Check List 6: Alves, M.M.E.; Aptroot, A.; Lacerda, S.R. & Cáceres, M.E S A new Eschatogonia species and two new Gassicurtia species from Chapada do Araripe, Ceará, NE Brazil. The Bryologist 117: Aptroot, A.; Ertz, D.; Lima, E.L.; Jesus, K.A.; Maia, L.C. & Cáceres, M.E.S Two new species of Roccellaceae (Ascomycota: Arthoniales) from Brazil, with the description of the new genus Sergipea. The Lichenologist 45: Aptroot, A. & Cáceres, M.E.S. 2014a. New lichen species from termite nests in rainforest in Brazilian Rondônia and adjacent Amazonas. Lichenologist 46: Aptroot, A. & Cáceres, M.E.S. 2014b. A key to the corticolous microfoliose, foliose and related crustose lichens from Rondônia, Brazil, with the description of four new species. Lichenologist 46: Aptroot, A. & Cáceres, M.E.S. 2014c. A refined species concept in the tropical lichen genus Polymeridium (Trypetheliaceae) doubles the number of known species, with a worldwide key to the species. Nova Hedwigia 98: Baltazar, J.M. & Gibertoni, T.B A checklist of the aphyllophoroid fungi (Basidiomycota) recorded from the Brazilian Atlantic Forest. Mycotaxon 109: Baltazar, J.M. & Gibertoni, T.B New combinations in Phellinus s.l. and Inonotus s.l. Mycotaxon 111: Baltazar, J.M.; Ryvarden, L. & Gibertoni, T.B The genus Coltricia in Brazil: new records and two new species. Mycologia 102: Baltazar, J.M.; Drechsler-Santos, E.R.; Ryvarden, L.; Cavalcanti, M.A.Q. & Gibertoni, T.B Contribution to the knowledge of polypores (Agaricomycetes) from the Atlantic Forest and Caatinga, with new records from Brazil. Mycosphere Online - Journal of Fungal Biology 3: Baltazar, J.M.; Ryvarden, L. & Gibertoni, T.B Diplomitoporus (Polyporales, Basidiomycota) in Brazil revisited. Mycological Progress 13: Barbosa, F.R. & Gusmão, L.F.P Conidial fungi from the semi-arid Caatinga biome of Brazil. Rare freshwater hyphomycetes and other new records. Mycosphere 2: Barbosa, F.R.; Raja, H.A.; Shearer, C.A. & Gusmão, L.F.P Some freshwater fungi from the Brazilian Semi-Arid Region, including two new species of Hyphomycetes. Cryptogamie. Mycologie 34: Barbosa, F.R.; Machiner, M.; Barbosa, G.C.K. & Gusmão, L.F.P A checklist of the fungi recorded from Serra da Jibóia, Bahia state, Brazil. Mycotaxon 129: Batista, A.C Publicações. Available in < batista.fungibrasil.net/>. Access on 6 May Bezerra, A.C.C. & Cavalcanti, L.H Diderma albo-columella (Myxomycetes) a new species in the Brazilian Atlantic Forest. Rodriguesia 61: Bezerra, M.F.; Farias, G.B. & Cavalcanti, L.H Mixobiota do Parque Nacional Serra de Itabaiana, SE, Brasil: Trichiales. Acta Botanica Brasilica 24: Bezerra, A.C.C.; Lima, V.X.; Tenório, J.C.G & Cavalcanti, L.H Myxomycetes from Alagoas state (Brazil) and notes on its distribution. Biotemas 27: Blackwell, M The Fungi: 1, 2, million species? American Journal of Botany 98: Cabral, T.S.; Marinho, P.; Goto, B.T. & Baseia, I.G Abrachium, a new genus in the Clathraceae, and Itajahya reassessed. Mycotaxon 119: Cáceres, M.E.S.; Vieira, T.S.; Jesus, L.S. & Lücking, R New and interesting lichens from the Caxiuanã National Forest in the Brazilian Amazon. Lichenologist 44: Cáceres, M.E.S.; Santos, V.M.; Góes, D.T.; Mota, D.A. & Aptroot, A Two new species of Malmidea from north-eastern Brazil. Lichenologist 45: Cáceres, M.E.S.; Aptroot, A. & Parnmen, S. 2014a. Remarkable diversity of the lichen family Graphidaceae in the Amazon rain forest of Rondônia, Brazil. Phytotaxa 189: Cáceres, M.E.S.; Ertz, D. & Aptroot, A. 2014b. New species and interesting records of Arthoniales from the Amazon, Rondônia, Brazil. Lichenologist 46: Cáceres, M.E.S.; Lima, E.L.; Aptroot, A. & Lücking, R. 2014c. Liquens brasileiros: novas descobertas evidenciam a riqueza no Norte e Nordeste do país. Boletim do Museu de Biologia Mello Leitão 35: Calaça, F.J.S.; Silva, N.C. & Xavier-Santos, S A checklist of coprophilous fungi and other fungi recorded on dung from Brazil. Mycotaxon 128: Carmo, L.T.; Monteiro, J.S.; Gusmão, L.F.P.; Sotão, H.M.P.; Gutierres, A.H. & Ruiz, R.F.C Anabahusakala, a new genus from the Brazilian Amazon rainforest. Mycotaxon 127: Rodriguésia 66(4):

111 1042 Maia, L.C. et al. Carvalho Jr., A.A. & Hennen, J.F New species and nomenclature in Prospodium (Uropyxidaceae, Pucciniales) and the new anamorphic genus Canasta in the Neotropics. Mycologia 102: Carvalho Jr., A.A. & Hennen, J.F The species of Puccinia on Piptocarpha and Vanillosmopsis in the Neotropics. Mycologia 104: Cavalcanti, L.H Biodiversidade e distribuição de mixomicetos em ambientes naturais e antropogênicos no Brasil: espécies ocorrentes nas Regiões Norte e Nordeste. In: Araujo, E.L. et al. (eds.). Biodiversidade, conservação e uso sustentável da flora do Brasil. Universidade Federal Rural de Pernambuco, Sociedade Botânica do Brasil, Recife. Pp Cavalcanti, L.H.; Ponte, M.P.P. & Mobin, M Myxomycetes, State of Piauí, Northeast Brazil. Check List 2: Cavalcanti, L.H.; Damasceno, G.; Bezerra, A.C.C. & Costa, A.A.A Mangrove myxomycetes: species occurring on Conocarpus erectus L. (Combretaceae). Sydowia 66: Cavalcanti, L.H.; Costa, A.A.A.; Barbosa, D.I.; Agra, L.A.N.N. & Bezerra, A.C.C Distribution and occurrence of Oligonema (Trichiales, Myxomycetes) in Brazil. Brazilian Journal of Botany 38: Coimbra, V.R.M. & Gibertoni, T.B First record of Trichopilus fasciculatus (Agaricales) from Brazil, with a key for the species of Entolomataceae from the Northern region. Mycoscience 56: Coimbra, V.R.M.; Gibertoni, T.B. & Wartchow, F Phaeocollybia nigripes (Agaricomycetes), a new species from Brazil. Mycotaxon 120: Coimbra, V.R.M.; Gibertoni, T.B. & Wartchow, F Megacollybia rimosa (Agaricales), a new species from Brazil. Mycoscience 54: Cortez, V.G.; Baseia, I.G. & Silveria, R.M.B Two noteworthy Phallus from southern Brazil. Mycoscience (Tokyo) 52: Costa, A.A.A.; Ferreira, I.N.; Bezerra, M.F.A. & Cavalcanti, L.H Mixobiota de Floresta Atlântica: novas referências de Physarales para o estado da Paraíba, Nordeste do Brasil. Revista Brasileira de Botânica 34: Costa, A.A.A.; Bezerra, A.C.C.; Lima, V.X. & Cavalcanti, L.H Diversity of Myxomycetes in an environmentally protected area of Atlantic Forest in northeastern Brazil. Acta Botanica Brasílica 28: Cruz, A.C.R.; Gusmão, L.F.P.; Castaneda Ruiz, R.F.; Stadler, M. & Minter, D.W Zelodactylaria, an interesting new genus from semi-arid northeast Brazil. Mycotaxon 119: Curvo, R.J.C Breve histórico da micologia e fungos referidos para Mato Grosso - Brasil. Revista Profiscientia 3: Damasceno, G.; Tenório, J.C.G.; Cavalcanti, L. H. & Andrade, L.H.C Stemonitaceae (Myxomycetes) in Brazilian mangroves. Sydowia 63: Drechsler-Santos, E.R.; Gibertoni, T. B.; Goes-Neto, A.; Cavalcanti, M.A.Q A re-evaluation of the lignocellulolytic Agaricomycetes from the Brazilian semi-arid region. Mycotaxon 108: Drechsler-Santos, E.R.; Cavalcanti, M.A.Q.; Loguercio- Leite, C. & Robledo, G. 2012a. On Neotropical Daedalea species: Daedalea ryvardenica sp. nov. Kurtziana 37: Drechsler-Santos, E.R.; Wartchow, F.; Coimbra, V.R.M.; Gibertoni, T.B. & Cavalcanti, M.A.Q. 2012b. Studies on lentinoid fungi (Lentinus and Panus) from the semi-arid region of Brazil. The Journal of the Torrey Botanical Society 139: Drechsler-Santos, E.R.; Wartchow, F.; Coimbra, V.R.M.; Gibertoni, T.B. & Cavalcanti, M.A.Q. 2012c. Studies on lentinoid fungi (and) from the semi-arid region of Brazil. The Journal of the Torrey Botanical Society 139: Drechsler-Santos, E.R.; Ryvarden, L.; Bezerra, J.L.; Gibertoni, T.B.; Salvador-Montoya, C.A. & Cavalcanti, M.A.Q New Records of Auriculariales, Hymenochaetales and Polyporales (Fungi, Agaricomycetes) for the Caatinga Biome. Check List 9: Feuerstein, S.C.; Cunha-Dias, I.P.R.; Aptroot, A.; Eliasaro, S. & Caceres, M.E.S Three new Diorygma (Graphidaceae) species from Brazil, with a revised world key. Lichenologist 46: Fidalgo, O Introdução à história da micologia brasileira. Rickia 3: Fiuza, P.O. & Gusmão, L.F.P. 2013a. Ingoldian fungi from the semi - arid Caatinga biome of Brazil. Mycosphere Online - Journal of Fungal Biology 4: Fiuza, P.O. & Gusmão, L.F.P. 2013b. Ingoldian fungi from semi-arid Caatinga biome of Brazil. The genus Campylospora. Mycosphere 4: Fiuza, P.O.; Gusmão, L.F.P.; Cruz, A.C.R. & Ruiz, R.F.C Conidial fungi from the semiarid Caatinga biome of Brazil: a new species of Pseudoacrodictys. Mycotaxon 127: Fiuza, P.O.; Ottoni-Boldrini, B.M.P.; Monteiro, J.S.; Catena, N.R.; Hamada, N. & Gusmão, L.F.P First records of Ingoldian fungi from the Brazilian Amazon. Brazilian Journal of Botany 38: 15 Forzza, R.C.; Baumgratz, J.F.A.; Bicudo, C.E.M.; Canhos, D.A.L.; Carvalho Jr., A.A.; Costa, A.F.; Costa, D.P.; Hopkins, M.; Leitman, P.M.; Lohmann, L.G.; Maia, L.C.; Martinelli, G.; Menezes, M.; Morim, M.P.; Nadruz-Coelho, M.A.; Peixoto, A.L.; Pirani, J.R.; Prado, J.; Queiroz, L.P.; Souza, V.C.; Stehmann, J.R.; Sylvestre, L.; Walter, B.M.T. & Zappi, D. (eds.) Catálogo de plantas e fungos Rodriguésia 66(4):

112 Diversity of Brazilian Fungi do Brasil. 2 vols. Andrea Jakobsson Estúdio / Jardim Botânico do Rio de Janeiro, Rio de Janeiro. 1699p. Gibertoni, T.B. & Drechsler-Santos, E.R Lignocellulolytic Agaricomycetes from the Brazilian Cerrado biome. Mycotaxon 111: Gibertoni, T.B.; Drechsler-Santos, E.R.; Baltazar, J.M.; Gomes-Silva, A.C.; Nogueira-Melo, G.S.; Ryvarden, L. & Cavalcanti, M.A.Q The genus Trichaptum (Agaricomycetes, Basidiomycota) in Brazil. Nova Hedwigia 93: Gibertoni, T.B., Martins-Júnior, A., Ryvarden, L. & Sotão, H Oxyporus mollis sp. nov. (Agaricomycetes) from the Eastern Brazilian Amazonia. Nova Hedwigia 94: Gibertoni, T.B. & Ryvarden, L Studies in Neotropical polypores 36. A note on the genus Henningsia. Synopsis Fungorum 32: Goes Neto, A. & Cavalcanti, L.H Myxomycetes of the State of Bahia: historical review and current situation. Mycotaxon 82: Gomes-Silva, A.C.; Ryvarden, L. & Gibertoni, T.B Two new species of Phellinus s.l. from the Brazilian Amazonia. Phytotaxa 67: Gomes-Silva, A.C.; Ryvarden, L. & Gibertoni, T.B Inonotus amazonicus sp. nov., I. calcitratus comb. nov. and notes on Phylloporia (Hymenochaetaceae, Agaricomycetes) from the Brazilian Amazonia. Mycoscience 54: Gomes-Silva, A.C.; Medeiros, P.S.; Soares, A.M.S.; Sotão, H.M.P.; Ryvarden, L. & Gibertoni, T.B Two new species of Rigidoporus (Agaricomycetes) from Brazil and new records from the Brazilian Amazonia. Phytotaxa 156: Goto, B.T.; Silva, G.A.; Yano-Melo, A.M. & Maia, L.C Checklist of the arbuscular mycorrhizal fungi (Glomeromycota) in the Brazilian semiarid. Mycotaxon 113: Goto, B.T.; Silva, G.A.; Maia, L.C.; Souza, R.G.; Coyne, D.; Tchabi, A.; Lawouin, L.; Hountondji, F. & Oehl, F Racocetra tropicana, a new species in the Glomeromycetes from tropical areas. Nova Hedwigia 92: Goto, B.T.; Silva, G.A.; Assis, D.M.A.; Silva, D.K.A.; Souza, R.G.; Ferreira, A.C.A.; Jobim, K.; Mello, C.M.A.; Vieira, H.E.E.; Maia, L.C. & Oehl, F Intraornatosporaceae (Gigasporales), a new family with two new genera and two new species. Mycotaxon 119: Goto, B.T.; Araújo, A.F.; Soares, A.C.F.; Ferreira, A.C.A.; Maia, L.C.; Sousa, C.S. & Silva, G.A Septoglomus titan, a new fungus in the Glomeraceae (Glomeromycetes) from Bahia, Brazil. Mycotaxon 124: Gusmão, L.F.P. & Maia, L.C (eds.). Diversidade e caracterização dos fungos do semi-árido brasileiro. Vol. 2. Instituto do Milênio do Semi-árido. MCT/ Associação Plantas do Nordeste, Recife. 219p Hennen, J.F.; Figueiredo, M.B.; Carvalho Jr., A.A. & Hennen, P.G Catalogue of the species of plant rust fungi (Uredinales) of Brazil. Jardim Botânico do Rio de Janeiro, Rio de Janeiro. 490p. Hibbett, D.S.; Binder, M.; Bischoff, J.F.; Blackwell, M.; Cannon, P.F.; Eriksson, O.E.; Huhndorf, S.; James, T.; Kirk, P.M.; Lücking, R.; Lumbsch, H.T.; Lutzoni, F.; Matheny, P.B.; McLaughlin, D.J.; Powell, M.J.; Redhead, S.; Schoch, C.L.; Spatafora, J.W.; Stalpers, J.A.; Vilgalys, R.; Aime, M.C.; Aptroot, A.; Bauer, R.; Begerow, D.; Benny, G.L.; Castlebury, L.A.; Crous, P.W.; Daí, Y.C.; Gams, W.; Geiser, D.M.; Griffith, G.W.; Gueidan, C.; Hawksworth, D.L.; Hestmark, G.; Hosaka, K.; Humber, R.A.; Hyde, K.D.; Ironside, J.E.; Köljalg, U.; Kurtzman, C.P.; Larsson, K.H.; Lichtwardt, R.; Longcore, J.; Miadlikowska, J.; Miller, A.; Moncalvo, J.M.; Mozley-Standridge, S.; Oberwinkler, F.; Parmasto, E.; Reeb, V.; Rogers, J.D.; Roux, C.; Ryvarden, L.; Sampaio, J.P.; Schüssler, A.; Sugiyama, J.; Thorn, R.G.; Tibell, L.; Untereiner, W.A.; Walker, C.; Wang, Z.; Weir, A.; Weiss, M.; White, M.M.; Winka, K.; Yao, Y.J. & Zhang, N A higher-level phylogenetic classification of the Fungi. Mycological Research 111: Hochgesand, E. & Gottsberger, G Myxomycetes from State of São Paulo. Boletim do Instituto de Botânica 10: Hoffmann, K.; Voigt, K. & Kirk, P.M Mortierellomycotina subphyl. nov., based on multi-genes genealogies. Mycotaxon 115: Humber, R.A Entomophthoromycota: a new phylum and reclassification for entomophthoroid fungi. Mycotaxon 120: Izabel, T.S.S.; Santos, D.; Almeida, D.A.C. & Gusmão, L.F.P Fungos conidiais do Bioma Caatinga. II. Novos registros para o Continente americano, Neotrópico e América do Sul e Brasil. Rodriguesia 62: Izabel, T.S.S.; Cruz, A.C.R. & Gusmão, L.F.P Conidial fungi from the semi-arid Caatinga biome of Brazil. Ellisembiopsis gen. nov., new variety of Sporidesmiella and some notes of Sporidesmium complex. Mycosphere Online - Journal of Fungal Biology 4: Jesus, A.L.; Marano, A.V.; Schoenlein-Crusius, I.H. & Pires-Zottarelli, C.L.A Diversidade de organismos zoospóricos do córrego Pirarungaua, Parque Estadual das Fontes do Ipiranga, São Paulo, Brasil: novas citações. Hoehnea 40: Kirk, P.; Cannon, P.F.; Minter, D.W. & Stalpers, J.A a ed. Dictionary of Fungi. CABI Publishing, Wallinford. 771p. Lima, E.L.; Maia, L.C.; Aptroot, A. & Cáceres, M.E.S. 2013a. New lichen species from Vale do Catimbau, Pernambuco, Brazil. The Bryologist 116: Lima, E.L.; Mendonça, C.O.; Maia, L.C.; Aptroot, A. & Cáceres, M.E.S. 2013b. Two new species of Rodriguésia 66(4):

113 1044 Maia, L.C. et al. Pyrenula with a red or orange thallus from Vale do Catimbau National Park, Pernambuco, Brazil. Lichenologist 45: Maia, L.C.; Melo, A.M.Y. & Cavalcanti, M.A Diversidade de fungos no estado de Pernambuco. In: Tabarelli, M. & Silva, J.M.C. da (orgs.). Diagnóstico da Biodiversidade de Pernambuco. Vol. 1. Editora Massangana, Recife. Pp Maia, L.C.; Drechsler-Santos E.R. & Cáceres, M Representatividade dos fungos nos herbários brasileiros. In: Maia, L.C.; Malosso, E. & Yano-Melo, A.M. (orgs.). Micologia: avanços no conhecimento. Universitária da UFPE, Recife. Pp Maia, L.C. & Carvalho Jr., A.A Fungos do Brasil. In: Forzza, R.C. et al. (orgs.). Catálogo de plantas e fungos do Brasil. Vol.1. Andrea Jakobsson Estúdio /Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro. Pp Maia, L.C.; Silva, G.A.; Yano-Melo, A.M. & Goto, B.T Fungos micorrízicos arbusculares no bioma Caatinga. In: Siqueira, J.O.; de Souza, F.A.; Cardoso, E.J.B.N. & Tsai, S.M. (orgs.). Micorrizas: 30 anos de pesquisas no Brasil. Vol. 1. Editora UFLA, Lavras. Pp Maimoni-Rodella, R.C Biodiversidade e distribuição de mixomicetos em ambientes naturais e antropogênicos no Brasil: espécies ocorrentes nas Regiões Sudeste e Centro-Oeste. In: Araujo, E.L. et al. (eds.). Biodiversidade, conservação e uso sustentável da flora do Brasil. Universidade Federal Rural de Pernambuco, Sociedade Botânica do Brasil, Recife. Pp Marinho, F.; Silva, G.A.; Ferreira, A.C.A.; Veras, J.S.N.; Souza, N.M.F.; Goto, B.T.; Maia, L.C. & Oehl, F Bulbospora minima, a new genus and a new species in the Glomeromycetes from semi-arid Northeast Brazil. Sydowia 66: Meijer, A.A.R Macrofungos Notáveis das Florestas de Pinheiro-do-Paraná. Embrapa, Colombo. 431p. Mello, C.M.A.; Silva, G.A.; Vieira, H.E.E.; Silva, I.R.; Maia, L.C. & Oehl, F Fuscutata aurea, a new species in the Glomeromycetes from cassava and maize fields in the Atlantic rainforest zone of Northeastern Brazil. Nova Hedwigia 95: Mello, C.M.A.; Silva, G.A.; Assis, D.M.A.; Pontes, J.S.; Ferreira, A.C.A.; Leão, M.P.C.; Vieira, H.E.E.; Maia, L.C. & Oehl, F Paraglomus pernambucanum sp. nov. and Paraglomus bolivianum comb. nov., and biogeographic distribution of Paraglomus and Pacispora. Journal of Applied Botany and Food Quality 86: Melo, R.F.R.; Miller, A.N.; Santiago, A.L.C. & Maia, L.C The genera Ascobolus and Saccobolus (Ascobolaceae, Pezizales) in Brazil. Mycosphere Online - Journal of Fungal Biology 5: Mendes, M.A.S.; Ferreira, M.A.S.V.; Dianese, J.C.; Santos, C.E.N.; Urben, A.F. & Castro, C Fungos em Plantas no Brasil. Embrapa-SPI/Embrapa- Cenargen, Brasília, DF. 569p. Mendes, M.A.S.; Urbem, A.F. & Dianese, J.C Fungos em plantas no Brasil. Vol. 1. 2ª ed. Embrapa, Brasília. 978p. Menezes, A.A.; Lima, E.L.; Leite, A.B.X.; Maia, L.C.; Aptroot, A. & Cáceres, M.E.S New species of Arthoniales from NE Brazil. Lichenologist 45: Menolli Jr., N.; Breternitz, B.S. & Capelari, M The genus Pleurotus in Brazil: a molecular and taxonomic overview. Mycoscience 55: Milanez, A.I.; Pires-Zotarelli, C.L.A. & Gomes, A.L Brazilian zoosporic fungi. Seção de Micologia/ IBt, São Paulo. 112p. Monteiro, J. S. & Gusmão, L.F.P An emendation of Fusticeps and two new species from the Brazilian Amazon Forest. Mycotaxon 123: Monteiro, J.S.; Gusmão L.F.P. & Ruiz, R.F.C. 2014a. Helicodochium, a new microfungus from submerged wood in Brazil. Mycotaxon 127: 5-9. Monteiro, J.S.; Gusmão, L.F.P. & Ruiz, R.F.C. 2014b. Two new microfungi from Brazilian Amazon Forest: Atrogeniculata submersa and Nigrolentilocus amazonicus. Mycotaxon 127: Nascimento, C.A.; Gomes, E.P.C.; Souza, J.I. & Pires- Zottarelli, C.L.A Zoosporic true fungi and heterotrophic straminipiles assemblages from soil of Brazilian Cerrado areas. 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114 Diversity of Brazilian Fungi Sá, M.C.A.; Pinheiro, F.G.B.; Silva, N.A.; Maia, L.C. & Wartchow, F Craterellus niger (Cantharellaceae, Cantharellales, Basidiomycota): a new species from Pernambuco, Brasil. Nova Hedwigia 99: Salazar-Yepes, M. & Carvalho Jr., A.A. 2010a. Novos registros de ferrugens (fungi, Uredinales) para o Brasil, coletados no Parque Nacional do Itatiaia. Acta Botanica Brasilica 24: Salazar-Yepes, M. & Carvalho Jr., A.A. 2010b. Ferrugens: diversidade de uredinales do Parque Nacional do Itatiaia, Brasil. Vol. 1. Technical Books, Rio de Janeiro. 201p. Salazar-Yepes, M. & Carvalho Jr., A.A Caetea, a new genus of Pucciniales on Piptadenia (Fabaceae) from Brazil. Mycologia 104: Salazar-Yepes, M. & Carvalho Jr., A.A Two new rust species on Fabaceae from Brazil. Mycotaxon 128: Santiago, A.L.C.; Benny, G. & Maia, L.C. 2011a. Syncephalis aggregata-a new species from the semiarid region of Brazil. Mycologia 103: Santiago, A.L.C.M. de A.; Trufem, S.F.B.; Malosso, E.; Santos, P.J.F. & Cavalcanti, M.A.Q. 2011b. Zygomycetes from herbivore dung in the ecological reserve of Dois Irmãos, Northeast Brazil. Brazilian Journal of Microbiology 42: Santiago, A.L.C.M. de A.; Santos, P J.P. & Maia, L.C Mucorales from the semiarid of Pernambuco, Brazil. Brazilian Journal of Microbiology 44: Santiago, A.L.C.; Hoffmann, K.; Lima, D.X.; Oliveira, R.J.; Vieira, H.E.E.; Malosso, E.; Maia, L.C. & Silva, G.A A new species of Lichtheimia (Mucoromycotina, Mucorales) isolated from Brazilian soil. Mycological Progress 13: Schüssler, A.; Schwarzott, D. & Walker, C A new fungal phylum, Glomeromycota: phylogeny and evolution. Mycological Research 105: Silva, M. & Minter, D.W Fungi from Brazil Recorded by Batista and Co-workers. Mycological Papers 169: Silva A.C.G. & Gibertoni, T.B Revisão do Herbário URM: novas ocorrências de Aphyllophorales para a Amazônia brasileira. Revista Brasileira de Botânica 32: Silva, C.F. & Cavalcanti, L.H Myxobiota of the Brazilian Atlantic Forest: species on oil palm tree (Elaeis guineensis, Arecaceae). Rodriguésia 61: Silva, N.A. & Cavalcanti, L.H Myxomycetes ocorrentes em áreas de Caatinga e Brejo de Altitude no sertão de Pernambuco, Brasil. Acta Botanica Brasilica 26: Silva, G.A.; Maia, L.C. & Oehl, F Phylogenetic systematics of the Gigasporales. Mycotaxon 122: Silva, C.R.; Castaneda-Ruiz, R.F. & Gusmão, L.F.P Anacraspedodidymum, a new genus from submerged wood in Brazil. Mycotaxon 128: Soares, A.M.S.; Sotão, H.M.P.; Medeiros, P.S. & Gibertoni, T Riqueza de fungos poliporoides (Agaricomycetes, Basidiomycota) em uma floresta ombrófila densa no Amapá, Amazônia brasileira. Boletim do Museu de Biologia Mello Leitao 35: Souza, J.I.; Pires-Zottarelli, C.L.A.; Santos, J.F.; Costa, J.P. & Harakava, R Isomucor (Mucoromycotina): a new genus from a Cerrado reserve in state of São Paulo, Brazil. Mycologia 104: Souza, J.I.; Marano, A.V.; Pires-Zottarelli, C.L.A.; Chambergo, F.S. & Harakava, R A new species of Backusella (Mucorales) from a Cerrado reserve in Southeast Brazil. Mycological Progress 13: Trierveiler-Pereira, L.; Calonge, F.D. & Baseia, I.G New distributional data on Geastrum (Geastraceae, Basidiomycota) from Brazil. Acta Botanica Brasilica 25: Viégas, A. P Verticillium lecanii (Zimm.) n. comb., causador do halo branco do Coccus viridis (Green). Revista do Instituto de Café 25: Viégas, A.P Môfo dos afídeos e aleirodídeos. Revista de Agricultura 15: Viégas, A.P Alguns fungos do cerrado. Bragantia 3: Viégas, A.P Alguns fungos do Brasil - Ascomycetos. Bragantia 4: Viégas, A.P Alguns fungos do Brasil - Cercospora. Boletim da Sociedade Brasileira de Agronomia 8: Viégas, A.P Alguns fungos do Brasil. XIII - Hifomicetos. Bragantia 6: Viégas, A.P Índice de fungos da América do Sul. Campinas, Instituto Agronômico de Campinas. 921p. Wartchow, F.; Maia, L.C. & Cavalcanti, M.A. 2013a. Studies on Amanita (Agaricomycetidae, Amanitaceae) in Brazil: two yellow gemmatoid taxa. Nova Hedwigia 96: Wartchow, F.; Maia, L.C. & Cavalcanti, M.A.Q. 2013b. Taxonomic studies of Amanita muscaria (L.) Lam (Amanitaceae, Agaricomycetes) and its infraspecific taxa in Brazil. Acta Botanica Brasilica 27: Artigo recebido em 04/06/2015. Aceito para publicação em 04/09/2015. Rodriguésia 66(4):

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116 Rodriguésia 66(4): DOI: / Update of the Brazilian floristic list of Algae and Cyanobacteria This paper was compiled by Mariângela Menezes 1, Carlos E. M. Bicudo and Carlos W. N. Moura With contributions by: Aigara M. Alves; Alana A. Santos; Alexandre de G. Pedrini; Andréa Araújo; Andrea Tucci; Aurelio Fajar; Camila Malone; Cecília H. Kano; Célia L. Sant Anna; Ciro Z. Branco; Clarisse Odebrecht; Cleto K. Peres; Emanuel B. Neuhaus; Enide Eskinazi-Leça; Eveline Aquino; Fabio Nauer; Gabriel N. Santos; Gilberto M. Amado Filho; Goia M. Lyra; Gyslaine C.P. Borges; Iara O. Costa; Ina de S. Nogueira; Ivania B. Oliveira; Joel. C.de Paula; José M. de C. Nunes; Jucicleide C. Lima; Kleber R.S. Santos; Leandro C. Ferreira;Lísia M.S. Gestinari; Luciana S. Cardoso; Marcia A.O. Figueiredo; Marcos H. Silva; Maria B.B.B. Barreto; Maria C.O. Henriques; Maria da G.G.S.Cunha; Maria E. Bandeira-Pedrosa; Maria F. Oliveira-Carvalho; Maria T.M. Széchy; Maria T.P. Azevedo; Mariana C. de Oliveira; Mariê M. Cabezudo; Marilene F. Santiago; Marli Bergesh; Mutuê T. Fujii; Norma C. Bueno; Orlando Necchi Jr.; Priscila B. Jesus; Ricardo G. Bahia; Samir Khader; Sandra M. Alves-da-Silva; Silvia M.P.B. Guimarães; Sonia M.B. Pereira; Taiara A. Caires; Thamis Meurer; Valéria Cassano; Vera R. Werner; Watson A. da Gama Jr. & Weliton J. da Silva. Abstract An updated synthesis of cyanobacteria and algae information is presented for Brazil aiming to refine the data gathered to date and evaluate the progress of the biodiversity knowledge about these organisms in the country since the publication of the Catálogo de Plantas e Fungos do Brasil. The results of 2015 showed an increase of 1,250 species (35.7%) when compared to 2010, reaching a total of 4,747 species. The most diverse classes in species number were the Bacillariophyceae, Conjugatophyceae, Florideophyceae, Cyanophyceae, Dinophyceae and Euglenophyceae. Bacillariophyceae and Cyanophyceae had the highest increase in species number in the five-year interval. The Southeast and South regions were the most diverse, however, the Northeast, with the states of Piauí and Sergipe, and the Central-west region, with Mato Grosso, Goiás and Distrito Federal, also stood out in the national algal biodiversity scenario. Despite the shortage of taxonomists and limited infrastructure, the results showed a significant improvement in the knowledge regarding the diversity of cyanobacteria and algae in the country during the study period, starting to even out regional geographical differences caused by subsampling. Key words: Biodiversity, phycology. Resumo Apresenta-se uma síntese atualizada de informações sobre algas no Brasil objetivando refinar os dados reunidos até o presente, bem como avaliar os avanços sobre o conhecimento da diversidade de algas no país desde a publicação do Catálogo de Plantas e Fungos do Brasil. Os resultados de 2015 mostraram um acréscimo de espécies (35.7%) a um total de em relação a As classes mais diversas em número de espécies foram Bacillariophyceae, Conjugatophyceae, Florideophyceae, Cyanophyceae, Dinophyceae e Euglenophyceae. Bacillariophyceae e Cyanophyceae tiveram o maior acréscimo de espécies no intervalo de cinco anos. A região Sudeste e Sul foram as mais diversas, porém, as regiões Nordeste com os estados do Piauí e Sergipe e Centro-Oeste com os estados de Mato Grosso, Goiás e Distrito Federal destacaram- -se no cenário da biodiversidade nacional. Apesar da escassez de taxonomistas e da infraestrutura limitada, os resultados obtidos evidenciaram um avanço significativo no conhecimento da diversidade de algas no país nesse período de cinco anos, iniciando uma mudança quanto as diferenças geográficas regionais. Palavras-chave: Biodiversidade, ficologia. For the complete list of authors, institutions and taxonomic groups see supplementary material < DOI: / ). 1 Author for correspondence: menezes.mariangela@gmail.com

117 1048 Menezes, M. et al. Introduction The publication of the Algae and Cyanobacteria (Bicudo & Menezes 2010) in the Catálogo de Plantas e Fungos do Brasil (Forzza et al. 2010) represents a milestone for the phycological knowledge in the country and worldwide. Until then, there was no single compilation presenting detailed information of these organisms diversity in different parts of Brazil. The knowledge was dispersed in regional or local lists (e.g. Cordeiro- Marino 1978; De-Lamonica-Freire 1989a, 1989b; Senna et al. 1998; Menezes & Dias 2001; Oliveira et al. 2002; Torgan et al. 2003) and/or restricted to particular algal groups (p.ex. Moreira-Filho et al. 1985; Torgan et al. 1999, 2001; Alves-da-Silva & Hahn 2001; Guimarães 2006; Tremarin et al. 2009; Procopiack et al. 2006). Since the publication of the Catálogo de Plantas e Fungos do Brasil, several new local and regional lists focusing on specific algal groups and general catalogues were published. Diatoms of central-western Brazil (da Silva et al. 2011); Chlorophyceae of Mato Grosso (Freitas & Loverde- Oliveira 2013); cyanobacteria and continental Algae of Pará (Costa et al. 2014); deep sea macroalgae associated to rodolith beds in coastal Espírito Santo (Amado Filho et al. 2010); bentonic algae from the Laje Santos, São Paulo (Jorge et al. 2012); macroalgae in the islands off the coast of Paraná (Pellizari et al. 2014); bentonic algae from Sergipe s coast (Pereira et al. 2014); and the revised and updated checklist of macroalgae from the Abrolhos Archipelago and Sebastião Gomes reef, state of Bahia (Torrano-Silva & Oliveira 2014a). Numerous articles regarding new records for Brazil and descriptions of new species have also been published. Increased collecting effort, especially in deep waters, and molecular biology studies have generated valuable knowledge. DNA barcoding tecniques helped to perform more precise identifications and also led to the correction of former determinations that incurred in erroneous citations of taxa that, in fact, do not occur along the Brazilian coast. Some examples are publications by Sutherland et al. (2011), Bahia et al. (2011), Cassano et al. (2012), Carvalho et al. (2012), Alves et al. (2012), Bahia et al. (2013), Jesus et al. (2013), Pellizzari et al. (2013), Rocha-Jorge et al. (2013), Bahia et al. (2014a, 2014b), Henriques et al. (2014), Moura et al. (2014), Nauer et al. (2014a, b), Nunes et al. (2014), Torrano-Silva et al. (2014b), Jesus et al. (2015) and Lyra et al. (2015). New occurrences and, more noticeably, the description of new taxa of microalgae were also a result of the increase in collection efforts, particularly the work regarding diatoms and cyanobacteria carried out by Wetzel et al. (2012a, b), Gama Jr. et al. (2012), Burliga et al. (2013), Caires et al. (2013), Santos et al. (2013) and Tremarin et al. (2013). The present paper aims to present a synthesis of the updated list of Brazilian Algae based in the new records, literature and herbarium data added between 2010 and The objective was to refine and update the data initially gathered, adjusting what was published for Algae in the Brazilian Catalogue (Bicudo & Menezes 2010) and to examine the information compiled in the Brazilian List over the past five years. Methods Methodology followed that published by Forzza et al. (2010). Taking the Algae list published by Bicudo & Menezes (2010) as a starting point, new taxa and occurrences for Brazil were included in the online system together with voucher information and/or literature reference stating the presence of a given taxon in Brazil, its geographic distribution, environment and life-form. The taxonomic status of all names was checked and nomenclatural updates followed Guiry & Guiry (2015) and other recent relevant articles, such as Krienitz & Bock (2012) and Gómez (2013). Changes in class circumscription meant that previously recognized classes Prymnesiophyceae and Rhodophyceae were not featured in the 2015 version of the Brazilian List. The first was included in Coccolithophyceae, while the latter was broken down into four different classes: Bangiophyceae, Florideophyceae, Porphyriophyceae and Stylonematophyceae. Genera Verdigellas D.L.Ballan. & J.N.Norris and Palmophyllum Kütz. (formerly included in Ulvophyceae), Micromonas Manton & Parke, Pseudoscourfieldia Manton and Pyramimonas Schmarda were placed in the Prasinophyceae. Partial resolution of the phylogenetic relationships of these genera was provided by Marin & Melkonian (2009) and Fučíková et al. (2014) leading to the adoption of the prasinophytes group concept, coined by Leliaert et al. (2012). The complete dataset used for the present analyses can be found at the Brazilian List of Algae (see supplementary material < Rodriguésia 66(4):

118 Brazilian Algae and Cyanobacteria org/ /m9.figshare > - DOI: / ). This new dataset includes, as well as the Brazilian states and regions of occurrence for the taxa, the distribution by hydrographic region for the continental Algae in order to provide data for biogeographic studies and conservation strategies. Under the definition of the Brazilian National Council for Water Resources (Conselho Nacional de Recursos Hídricos - CNRH) an hydrographic region represents the territorial space comprised by a drainage basin or group of basins or sub-basins close to each other and with homogeneous or similar natural, social and economic attributes, established through Brazilian law (Resolution 32 of the CNRH, published 15/10/03). Each hydrographic region constitutes an administrative region and also a main unit for planning and management of water resources, falling under the responsibility of the CNRH. Furthermore, the integrated management of combined hydrographic basins, such as the implantation of a network of conservation units, is an example of an adequate model to select suitable conservation regions, favouring connectivity of the habitats within and between the hydrographic basins (Hero & Hidway 2006). Therefore, according to the CNRH (see above), Brasil is currently divided into 12 hydrographic regions: 1) Amazonian (states of Acre, Amazonas, Rondônia and Roraima, Pará, Mato Grosso and Amapá); 2) East Atlantic (Sergipe, Bahia, Minas Gerais and of Espírito Santo); 3) Northeast Occidental (Pará and Maranhão); 4) Northeast Oriental (Piauí, Ceará, Paraíba, Rio Grande do Norte, Alagoas and Pernambuco); 5) Southeast Atlantic (Minas Gerais, Espírito Santo, Rio de Janeiro, São Paulo (25% of Southeastern Brazilian region) and Paraná); 6) South Atlantic (Paraná, Santa Catarina and Rio Grande do Sul); 7) Paraguay (Mato Grosso do Sul and Mato Grosso); 8) Paraná (São Paulo, Paraná, Mato Grosso do Sul, Minas Gerais, Goiás, Santa Catarina and Distrito Federal); 9) Parnaíba (Piauí, Maranhão and Ceará); 10) São Francisco (Minas Gerais, Distrito Federal, Goiás, Bahia, Pernambuco, Alagoas and Sergipe); 11) Tocantins- Araguaia (Goiás, Tocantins, Pará, Maranhão, Mato Grosso and Distrito Federal); and 12) Uruguay (Rio Grande do Sul and Santa Catarina). Results The analysis of the 2015 Brazilian List of Algae (see supplementary material < dx.doi.org/ /m9.figshare > - DOI: / ) shows the algae and cyanobacteria diversity divided in 29 classes, 1,018 genera, 4,747 species, 12 subspecies and 1,424 varieties. The Incertae sedis category was added in 2014 to include taxa referred for Brazil in 19th century publications for which the systematic positioning remains unsure (39 species). When compared with the data from 2010 (Fig. 1) an increase of 1,711 taxa, 12 classes, 189 genera, 1,250 species, four subspecies and 457 varieties. Of these, 33 are new taxa described for Brazil by a total of 48 Brazilian researchers in the following groups: Cyanophyceae (13), Bacillariophyceae (12), Florideophyceae (11), Conjugatophyceae (four), Trebouxiophyceae (three), Florideophyceae (nine), Bangiophyceae (three), Ulvophyceae (seven) (Tab. 1). The ten most diverse classes were: Bacillariophyceae with 1,247 species, followed by the Conjugatophyceae (610), Florideophyceae (489), Cyanophyceae (462), Dinophyceae (420), Euglenophyceae (367), Chlorophyceae (352), Ulvophyceae (217), Phaeophyceae (103) and Coccolithophyceae (91). There were no changes regarding the diversity of classes Chrysophyceae, Cryptophyceae, Dictyocophyceae, Raphidophyceae, Synurophyceae and Xanthophyceae. On the other hand, a drop in the number of taxa in the Figure 1 Number of algae and cyanobacteria for Brazil distributed by taxonomic levels in 2010 and Table 1 Distribution of the number of new species of algae and cyanobacteria described by their publishing Brazilian authors between 2010 and Number of new species Number of authors Rodriguésia 66(4):

119 1050 Menezes, M. et al. Table 2 Number of algae and cyanobacteria genera and species recorded in 2010 and 2015, distributed by class. (-) - Not included in 2010 or 2015; (*) - Included in Coccolithophyceae in 2014; (**) - subdivided in 2015 in Bangiophyceae, Compsopogonophyceae, Florideophyceae, Porphyriophyceae and Stylonematophyceae. GEN: genera, SP: species. Classes Bacillariophyceae Charophyceae Chlorophyceae Cyanophyceae Dinophyceae Euglenophyceae Phaeophyceae Prasinophyceae Ulvophyceae Chrysophyceae Cryptophyceae Dictyocophyceae Raphidophyceae Synurophyceae Xanthophyceae Bangiophyceae Categorias Ano GEN SP GEN 35 3 SP GEN SP GEN SP GEN SP GEN SP GEN SP GEN 8 5 SP GEN SP GEN SP GEN 9 9 SP GEN 2 2 SP 4 4 GEN 6 6 SP 9 9 GEN 3 3 SP GEN SP GEN - 3 SP - 7 Rodriguésia 66(4):

120 Brazilian Algae and Cyanobacteria 1051 Classes Chlorodendrophyceae Coccolithophyceae Coleochaetophyceae Compsopogonophyceae Conjugatophyceae Florideophyceae Klebsormidiophyceae Mamiellophyceae Nephroselmidophyceae Pedinophyceae Porphyridiophyceae Stylonematophyceae Prymnesiophyceae* Rhodophyceae* Categorias Ano GEN - 2 SP - 2 GEN - 49 SP - 91 GEN - 1 SP - 3 GEN - 4 SP - 5 GEN - 44 SP GEN SP GEN - 1 SP - 5 GEN - 1 SP - 1 GEN - 1 SP - 3 GEN - 2 SP - 2 GEN - 1 SP - 2 GEN - 3 SP - 4 GEN 46 - SP 93 - GEN SP Charophyceae followed the exclusion of the Desmidiaceae, nowadays considered as part of the Conjugatophyceae. The Rodophyceae were increased by 36 genera and 111 species. The classes that saw the largest increases in species numbers in 2015 were the Bacillariophyceae and the Cyanophyceae (Tab. 2). In terms of epicontinental or marine environment, eleven classes are exclusively epicontinental (Charophyceae, Chlorodendrophyceae, Chlorophyceae, Coleochaetophyceae, Conjugatophyceae, Klebsormidiophyceae, Mamiellophyceae, Porphyridiophyceae, Synurophyceae, Rodriguésia 66(4):

121 1052 Menezes, M. et al. Trebouxiophyceae and Xanthophyceae), six are exclusively marine (Coccolithophyceae, Bangiophyceae, Dictyochophyceae, Prasinophyceae, Phaeophyceae and Stylonematophyceae), while the remaining 13 classes occur in both environments (Tab. 3). Algae and cyanobacteria distribution by Brazilian geopolitical regions is shown in Figure 2. When compared to the 2010 data, the new data show an increase in the number of taxa in all regions with Southeast Brazil as the most diverse, followed by South, Northeast, Centralwest and Northern. When we analyse the state records, only Amapá, Acre, Roraima, Rondônia and Tocantins did not show expressive increase in species numbers from 2010 to 2015 (Tab. 4). The states with the higher increase in number of taxon records were Mato Grosso and Piauí, where the records were respectively 81% and 79% higher, followed by Sergipe, with 54%, Goiás 50% and Distrito Federal with 40%. Despite the increases, species distribution between the states and the geopolitical regions continued to be heterogeneous (Fig. 3). Taking into account only the macroalgae and marine cyanobacteria (Fig. 4), number of species per class continued to be higher in the southeast and northeast regions. Excepting Cyanophycae and macroscopic Prasinophyceae, 762 species of red, brown and green algae were recorded in 2015, amounting to 12.6% more than found in The states where the marine macroalgae have shown higher number of species were the same found by Bicudo & Menezes (2010), namely Espírito Santo, Rio Janeiro, Bahia, São Paulo, Pernambuco and Rio Grande do Norte, and the ones where the species number increased more Figure 2 Number of epicontinental algae and cyanobacteria taxa distributed by Brazilian geopolitical region in 2010 and markedly were Rio Grande do Norte (91), Bahia (87), Espírito Santo (79), Sergipe (75), São Paulo (71) and Pernambuco (63) (Tab. 5). Regarding marine cyanobacteria, 2015 saw 73 species while in 2010 there were only 17 recorded for Brazil, and the state with higher number of species was São Paulo (63), followed by Rio de Janeiro (26), Bahia (23) and Rio Grande do Sul (16). Other states such as Pará, Maranhão, Ceará, Rio Grande do Norte, Espírito Santo, Paraná and Santa Catarina have. There were no changes in the number of species or the distribution of Verdigellas and Palmophyllum (Prasinophyceae) between 2010 and 2015, with the species distribution being restricted to deep environments off the coast of Bahia, Espírito Santo and Rio de Janeiro (Tab. 5). The marine microalgae (Tab. 3) had an increase of 251 species (1,175 in 2015 vs. 924 in 2010). The states with the largest growth in the number of records were Paraná (37%), Pernambuco (32%), Rio de Janeiro (29%), Pará (27%), Rio Grande do Sul (25%), Espírito Santo (21%) and Bahia (21%). The remaining states had inexpressive alterations on the number of taxa, growing up to 16%, e.g. Bahia, Alagoas, Sergipe, Piauí and Amazonas had records equal or lower than five taxa. South and Northeast regions continued to be the most diverse, while, at state level, São Paulo (549 species), Rio Grande do Sul (529 species), Rio de Janeiro (491 species), Paraná (364 species), Pernambuco (348 species) and Bahia (303) are the leading states. The Bacillariophyceae and Dinophyceae continued to be the microalgae classes with the highest number of taxa (Tab. 6). Epicontinental algae (Tab. 3) have reached 2,808 species records in 2015, an additional 918 species to the 2010 figure of 1,890. Central-west, South, and Southeast regions have had the highest species number increases since 2010, followed by Northeast and North. The states of Bahia (85%), Minas Gerais (42%), Goiás (38.3%), Amazonas (28.2%), Distrito Federal (27%), Mato Grosso (25%) and Paraná (24%) had the highest species number increases. The most diverse states in terms of species numbers were Rio de Janeiro, São Paulo, Paraná, Rio Grande do Sul, Amazonas, Mato Grosso, Goiás, Distrito Federal and Bahia. The most diverse group continues to be the Bacillariophyceae followed by Conjugatophyceae (formerly included in the Charophyceae), Euglenophyceae and Chlorophyceae (Tab. 6). Rodriguésia 66(4):

122 Brazilian Algae and Cyanobacteria Cyanobacteria species number added up to 389 in 2015 from a total of 294 in 2010, and the regions with highest increase in records were the Southeast and South (83% each), and the Central-western (53.5%). At state level, São Paulo (48.2%), Rio de Janeiro (42%), Rio Grande do Sul (42%), Amazonas (37.2%) and Distrito Federal (31.4%) have shown the largest increase in specie records (Tab. 3, Tab. 6). São Paulo, Rio de Janeiro and Rio Grande do Sul feature as the most diverse states in terms of number of taxa (Tab. 6). Amongst the 1,018 genera currently recognized in 2015, 41 (4%) concentrate 1,852 (39%) of the species listed (Tab. 7). The four genera 1053 with largest number of taxa were Cosmarium Corda ex Ralfs (Conjugatophyceae) with 156 species, Pinnularia Ehrenb. (Bacillariophyceae) with 125, Trachelomonas Ehrenb. (Euglenophyceae) with 107 and Staurastrum Meyen ex Ralfs (Conjugatophyceae), with 93 species. Regarding the occurrence of algae and cyanobacteria by hydrographic region (Fig. 5), the most representative were the Southeast Atlantic and Paraná, followed by the South Atlantic and the Amazonian Hidrographic Regions. The lowest number of records was found in the Northeast Occidental Atlantic and Parnaíba regions, which are also less well known territories in terms of their algal biodiversity. Figure 3 Number of algae and cyanobacteria species distributed by state and Brazilian geopolitical region. Figure 4 Seaweeds and marine cyanobacteria species richness distributed by Brazilian geopolitical region and taxonomic class. Rodriguésia 66(4):

123 1054 Menezes, M. et al. Table 3 Species richness of algae and cyanobacteria classes recorded in 2010 and 2015, distributed by environment. (*) Updates of the data published in 2010; (-) - Not included in 2010; (**) - Including Conjugatophyceae taxa; (1) - previously included in Rhodophyceae; (2) - including 2010 Prymnesiophyceae taxa. Classes Epicontinental Ambiente Marinho 2010* * 2015 Bacillariophyceae Bangiophyceae Charophyceae 419** Chlorodendrophyceae Chlorophyceae Chrysophyceae Coccolithophyceae Coleochaetophyceae Compsopogonophyceae Conjugatophyceae Cryptophyceae Cyanophyceae Dictyocophyceae Dinophyceae Euglenophyceae Florideophyceae Klebsormidiophyceae Mamiellophyceae Nephroselmidophyceae Pedinophyceae Phaeophyceae Porphyridiophyceae Prasinophyceae Raphidophyceae Stylonematophyceae Synurophyceae Trebouxiophyceae Ulvophyceae Xanthophyceae Total Rodriguésia 66(4):

124 Brazilian Algae and Cyanobacteria 1055 Table 4 Species richness of algae and cyanobacteria by Brazilian geopolitical region in 2010 and Year North Northeast Central-West Southwest South AM AP AC PA RO RR TO MA PI CE RN PB PE AL BA SE MT MS GO DF MG ES RJ SP PR SC RS Table 5 Species number of marine macroalgae and cyanobacteria by Brazilian geopolitical region and state in Region/States North Northeast Southwest South Class AP PA MA PI CE RN PB PE AL SE BA ES RJ SP PR SC RS Bangiophyceae Compsopogonophyceae Cyanophyceae Florideophyceae Stylonematophyceae Phaeophyceae Prasinophyceae Ulvophyceae Rodriguésia 66(4):

125 1056 Menezes, M. et al. Table 6 Species number of epicontinental algae and cyanobacteria by Brazilian geopolitical region and state in Regions/States North Central-West Northeast Southwest South Class AC AP RO RR AM PA TO GO DF MT MS MA PI CE RN PB PE AL SE BA MG ES RJ SP PR SC RS Bacillariophyceae Charophyceae Chlorophyceae Chrysophyceae Coleochaetophyceae Compsopogonophyceae Conjugatophyceae Cryptophyceae Cyanophyceae Dinophyceae Euglenophyceae Florideophyceae Klebsormidiophyceae Mamiellophyceae Nephroselmidophyceae Porphyridiophyceae Raphidophyceae Synurophyceae Trebouxiophyceae Ulvophyceae Xanthophyceae Rodriguésia 66(4):

126 Brazilian Algae and Cyanobacteria Table 7 Species number of the most species rich genera of algae and cyanobacteria recorded in Brazil in Class Genus Species number Conjugatophyceae Cosmarium 156 Bacillariophyceae Pinnularia 125 Euglenophyceae Trachelomonas 107 Conjugatophyceae Staurastrum 93 Bacillariophyceae Eunotia 72 Euglenophyceae Phacus 69 Dinophyceae Protoperidinium 66 Chlorophyceae Oedogonium 65 Bacillariophyceae Navicula 63 Conjugatophyceae Closterium 60 Dinophyceae Tripos 56 Bacillariophyceae Chaetoceros 50 Euglenophyceae Strombomonas 48 Conjugatophyceae Spirogyra 47 Conjugatophyceae Euastrum 47 Bacillariophyceae Nitzschia 46 Bacillariophyceae Surirella 41 Euglenophyceae Euglena 39 Bacillariophyceae Amphora 36 Dinophyceae Dinophysis 35 Synurophyceae Mallomonas 33 Cyanophyceae Phormidium 32 Conjugatophyceae Micrasterias 31 Conjugatophyceae Staurodesmus 29 Bacillariophyceae Coscinodiscus 28 Euglenophyceae Lepocinclis 28 Euglenophyceae Entosiphon 28 Chlorophyceae Chlamydomonas 27 Charophyceae Nitella 27 Bacillariophyceae Diploneis 25 Chlorophyceae Scenedesmus 25 Ulvophyceae Cladophora 24 Conjugatophyceae Pleurotaenium 23 Charophyceae Chara 23 Ulvophyceae Caulerpa 22 Cyanophyceae Scytonema 22 Dinophyceae Oxytoxum 22 Bacillariophyceae Gyrosigma 21 Bacillariophyceae Cocconeis 21 Dinophyceae Gymnodinium 21 Florideophyceae Ceramium 19 Total 1852 Rodriguésia 66(4):

127 1058 Menezes, M. et al. Figure 5 Species richness of epicontinental algae and cyanobacteria by Brazilian hydrographic region Discussion The number of records (1,250) added to the latest version of the Algae and Cyanobacteria (Bicudo & Menezes 2010) represents another important advancement for neotropical phycology, however we believe that there is still much research to be carried out. The present total of 4,747 species recorded for different environments and Brazilian states is becoming close to previous estimates by Bicudo et al. (1998) and Menezes & Bicudo (2009), of respectively 5,000 and 5,614 species. These estimates are still fairly distant from the real algal diversity to be found in the country, when the extense geographic gaps that have not been studied are taken into account. Some taxonomic groups have been subestimated as it is obviated below. Using the cyanobacteria as an example, taking worldwide figures of 5,000 known species and 3,000 species still to be described estimated by Guiry (2012), the 462 cyanobacteria species currently recorded for Brazil represent less than 10% of the known species, or slightly more than 5% of the total estimated. However Rocha (2006) estimated that there would be around 1,200 species in this group, and the figure recorded so far for Brazil is around 38% of this estimate. Of the new species of cyanobacteria and microalgae included in the Brazilian List between 2010 and 2015, seven belong to genus Brasilonema Fiore et al. (Sant Anna et al. 2011), one of each Scytonema C.Agardh ex Bornet & Flahault (Hentschke & Komarék 2014), Lemmermaniella Geitler (Gama Jr. et al. 2014), Chroococcus Nägeli (Gama Jr et al. 2014) and Eunotia Ehrenb. (Furhmann et al. 2013) for the Paraná hydrographic region, São Paulo state. A species of Symploca Kütz. ex Gomont was described for coastal Bahia (Caires et al. 2013). One species of Urosolenia Round & R.M.Crawford (Tremarin et al. 2013a) and two Encyonema Kütz. (da Silva et al. 2015) were described for For the Paraná hydrographic region while a species of Conjugatophyceae, Staurastrum Meyen ex Ralfs (Santos et al. 2013) and a Cyanophyceae belonging to the genus Pannus Hickel (Malone et al. 2014) were described for the Paraguay hydrographic region (pantanal of Nhecolândia, Mato Grosso do Sul state). For the Amazon hydrographic region the monotypic new genera Eunotioforma Kociolek & Burliga and Bicudoa C.E.Wetzel et al. (Wetzel et al. 2012a; Burliga et al. 2013), as well as a species of Tursiocola R.W.Holmes et al. (Wetzel et al. 2012b) and another of Aulacoseira Thwaites (Tremarin et al. 2013b) were described. A new species of Oocystis A.Braun (Ramos et al. 2015) was described from the West Atlantic hydrographic region. Three new species of Diatomaceae (one Eunotia and two Aulacoseira) described have wider distribution and span through two or more hydrographic regions (Metzeltin & Tremarin 2011; Tremarin et al. 2012; Tremarin et al. 2014). Regarding marine macroalgae, genus Laurenciella Cassano et al. (Florideophyceae) was proposed with basis on molecular evidence (Cassano et al. 2012), while four species of Hypnea J.V.Lamour. (Nauer et al. 2014a, b; Jesus et al. 2015), one of Osmundea Stackh. (Rocha-Jorge et al. 2013) and two Ulvophyceae, Codium pernambucensis Oliveira & S.M.B.Pereira (Carvalho et al. 2012) and Gayralia brasiliensis Pellizari et al. (Pellizzari et al. 2013) were described during the last five years. The Bacillariophyceae and the Florideophyceae are the most speciose groups of respectively microalgae and macroalgae, both showing an increase in species number during the last period, while the records of other groups, considered underestimated already in 2010 (Bicudo & Menezes 2010), such as Coccolithophyceae, Prasinophyceae and Xanthophyceae, remain unaltered. The number (33) of new species described for Brazil during the period means that around six species were described per year. When analysing the authors against the new species described, we found that the number of new species dividided by the number of Brazilian authors (48), gives a proportion close to one (0.7) over the last Rodriguésia 66(4):

128 Brazilian Algae and Cyanobacteria five years. However, the breakdown of species described per authors is heterogeneous (Tab. 1), showing that the productivity is concentrated by a handful of taxonomists with enough experience and/or access to the collections needed to perform this task. This highlights the insufficiency of specialized, well trained researchers able to describe the overwhelming diversity of organisms found in Brazil. On the other hand, according to Sangster & Luksenburg (2015), a new trend of increased time and effort employed by taxonomists to produce good quality, complete new descriptions implies in less frequent subsequent revisions of taxonomic concepts. Studies suggest that, while new taxa continue to be described at a steady rate, the number of taxonomists has actually increased (Pimm et al. 2010; Jope et al. 2011; Tancoigne & Dubois 2013) therefore the number of species described per taxonomist has decreased. Such studies conclude that the number of taxonomy researchers is not declining, proposing that the decline seen is in the productivity per taxonomist (Sangster & Luksenburg 2015). Summarizing, our results show an increase in the knowledge regarding algal biodiversity in Brazil during the last five years that has started to close the gap between better studied and less known regions. Even with Southeast and South regions maintaining their position as the most diverse areas through our better understanding of their algal flora (Bicudo & Menezes 2010), the present records for some states within the Northeast, such as Piauí and Sergipe, and the Central-western states of Mato Grosso, Goiás and Distrito Federal have shown that they also hold considerable algal biodiversity. However relatively modest, the discovery of new genera and species has increased and spread through the less known regions such as the Amazon, Northeast and Central-western. As seen for other biodiverse countries, increased rate of description of new taxa started to address the geographic bias caused by the polarization of taxonomic research in and around the largest economic centres worldwide (Tancoigne et al. 2011; Grieneisen et al ). On the other hand, in the case of algae and cyanobacteria, Brazil continues to face obstacles for the advancement in the knowledge of phycological diversity, be it because of global issues, such as the shortage of taxonomists specializing in this area (Guiry 2012; De Clerck et al. 2013) or, in the regional scale, such as the lack of laboratory facilities and nonavailability of biological collections where the 1059 diversity is higher (Grieneisen et al. 2014). Strategic policies have already been launched by the Brazilian government in order to accelerate the study of the country s biodiversity, however, these need to be strengthened and widened to ensure its mapping is more homogeneous and complete. Acknowledgements We would like to thank the coordinators of the Brazilian Floristic List for their support of the updating of the online list. To Daniela Zappi and the two anonymous reviewers for helpful comments and suggestions to the manuscript. References Alves, A.M.; Gestinari, L.M.S.; Oliveira, I.S. Moniz- Brito, K.L. & Moura, C.W.N The genus Cladophora (Chlorophyta) in the littoral of Bahia, Brazil. Nova Hedwigia 95: Alves-Da-Silva, S.M. & Hahn, A.T Lista das Euglenophyta registradas em ambientes de águas continentais e costeiras do Estado deo Rio Grande do Sul, Brasil. Iheringia, Série Botânica 55: Amado Filho, G.M.; Maneveldt, G.; Pereira Filho, G.; Manso, R.C.; Bahia, R.; Barreto, M.B.B. & Guimaraes, S.M.P.B Seaweed diversity associated with a Brazilian tropical rhodolith bed. Ciencias Marinas 36: Bahia, R.G.; Riosmena-Rodriguez, R.; Maneveldt, G.W. & Amado Filho, G.M Research note: First report of Sporolithon ptychoides (Sporolithales, Corallinophycidae, Rhodophyta) for the Atlantic Ocean. Phycological Research 59: Bahia, R.G.; Amado-Filho, G.M.; Maneveldt, G.W.; Adey, W.H.; Johnson, G.; Marins, B.V. & Longo, L.L Sporolithon tenue sp. nov. (Sporolithales, Corallinophycidae, Rhodophyta): A new rhodolithforming species from the tropical southwestern Atlantic. Phycological Research 62: Bahia, R.G.; Amado Filho, G.M. & Maneveldt, G.W. 2014a. Sporolithon molle (Heydrich) Heydrich (Sporolithales, Corallinophycidae, Rhodophyta): An addition to the atlantic flora found on a remote oceanic island. Cryptogamie. Algologie 35:7-14. Bahia, R.G.; Amado Filho, G.M.; Azevedo, J. & Maneveldt, G.W. 2014b. Porolithon improcerum (Porolithoideae, Corallinaceae) and Mesophyllum macroblastum (Melobesioideae, Hapalidiaceae): new records of crustose coralline red algae for the Southwest Atlantic Ocean. Phytotaxa 190: Bicudo, C.E.M.; Sant Anna, C.L.; Bicudo, D.C.; Pupo, D.; Castro-Pinto, L.S.; Azevedo, M.T.P.; Xavier, M.B.; Fujii, M.T.; Yokoya, N.S. & Guimarães, S.M.P.B O estudo das algas no Estado de São Paulo. In: Bicudo, C.E.M. & Shepherd, G.J. (eds.). Rodriguésia 66(4):

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132 Rodriguésia 66(4): DOI: / Bryophytes diversity in Brazil Denise P. Costa 1 & Denilson F. Peralta 2 Abstract The bryoflora of Brazil comprises 1,524 species, 117 families, and 413 genera (11 hornworts, 633 liverworts, and 880 mosses). The most diverse families of liverworts are: Lejeuneaceae (285 species), Lepidoziaceae (48), Frullaniaceae (37), Ricciaceae (36), Plagiochilaceae (27), Radulaceae and Metzgeriaceae (26 each), Lophocoleaceae (18), Aneuraceae (15), and Calypogeiaceae (13); while, for the mosses, we have: Sphagnaceae (83 species), Fissidentaceae (65) Pottiaceae (63), Dicranaceae (54), Bryaceae and Sematophyllaceae (53 each), Orthotrichaceae and Pilotrichaceae (51 each), Calymperaceae (48), and Hypnaceae (28). These large groups account for 71% of the Brazilian bryophyte species. Lejeuneaceae and Sphagnaceae are the families with highest number of endemic taxa (54 and 60 species). The Atlantic Rainforest presents the greatest number of species (1,337), followed by the Amazon Rainforest (570) and Cerrado (478). The highest number of endemic species (242) is associated with the Atlantic Rainforest, where the Dense Ombrophilous Forest concentrates 73% of the species with 62% endemism. The Southeastern region is the most diverse in number of species (1,228) and with more endemism (219). Most endangered species are restricted to the Atlantic Rainforest of southeastern Brazil, which is the diversity and endemism centre for mosses and liverworts. The information currently presented by the Brazilian List is close to the real bryophyte diversity found in the country. Key words: hornworts, liverworts, mosses, bryoflora, South America. Resumo A brioflora do Brasil conta com espécies, 117 famílias e 413 gêneros (11 antóceros, 633 hepáticas e 880 musgos). As famílias mais diversas de hepáticas são: Lejeuneaceae (285 espécies), Lepidoziaceae (48), Frullaniaceae (37), Ricciaceae (36), Plagiochilaceae (27), Radulaceae e Metzgeriaceae (26 cada), Lophocoleaceae (18), Aneuraceae (15) e Calypogeiaceae (13); e de musgos são: Sphagnaceae (83), Fissidentaceae (65), Pottiaceae (63), Dicranaceae (54), Bryaceae e Sematophyllaceae (53 cada), Orthotrichaceae e Pilotrichaceae (51 cada), Calymperaceae (48) e Hypnaceae (28), totalizando 71% das espécies de briófitas do país. Lejeuneaceae e Sphagnaceae são as famílias com maior número de endemismo (54 e 60 espécies). A Mata Atlântica apresenta o maior número de espécies (1.337), seguida pela Amazônia (570) e pelo Cerrado (478). Também apresenta o maior número de espécies endêmicas (242), com a Floresta Ombrófila Densa concentrando 73% dos táxons e 62% dos endemismos. A Região Sudeste é a mais diversa (1.228) e com maior endemismo (219). A maioria das espécies ameaçadas de extinção está restrita a Mata Atlântica do sudeste, que é um centro de diversidade e endemismo para musgos e hepáticas no país. As informações da Lista de Espécies da Flora do Brasil estão próximas da real diversidade de briófitas do país. Palavras-chave: antóceros, hepáticas, musgos, brioflora, América do Sul. Introduction The first checklist of the Brazilian bryophytes was published by Costa & Luizi-Ponzo (2010) treating three Divisions, or Phyla, in the country, Anthocerotophyta (hornworts), Marchantiophyta (liverworts), and Bryophyta (mosses), and represented a progress for the knowledge of the bryoflora in the country. The first detailed publication for Anthocerotophyta and Marchantiophyta featured in the Pars Prior of Martius and Eichler Flora Brasiliensis, with the treatments prepared by Nees ab Esenbeck (1833), and, slightly later, the treatment of Bryophyta published by Hornschuch (1840) in the same encyclopaedia. At that stage 275 species of bryophytes, comprising 81 liverworts 1 Rio de Janeiro Botanical Garden, R. Pacheco Leão 915, , Rio de Janeiro, RJ, Brazil. 2 Institute of Botany of São Paulo, Av. Miguel Estéfano 3687, , São Paulo, SP, Brazil. 3 Author for correspondence: denisepinheirodacosta@gmail.com

133 1064 Costa, D.P. & Peralta, D.F. and 194 mosses were recognized, representing around 18% of the currently recognized species. Following these publications, the major advances of the knowledge of these three divisions of plants have been published in a disperse manner, in different vehicles such as checklists, regional floras, and few taxonomic revisions and descriptions of new taxa. Catalogues compiling published data were published by Yano (1981, 1984, 1989, 1995) and Costa et al. (2011). Such works summarized all citations and treatments of species recorded for Brazil, and served as the base for Gradstein & Costa (2003) who recognized 600 species of liverworts and hornworts for Brazil, a number very close of the actually recognized to the country (94%). Concerning the mosses, the Brazilian Moss Guide has already been accepted for publication by The New York Botanical Garden Press. Around 800 species of mosses are recognized, a close number to what is actually recognized to the country (90%). Among the new studies who brought taxonomic and nomenclatural novelties after Costa & Luizi-Ponzo (2010) we can cite: Câmara (2011) completed the World revision of Taxithelium (Pylaisiadelphaceae), Majestyk (2011) published the Neotropical revision of Daltonia (Daltoniaceae), Reiner-Drehwald (2011) new synonyms to Ceratolejeunea (Lejeuneaceae), Olson et al. (2011) accomplished the World revision of Neckeraceae, Suarez & Schiavone (2011) revised South American Pohlia (Bryaceae), Uribe (2011) revised types of Frullania subgenus Meteoriopsis, Costa (2012) revised genus Streptopogon (Pottiaceae), Bastos (2012a,b,c,d and 2013) made new combinations and synonyms to Cheilolejeunea (Lejeuneaceae), Gradstein (2013) presented new synonyms to Omphalanthus and Aureolejeunea (Lejeuneaceae), Cano & Jimenez (2013) revised tribe Pleuroweisiae (Pottiaceae), Söderstrom et al. (2013) added synonyms to Lophocoleaceae, Costa (2014) presented synonyms to Brazilian Pottiaceae, and Gradstein & Ilkiu-Borges (2015) completed the World revision of the genus Odontoschisma (Cephaloziaceae). The addition of new records to the list of bryophytes in Brazil increased the knowledge of the diversity in this group, and emphasized the importance of new records or new taxa published by Costa et al. (2008), Ilkiu-Borges & Pereira Alvarenga (2008), Ilkiu- Borges & Gradstein (2008), Bastos (2011), Bordin et al. (2011), Câmara & Carvalho-Silva (2011), Peralta (2011), Ilkiu-Borges (2011), Câmara & Carvalho-Silva (2012), Brito & Ilkiu-Borges (2012a,b), Peralta & Boas-Bastos (2012), Peralta & Reiner-Drewald (2013), and Ayub et al. (2014). Floristic surveys have also contributed to the bryophyte knowledge and added new records to the species list of bryophytes in Brazil, such as: Ayub et al. (2014) that recorded Riccia boliviensis Jovet-Ast and R. iodocheila M. Howe to the country, and R. squamata Nees and R. subplana Steph. to Rio Grande do Sul State. This new species list of bryophytes has the main objectives add new data, including new informations and performing the correction of possible errors published in Costa & Luizi-Ponzo (2010). Methods The methodology used in this study was the same adopted for the other groups of plants and fungi of Brazil, summarized in Forzza et al. (2010). Regional lists and selected databases were evaluated by the specialists who accepted the invitation to contribute online to the project website at the Jardim Botânico do Rio de Janeiro (< jbrj.gov.br>). Each expert was tasked with the inclusion of new records and/or the revision and modification of previously entered records, in the light of the actual knowledge for his/her speciality group, including information about life forms, substrates, vegetation types, vouchers, etc. All information was based on checked herbaria species, therefore, for a taxon ranging throughout the Brazilian states, the specialist would have seen at least one specimen per state. The complete dataset used for the present analyses can be found as the Brazilian List of Bryophytes (see supplementary material < dx.doi.org/ /m9.figshare > - DOI: / ). Eight experts participated in data collection and contributed data in the following families and/ or genera (authors are listed in order of authorship): Entodontaceae (Diego K. Henriques & Denise Pinheiro da Costa), Fissidentaceae (Juçara Bordin), Frullaniaceae (Denilson F. Peralta), Lejeuneaceae (Denise Pinheiro da Costa), Cheilolejeunea and Rectolejeunea (Cid José Passos Bastos), Lepidoziaceae (Denilson F. Peralta), Lophocoleaceae (Denilson F. Peralta), Pilotrichaceae (Denise Pinheiro da Costa), Pottiaceae (Denise Pinheiro da Costa), Sematophyllaceae (Paulo Eduardo Aguiar Saraiva Câmara), Sphagnaceae (Denise Pinheiro da Costa) and Thuidiaceae (Denise Pinheiro da Costa & Abel Eustáquio R. Soares). The species data for the 108 remaining families were added by the two bryophyte coordinators, Denise Rodriguésia 66(4):

134 Bryophytes diversity in Brazil Pinheiro da Costa (liverworts and hornworts) and Denilson F. Peralta (mosses). It was also the coordinators responsibility to check and include information that was, for any reason, not complete by the end of March Results According to the latest dataset (see Brazilian List of Bryophytes; supplementary material < dx.doi.org/ /m9.figshare > - DOI: / ), the bryophyte flora of Brazil comprises 1,524 species, of which 11 are hornworts, 633 liverworts and 880 mosses. This number is only 1% higher than the previous record of 1,521 species (Costa & Luizi-Ponzo 2010) (Fig. 1). Due to recent advances in the classifications of bryophyte groups, 117 families and 413 genera are currently recognized versus 109 families and 395 genera recorded by Costa & Luizi-Ponzo (2010). The ten most diverse families of liverworts are: Lejeuneaceae (285 species), Lepidoziaceae (48), Frullaniaceae (37), Ricciaceae (36), Plagiochilaceae (27), Radulaceae (26), Metzgeriaceae (26), Lophocoleaceae (18), Aneuraceae (15), and Calypogeiaceae (13) (Fig. 2). For mosses, the ten most diverse families are: Sphagnaceae (83 species), Fissidentaceae (65), Pottiaceae (63), Dicranaceae (54), Bryaceae (53), Sematophyllaceae (53), Orthotrichaceae (51), Pilotrichaceae (51), Calymperaceae (48), and Hypnaceae (28) (Fig. 2). The results in 2015 differ in relation to data presented by Costa & Luizi-Ponzo (2010), with the liverworts of the Ricciaceae family exceeding Plagiochilaceae in number of species, Aneuraceae and Calypogeiaceae surpassing Jungermanniaceae, which now has less species (Fig. 3). For the mosses, Bryaceae, Pilotrichaceae, and Hypnaceae exceeded Fissidentaceae and Calymperaceae (Fig. 3). Regarding the richest families of liverworts in Brazil, we highlight Lejeuneaceae (285 species) containing 45% of all species recognized for this division in Brazil, while, for the mosses, three families, Sphagnaceae (83 species), Fissidentaceae (65 species), and Pottiaceae (63 species) together total 24% of the species recognized for this division in the country. Comparing the information published by Costa & Luizi-Ponzo (2010) to the present list, there was a small increase in the percentage of endemic species, from 18% to 19.6%. However, among the ten families of liverworts with highest number of endemic taxa, Lejeuneaceae remained the first 1065 one, with 54 species, followed by Ricciaceae (11), Metzgeriaceae (6), Lepidoziaceae (6), Frullaniaceae (5), Plagiochilaceae e Aneuraceae (2 each), Radulaceae, Lophocoleaceae, and Calypogeiaceae (1 each) (Fig. 2). Figure 1 Comparison between the number of species of Mosses, Liverworts and Hornworts between Costa & Luizi-Ponzo (2010) and the 2015 list. Figure 2 Total number of species in the ten most diverse families of liverworts, comparing data from Costa & Luizi-Ponzo (2010) and the 2015 list. Figure 3 Total number of species in the ten most diverse families of mosses, comparing data from Costa & Luizi-Ponzo (2010) and the 2015 list. Rodriguésia 66(4):

135 1066 Costa, D.P. & Peralta, D.F. Regarding the mosses, among the ten families with the highest number of endemic taxa, Sphagnaceae remained the first one, with 60 species, followed by Sematophyllaceae (17), Dicranaceae (15), Pottiaceae and Orthotrichaceae (11 each), Bryaceae and Pilotrichaceae (9 each), Hypnaceae (4), Calymperaceae (2), and Fissidentaceae (1) (Fig. 3). The present study recognizes 413 genera, with the 20 most diverse genera including 600 species, of which 149 species are endemic (Tab. 1). As in the previous study (Costa & Luizi-Ponzo 2010), the three most diverse genera continue to be Sphagnum (83 species), Fissidens (65), and Lejeunea (40). However, it is noteworthy that the total number of species in Sphagnum and Lejeunea has been reduced from respectively 93 and 42 species in 2010 to 83 and 40 in Allied to this decrease, the number of endemic species in these genera also changed, and in Sphagnum falling from 65 to 60, while in Lejeunea it increased from five to six species. Other variations were recorded in the following moss genera: Fissidens, Campylopus, and Sematophyllum with respectively three, seven, and six endemic species in 2010 (from a total of 16 taxa) and had a respective increase to six, eight, and eight species (from a total of 22 taxa) in Liverwort genera such as Cololejeunea and Cheilolejeunea had a large increase in the number of species and endemic taxa, with an increase from 17 species (one endemic) in 2010 to 37 (eight endemic) in Cololejeunea, and an increase from the 16 species (1 endemic) previously recorded for Cheilolejeunea to 37 species (14 endemic) in Table 1 List of the twenty richest genera of bryophytes (10 of mosses and 10 of liverworts), with their numbers of species and endemism, comparing data from Costa & Luizi-Ponzo (2010) and Genera Species number 2015 Endemism 2015 Species number 2010 Endemism 2010 Mosses Sphagnum Fissidens Campylopus Syrrhopodon Sematophyllum Bryum Macromitrium Calymperes Schlotheimia Philonotis Liverworts Lejeunea Cololejeunea*** Cheilolejeunea Frullania Riccia Plagiochila Metzgeria Radula Drepanolejeunea Bazzania Total *** formerly included genus Aphanolejeunea was transferred to Cololejeunea (Pócs et al. 2014). --- Genera not listed by Costa & Luizi-Ponzo (2010). Rodriguésia 66(4):

136 Bryophytes diversity in Brazil Classification changes in some genera of Lejeuneaceae meant that several species had to be transferred to other genera, with Aphanolejeunea A. Evans being subsumed into Cololejeunea (Spruce) Schiffn. (Pócs et al. 2014), (Tab. 1). The biomes with the highest number of species continues to be the Atlantic Rainforest, with 1,337 species, followed by the Amazon Rainforest (570 species), Cerrado (478), Pantanal (176), Pampa (120), and Caatinga (96). It is worth mentioning that the bryophyte diversity for the Caatinga biome were included in the introduction of Forzza et al. (2010) but were not part of the analysis presented by Costa & Luizi-Ponzo (2010) in the same book. The Atlantic Rainforest is the biome with the largest number of endemic species (242), followed by Cerrado (63), Amazon Rainforest (52), Pampa (16), Caatinga (8), and Pantanal (6) (Fig. 4). The greatest diversity in the Atlantic Rainforest is found in the Ombrophilous Forest, and this vegetation type concentrates 73% of the total species and 62% of the endemism found in this biome. Ombrophilous Dense Forest and Ombrophilous Mixed Forest contain the highest richness (1,138 and 632 species, respectively), followed by the Amazonian Terra Firme Forest (463 species) vegetation, where 38 endemic species occur. In the Cerrado, the Gallery Forest presents a large number of species (309), with 37 endemic from this vegetation type. Regarding the geopolitical regions, there was a significant increase in the number of new records, however the Southeast region continued to be the most diverse one, with 1,228 species in the present survey (vs. 1,205 in Costa & Luizi-Ponzo 2010), as well as with the high number of endemism (219 taxa). The other regions also increased their representation: the South showed 843 species (vs. 796 in Costa & Luizi-Ponzo 2010); Northeast, 689 (vs. 609); North, with 601 species (vs. 585.); and Central-west, with 496 (vs. 460) (Fig. 5). Taking into account the diversity within the federation states, changes over the last five years did not change the position of Rio de Janeiro, São Paulo and Minas Gerais as the leading states in terms of species richness. Rio de Janeiro and São Paulo states count with 900 species each, while Minas Gerais has 766 species records (vs. 873, 840 and 687 in the previous survey), confirming the Southeast region as the one with presents the highest richness. When compared to previous data (Costa e Luizi-Ponzo 2010), the largest additions occurred in Minas Gerais 1067 state (79 species), followed by Paraná (72 species), Pernambuco (67 species), and Bahia (66 species), probably due to increased knowledge of the flora in these states which are among the ten richest in the country. The smallest additions correspond to the North region states, Amapá (two species), Acre and Roraima (five species each), and Tocantins (10 species), and the northeastern states as Piauí (seven species), Sergipe and Paraíba (12 species each), Maranhão (17 species), and Rio Grande do Norte (19 species), the latter state with the lowest diversity, adding up to only 35 species (Tab. 2). The analysis of life-forms shows that most species that occur in Brazil have foliose habit (717). In relation to the remaining life-forms, a predominance of tuft (421 species), followed by mat (371 species), and weft (237 species) was found (Fig. 6). Regarding the type of substrate colonized by bryophytes the predominance of corticicolous (797 species), followed by terricolous (640 species), rupicolous (550 species), and epixilous (210 species). Meanwhile, 120 leaf-dwelling, or epiphyllous species belong mostly Figure 4 Total number of bryophyte species and endemic taxa by Biome. Figure 5 Comparison of the bryophytes species number by geographic regions, comparing data from Costa & Luizi-Ponzo (2010) and Rodriguésia 66(4):

137 1068 Costa, D.P. & Peralta, D.F. Table 2 Number of species and endemism of bryophytes by state and federal district, comparing comparing data from Costa & Luizi-Ponzo (2010) and The endemism refers to species that occur only in these states. States Species number 2015 Endemism 2015 Species number 2010 Endemism 2010 Rio de Janeiro São Paulo Minas Gerais Rio Grande do Sul Santa Catarina Paraná Bahia Amazonas Espírito Santo Pernambuco Pará Mato Grosso Goiás Mato Grosso do Sul Roraima Acre Distrito Federal Ceará Rondônia Amapá Paraíba Alagoas Maranhão Piauí Tocantins Sergipe Rio Grande do Norte to the family Lejeuneaceae, especially in the genera Ceratolejeunea, Cololejeunea, Drepanolejeunea, Lejeunea, Leptolejeunea, and Prionolejeunea (Fig. 7). Considering the three biomes with the highest number of mosses and liverworts species, the Atlantic Rainforest is the one with the greatest diversity of life forms and types of substrates, followed by the Amazon Rainforest and Cerrado (Figs. 8 and 9), where the foliose, tuft and mat life-forms predominate in conjunction with corticicolous, epixilous and terricolous substrates. In the Caatinga, Pantanal, and Pampa the commonest life-forms are tuft and weft, and the substrates are mostly corticicolous, terricolous and rupicolous (Figs. 8 e 9). The majority of threatened species found in the Brazilian Red Book (Martinelli & Moraes 2013) occur restricted to only one or two states in southeastern Brazil. Many of these species have reduced Occupation Area of Occurrence (AOO) or Extension Area of Occurrence (EOO), and occur in the Atlantic Rainorest, alongside all endangered species listed as threatened. Rodriguésia 66(4):

138 Bryophytes diversity in Brazil Figure 6 Total number of bryophytes species by life-form. Figure 7 Total number of bryophytes species by substrate. Figure 8 Species distribution by life-form and biome. Figure 9 Species distribution by substrate and biome Discussion The recognition of 1,524 species of mosses, liverworts, and hornworts for Brazil represents an advance in the knowledge of these three divisions. These figures are certainly close to the real diversity as they approach the estimates presented by Gradstein & Costa (2003) for liverworts and hornworts ( species) and Costa et al. (2011) for mosses (892 species), with an addition of only three species over what was previously recorded by Costa & Luizi-Ponzo (2010). The difference in species number is small however the number of families and genera recognized are consistently different, with 117 families and 413 genera versus 109 families and 395 genera listed in Costa & Luizi-Ponzo (2010). The difference in family and genera number is due to recently published studies that focused on large, previously understudied groups including changes of circumscription at genus and family level and presenting vast synonymy. These studies clarified important gaps related to the taxonomy and geographic distribution of taxa. In the genus Sphagnum, for example, 93 species were recognized in the list of Costa & Luizi-Ponzo (2010), while only 83 are recognized in the present list. This genus is currently under revision (Costa, unpublished). On the other hand, many groups still are in need of detailed study, not only for species circumscription, but also to fully understand the geographic distribution of their taxa. These include liverworts such as Bazzania, Frullania, Lejeunea p.p., Plagiochila p.p., and Riccardia and moss genera Callicostella, Cyclodictyon, Hyophila, Macromitrium, Mittenothamnium, Schlotheimia, and Sematophyllum p.p.. These groups lack recent taxonomic treatments including identification keys, descriptions, and illustrations. Some of these genera are already under study, but the data are not yet available to be included in the present work. It is probable that the number of species featured for these groups is underestimated in our list. The Atlantic Rainforest, the biome with the highest number of species including the largest number of endemic and endangered species, is recognized as one of the hotspots in the planet (Mittermeier et al. 1998, 2004), and has already been pointed out as one of the main centres of diversity and endemism for mosses and liverworts Rodriguésia 66(4):

139 1070 Costa, D.P. & Peralta, D.F. in the Brazil (Gradstein & Costa 2003; Costa et al. 2011). This high diversity is attributed to different environmental conditions occurring in mountainous tropical areas (humid climate with rains distributed throughout the year; fog in the mountains; ample variation of average temperatures from the lowland to the high mountains), favouring a greater number of microhabitats, high rainfall, and topographic complexity (mountain ranges with cloud forest in the Altitude Grasslands; shoreline fringed by sandy beaches, rocky shores, salt marshes and mangroves), which leads consequently to greater floristic diversity (Pócs 1982). Conclusions We believe that the information compiled for this paper approaches closely the real diversity of bryophytes in the country. However, gaps are still recognized in regions that need increased collection efforts and remain insufficiently sampled. Further studies developed in these regions will certainly yield new records possibly new species of mosses, hornworts and liverworts for the Brazilian Flora. References Ayub, D.M.; Costa, D.P. & Santos, R.P Additions to the Ricciaceae flora of Rio Grande do Sul, including two remarkable records for the Brazilian liverwort flora. Phytotaxa 161: Bastos, C.J.P Cheilolejeunea ornata (Lejeuneaceae), a new species from Brazilian Atlantic Forest. Journal of Bryology 33: Bastos, C.J.P. 2012a. Nomenclatural notes on the genus Rectolejeunea A. Evans (Lejeuneaceae, Marchantiophyta). Journal of Bryology 34: Bastos, C.J.P. 2012b. New combinations and synonyms in Cheilolejeunea (Spruce) Schiffn. (Lejeuneaceae, Marchantiophyta). Journal of Bryology 34: Bastos, C.J.P. 2012c. Synonymy and notes on the occurrence of Cheilolejeunea intertexta (Lindenb.) Steph. (Lejeuneaceae, Marchantiophyta) in Neotropics. Journal of Bryology 34: Bastos, C.J.P. 2012d. Type studies on Cheilolejeunea (Spruce) Schiffn. (Lejeuneaceae): Brazilian species described by Stephani. Journal of Bryology 34: Bastos, C.J.P Cheilolejeunea. In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. Available in ( floradobrasil/ FB126233). Access on 6 May Bastos, C.J.P. & Gradstein, S.R Two new species of Cheilolejeunea (Spruce) Schiffn. (Lejeuneaceae) from Brazil: C. lacerata sp. nov. and C. rupestris sp. nov. Journal of Bryology 28: Bordin, J.; Pursell, R.A. & Yano. O Fissidens pseudoplurisetus sp. nov. (Fissidentaceae, subgenus Aloma), from the Atlantic forest, Brazil. The Bryologist 114: Brito, E.S. & Ilkiu-Borges, A.L. 2012a. A new species of Ceratolejeunea Jack & Steph. (Lejeuneaceae, Jungermanniopsida) from a remnant of Amazonian forest in Maranhão, Brazil. Nova Hedwigia 95: Brito, E.S. & Ilkiu-Borges, A.L. 2012b. Primeiro registro de Cololejeunea panamensis para a America do Sul. Rodriguésia 63: 1-3. Câmara, P.E.A.S A review of Taxithelium (Pylaisiadelphaceae). Systematic Botany 36: Câmara, P.E.A.S. & Carvalho-Silva, M Taxithelium juruense (Broth.) Broth. (Pylaisiadelphaceae) an endangered Brazilian endemic, with notes on the genus Taxithelium for Brazil. Acta Botanica Brasilica 25: Câmara, P.E.A.S. & Carvalho-Silva, M A new species of Taxithelium (Pylaisiadelphaceae) from Brazil. The Bryologist 115: Cano, M.J. & Jimenez, J.A A taxonomic revision of the tribe Pleuroweisiae (Pottiaceae, Bryophyta) in South America. Phytotaxa 143: Costa, D.P Validation of the new species of Streptopogon (Pottiaceae, Bryophyta) and a synopsis of the genus for Brazil. Systematic Botany 37: Costa, D.P New synonyms for South American/ Brazilian Pottiaceae (Bryophyta). Phytotaxa 167: Costa, D.P.; Pôrto, K.C.; Luizi-Ponzo, A.P.; Ilkiu-Borges, A.L.; Bastos, C.J.P.; Câmara, P.E.A.S.; Peralta, D.F.; Bôas-Bastos, S.B.V.; Imbassahy, C.A.A.; Henriques, D.K.; Gomes, H.C.S.; Rocha, L.M.; Santos, N.D.; Siviero, T.S.; Vaz-Imbassahy, T.F. & Churchill, S.P Synopsis of the Brazilian moss flora: checklist, distribution and conservation. Nova Hedwigia 93: Costa, D.P. & Luizi-Ponzo, A.P As briófitas do Brasil. In: Forzza et al. (orgs.). Catálogo de Plantas e Fungos do Brasil. Vol.1. Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro. Pp Costa, D.P.; Pôrto, K.C.; Luizi-Ponzo, A.P.; Ilkiu-Borges, A.L.; Bastos, C.J.P.; Câmara, P.E.A.S.; Peralta, D.F.; Bôas-Bastos, S.B.V.; Imbassahy, C.A.A.; Henriques, D.K.; Gomes, H.C.S.; Rocha, L.M.; Santos, N.D.; Siviero, T.S.; Vaz-Imbassahy, T.F. & Churchill, S.P Synopsis of the Brazilian moss flora: checklist, distribution and conservation. Nova Hedwigia 93: Costa, D.P.; Santos, N.D. & Váña, J A New Species of Cylindrocolea (Cephaloziellaceae) from Brazil. The Bryologist 111: Rodriguésia 66(4):

140 Bryophytes diversity in Brazil Gradstein, S.R Notes on Early land plants today 22. New combinations and new synonymy in Omphalanthus and Aureolejeunea (Lejeuneaceae, Marchantiophyta). Phytotaxa 76: Gradstein, S.R. & Costa, D.P The Hepaticae and Anthocerotae of Brazil. Memoirs of The New York Botanical Garden 87: Forzza, R.C.; Baumgratz, J.F.A.; Bicudo, C.E.M.; Canhos, D.A.L.; Carvalho Jr., A.A.; Costa, A.F.; Costa, D.P.; Hopkins, M.; Leitman, P.M.; Lohmann, L.G.; Maia, L.C.; Martinelli, G.; Menezes, M.; Morim, M.P.; Nadruz-Coelho, M.A.; Peixoto, A.L.; Pirani, J.R.; Prado, J.; Queiroz, L.P.; Souza, V.C.; Stehmann, J.R.; Sylvestre, L.; Walter, B.M.T. & Zappi, D. (eds.) Catálogo de plantas e fungos do Brasil. 2 vols. Andrea Jakobsson Estúdio / Jardim Botânico do Rio de Janeiro, Rio de Janeiro. 1699p. Gradstein, S.R. & Ilkiu-Borges, A.L A taxonomic revision of the genus Odontoschisma (Marchantiophyta, Cephaloziaceae). Nova Hedwigia 99: Hornschuch, C.F Musci. In: C.F.P. Martius (ed.). Flora Brasiliensis seu enumeratio plantarum in Brasilia hactenus detectarum quas suis aliorumque botanicorum studiis descriptas et methodo naturali digestas partim icone illustratas. Stuttgartiae et Turbingae, J.G. Cottal. Vol. 1, part 2, pp Ilkiu-Borges, A.L On Pycnolejeunea gadsteinnii (Lejeuneaceae) a new species from Brazil. Boletim do Instituto de Botânica de São Paulo 21: 1-4. Ilkiu-Borges, A.L. & Gradstein, S.R A new species of Cheilolejeunea (Spruce) Schiffn. (Lejeuneaceae) from Cerro de la Neblina, Venezuela. Nova Hedwigia 87: Ilkiu-Borges, A.L. & Pereira-Alvarenga, L.D On Ceratolejeunea atlantica, a new species of Lejeuneaceae (Jungermanniopsida) from Brazil. Nova Hedwigia 86: Majestyk, P A taxonomic treatment of Daltonia (Musci: Daltoniaceae) in the Americas. Journal of the Botanical Research Institute of Texas 5: Martinelli, G. & Moraes, M.A. (orgs.) Livro vermelho da flora do Brasil. Andrea Jakobsson & Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, Rio de Janeiro. 1100p. Mittermeier, R.A.; Myers, N.; Thomsen, J.B.; Fonseca, G.A.B. da & Olivieri, S Biodiversity hotspots and major tropical wilderness areas: approaches to setting conservation priorities. Conservation Biology 12: Mittermeier, R.A.; Robles-Gil, P.R.; Hoffmann, M.; Pilgrim, J.; Brooks, T.; Mittermeier, C.G.; Lamoreux, J. & Fonseca, G.A.B Hotspots revisited. Earth s biologically richest and most endangered terrestrial ecoregions. CEMEX, Washington, DC. 392p Nees ab Esenbeck, C.G Hepaticae. In: Martius, C.F.P. (ed.). Flora Brasiliensis seu enumeratio plantarum in Brasilia tam sua ponte quam accedente cultura provenientium quas in itineré auspicius Maximilianai Joseph I. bavariae regis annis peracto collegit, partim descript. Stuttgartiae et Turbingae, J.G. Cottal. Vol. 1, pars prior, pp Olsson, S.; Enroth, J.; Buchbender, V.; Hedenäs, L.; Huttunen, S. & Quandt, D Neckera and Thamnobryum (Neckeraceae, Bryopsida) paraphyletic assemblages. Taxon 60: Peralta, D.F Ocorrência do gênero Conostomum Sw. (Bartramiaceae) no Brasil. Acta Botanica Brasilica 25: Peralta, D.F. & Boas-Bastos, S.V Occurrence of the genus Pterobryopsis M. Fleisch. (Pterobryaceae) in Brazil. Revista Brasileira de Botânica 35: Peralta, D.F. & Reiner-Drewald, M.E Cheilolejeunea laciniata (Lejeuneaceae, Marchantiophyta), a new species from Southeastern Brazil. The Bryologist 116: Pócs, T Tropical Forest Bryophytes. In: Smith, A.J.E. Bryophyte Ecology. Chapman and Hall, New York. Pp Pócs, T.; Bernercker, A. & Tixier, P Synopsis and key to species of Neotropical Cololejeunea (Lejeuneaceae). Acta Botanica Hungarica 562: Reiner-Drehwald, M.E Studies on Neotropical Lejeuneaceae (Jungermanniopsida). New synonyms and Ceratolejeunea temnantha (Spruce) comb. nov. Cryptogamie, Bryologie 32: Soderstrom, L.; Váňa, J.; Crandall-Stotler, B.; Stotler, R.; Hagborg, A. & Konrat, M Notes on land plants today 43. New combinations in Lophocoleaceae (Marchantiophyta) Phytotaxa 122: Suarez, G.M. & Schiavone, M.S Pohlia Hedw section Pohlia (Bryaceae) in Central and South America. Nova Hedwigia 92: Uribe-M, J Type studies on Frullania subgenus Meteoriopsis. VI. Subgeneneric affiliation of selected Asiatic species previously assigned to subg. Meteoriopsis. Caldasia 33: Yano, O Checklist of Brazilian mosses. The Journal Hattori Botanical Laboratory 50: Yano, O Checklist of Brazilian liverworts and hornworts. The Journal Hattori Botanical Laboratory 56: Yano, O An additional checklist of Brazilian bryophytes. The Journal Hattori Botanical Laboratory 66: Yano, O A new additional annoted checklist of Brazilian bryophytes. The Journal Hattori Botanical Laboratory 78: Artigo recebido em 06/04/2015. Aceito para publicação em 02/09/2015. Rodriguésia 66(4):

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142 Rodriguésia 66(4): DOI: / Diversity of ferns and lycophytes in Brazil Jefferson Prado 1, Lana da S. Sylvestre, Paulo H. Labiak, Paulo G. Windisch, Alexandre Salino, Iva C.L. Barros, Regina Y. Hirai, Thaís E. Almeida, Augusto C.P. Santiago, Maria A. Kieling-Rubio, Anna Flora de N. Pereira, Benjamin Øllgaard, Carla G.V. Ramos, John T. Mickel, Vinicius A.O. Dittrich, Claudine M. Mynssen, Pedro B. Schwartsburd, João Paulo S. Condack, Jovani B.S. Pereira, Fernando B. Matos Abstract This compilation of ferns and lycophytes in Brazil is an update of the one published in 2010 in Catálogo de Plantas e Fungos do Brasil. The methodology consisted in collecting data from regional checklists, taxonomic revisions, and selected databases. Invited specialists improved the list accessing a website housed at the Jardim Botânico do Rio de Janeiro. The results show 1,253 species: 1,111 of ferns and 142 of lycophytes. This number is 6.5% higher than the previous one (1,176 spp.). The percentage of endemic species decreased from 38.2% to 36.7%. We recognized 36 families and 133 genera (vs. 33 families, 121 genera in 2010). The 10 most diverse families are Pteridaceae (196 spp.), Dryopteridaceae (179), Polypodiaceae (164), Hymenophyllaceae (90), Thelypteridaceae (86), Aspleniaceae (78), Lycopodiaceae (64), Selaginellaceae (55), Anemiaceae (51), and Cyatheaceae (45). The three most diverse genera are still Elaphoglossum (87 spp.), Thelypteris (85), and Asplenium (74). The richest phytogeographic domain continues to be in the Atlantic Rainforest with 883 species which also has the largest number of endemic and threatened species, followed by the Amazon Rainforest (503), Cerrado (269), Pantanal (30), Caatinga (26), and Pampa (eight). Minas Gerais remains as the richest state (657 spp. vs. 580 in 2010). Key words: Flora, Taxonomy, Endemic species, Atlantic Rainforest, Amazon Rainforest. Resumo Esta compilação de samambaias e licófitas do Brasil é uma atualização daquela de 2010, no Catálogo de Plantas e Fungos do Brasil. A metodologia consistiu na reunião de dados de listas regionais, revisões de grupos e bancos de dados selecionados. Especialistas convidados melhoraram a lista através do acesso a um sítio da web do Jardim Botânico do Rio Janeiro. Os resultados apontam uma diversidade de espécies, sendo samambaias e 142 licófitas. Este número é 6,5% maior que o anterior (1.176 espécies). As espécies endêmicas decresceram de 38,2% para 36,7%. Foram reconhecidas 36 famílias e 133 gêneros (vs. 33 famílias, 121 gêneros em 2010). As dez famílias mais diversas são: Pteridaceae (196 espécies), Dryopteridaceae (179), Polypodiaceae (164), Hymenophyllaceae (90), Thelypteridaceae (86), Aspleniaceae (78), Lycopodiaceae (64), Selaginellaceae (55), Anemiaceae (51) e Cyatheaceae (45). Os três gêneros mais diversos continuam sendo Elaphoglossum (87 espécies), Thelypteris (85) e Asplenium (74). O Domínio Fitogeográfico mais rico continua sendo a Mata Atlântica (883 espécies) e também com mais espécies endêmicas e ameaçadas, seguido pela Amazônia (503 espécies), Cerrado (269), Pantanal (30), Caatinga (26) e Pampa (oito). Minas Gerais permanece como o estado com maior riqueza (657 espécies vs. 580 em 2010). Palavras-chave: Flora, Taxonomia, Espécies endêmicas, Floresta Atlântica, Floresta Amazônica. The full authorship of this article can be found in Appendix 1. 1 Author for correspondence: jprado.01@uol.com.br

143 1074 Prado, J. et al. Introduction The publication of the list of ferns and lycophytes of Brazil in 2010 (Prado & Sylvestre 2010) represented a major advance in the knowledge of these groups of plants in the country. Until that date there was no detailed compilation of the diversity of these plants, other than that published in the Flora Brasiliensis by Martius & Eichler, whose treatments were developed by Spring (1840 Lycopodineae), Sturm (1859 Ophioglossaceae, Marattiaceae, Osmundaceae, Schizaeceae, Gleicheniaceae, and Hymenophyllaceae), and Baker (1870, Cyatheaceae and Polypodiaceae). This flora had recognized 576 species, representing less than half of the number of species currently known. Since the publication of the Flora Brasiliensis, major advances in the knowledge of ferns and lycophytes have been reported in a fragmented way, through the publication of regional floras, taxonomic revisions of groups, checklists, as well as descriptions of new taxa. These data were compiled and included in the first version of the Catálogo de Plantas e Fungos do Brasil, which reported the occurrence of 1,176 species, being 123 of lycophytes and 1,053 of ferns (Prado & Sylvestre 2010). After this compilation, new studies appeared, presenting taxonomic and nomenclatural novelties, such as (in alphabetical order of family): Mickel (2015, in press) which dealt with the taxonomic revision of Anemia (Anemiaceae), a very diverse and well represented group in Brazil; Lehnert (2012) studied species of Cyathea 1 2-pinnate (Cyatheaceae) and Lehnert & Weigand (2013) presented a review of Cyathea corcovadensis group; Schwartsburd & Prado (2014) presented a review of Hypolepis rugosula group (Dennstaedtiaceae) and Schwartsburd et al. (2014) revealed the occurrence of new taxa of Pteridium (Dennstaedtiaceae) in Brazil. Regarding the Dryopteridaceae, one of the richest families in Brazil, we highlight the studies by Vasco (2011) and Vasco et al. (2013) on Elaphoglossum subsection Muscosa and section Squamipedia; Matos & Mickel (2014), who reviewed Elaphoglossum section Polytrichia; Sundue et al. (2013) presented a synopsis of Rumohra in Brazil, with the description of a new species and Prado et al. (2014) demonstrated the occurrence of another species of Dryopteris in Brazil, also treating the genus in the country. In Gleicheniaceae, the revision of Sticherus published by Gonzales & Kessler (2011), brought new combinations that have altered the diversity of this group in Brazil. In Polypodiaceae, several taxonomic revisions have been published, especially for the group of the grammitid ferns, with the description of new genera and species. The new genera Leucotrichum (Labiak et al. 2010) and Stenogrammitis (Labiak 2011) were segregated from Lellingeria. Hirai et al. (2011) described Moranopteris and later Hirai & Prado (2012) presented a taxonomic revision for this genus. Finally, Labiak (2013), Lehnert (2013), and Moguel-Velázquez & Kessler (2013) presented the revisions of the genera Lellingeria, Melpomene, and Alansmia, respectively, in a single volume of the Flora Neotropica series. For the Pteridaceae, Link-Pérez & Hickey (2011) reviewed the group of Adiantopsis radiata and for Brazil they described a new species (A. timida Link-Pérez & Hickey) and recognized another one (A. trifurcata (Baker) Link-Pérez & Hickey). Cochran et al. (2014) described a new genus (Tryonia), segregated from Eriosorus/ Jamesonia to the Brazilian Atlantic Rainforest. More specific contributions, such as descriptions of new taxa, also added new names to the list of ferns and lycophytes of Brazil and improved the estimates of the diversity of the group: in Athyriaceae, new species in Diplazium (Mynssen & Matos 2012; Mynssen & Sylvestre 2013); in Blechnaceae, a new species of Blechnum (Dittrich et al. 2012); in Dennstaedtiaceae, new species of Hypolepis (Schwartsburd et al. 2012); in Dryopteridaceae, a new genus (Mickelia) and several new combinations (Moran et al. 2010), and new species for Elaphoglossum (Kieling-Rubio & Windisch 2012; Melo & Salino 2012); in Isoetaceae, new species of Isoetes (Pereira et al. 2012; Pereira & Labiak 2013); in Lycopodiaceae, new species and new combinations of Diphasiastrum, Lycopodiella, Phlegmariurus, and Pseudolycopodiella (Øllgaard 2012a; Øllgaard 2012b); in Polypodiaceae, nomenclatural novelties in Pleopeltis (Prado & Hirai 2011; Schwartsburd 2014), and nomenclatural novelties and new species of Serpocaulon (Schwartsburd & Smith 2013) and Stenogrammitis (Labiak 2011); in Pteridaceae, also nomenclatural novelties in Adiantopsis (Schuettpelz et al. 2014), a new species in Adiantum (Prado & Hirai 2013), and nomenclatural novelties in Jamesonia (Christenhusz et al. 2011); in Thelypteridaceae, new species in Thelypteris (Matos et al. 2010; Salino et al. 2011; Salino et al. 2014). Rodriguésia 66(4):

144 Diversity of ferns and lycophytes in Brazil Floristic surveys have also contributed to the increased knowledge of the group, contributing new names to the list of ferns and lycophytes of Brazil. Biral & Prado (2012) revealed the occurrence of Pellaea ovata (Desv.) Weath. in São Paulo State, previously known only to the Andean region. Carvalho et al. (2012) reported the occurrence of several species of Pico da Neblina Massif region known only from the Venezuelan side that had not been yet cited for Brazil. Cnemidaria spectabilis (Kunze) R.M. Tryon var. spectabilis [Cyathea spectabilis (Kunze) Domin] was recorded for the Brazilian Amazon area for the first time by Góes-Neto & Pietrobom (2012). It was previously known only for neighbouring countries of Brazil, such as: Venezuela, Guyana, Suriname, and French Guiana. The main goals of this new compilation for ferns and lycophytes of Brazil are to refine the data gathered and correct any misconceptions published for the group in the Catálogo de Plantas e Fungos do Brasil (Prado & Sylvestre 2010), and contribute with new information on Brazilian plant diversity. Methodology The methodology used in this study was basically the same as that adopted for the other groups enumerated by Forzza et al. (2010). Regional lists and selected databases were consolidated and made available to the invited experts on a website at the Jardim Botânico do Rio de Janeiro. Each expert had the task to include new records and modify data from previous records in the light of the latest knowledge for his/her speciality group. The responsibility of the coordinators of the group of ferns and lycophytes was to check and complete the data that eventually was not included by experts. Further details regarding the preparation of the new version of the Brazilian List, which was updated until March 2015 can be found in BFG (2015). The complete dataset used for the present analyses can be found as the Brazilian List of Ferns and Lycophytes (see supplementary material < dx.doi.org/ /m9.figshare > - DOI: / ). The 20 experts who participated in this step contributed to the updates in the following families (authors are listed in order of authorship): Anemiaceae (J.T. Mickel, I.C.L. Barros, A.C.P. Santiago, A.F.N. Pereira & P.H. Labiak), Aspleniaceae (L.S. Sylvestre), Athyriaceae (C.M. Mynssen), Blechnaceae (V.A.O. Dittrich & A. Salino), Culcitaceae (J.P.S. Condack), Cyatheaceae (P.G. Windisch & A.C.P. Santiago), 1075 Cystopteridaceae (C.M. Mynssen), Dennstaedtiaceae (P.B. Schwartsburd), Dicksoniaceae (J.P.S. Condack), Dryopteridacae (J. Prado, L.S. Sylvestre, A. Salino, J.P.S. Condack, P.G. Windisch, R.Y. Hirai, T.E. Almeida & M.A. Kieling-Rubio), Equisetaceae (A. Salino & T.E. Almeida), Gleicheniaceae (F.B. Matos), Hemidictyaceae (C.M. Mynssen), Hymenophyllaceae (P.G. Windisch), Isoetaceae (J.B. Pereira, J. Prado & L.S. Sylvestre), Lindsaeaceae (J. Prado), Lomariopsidaceae (I.C.L. Barros, A.C.P. Santiago & A.F.N. Pereira), Lycopodiaceae (P.G. Windisch, C.G.V. Ramos & B. Øllgaard), Lygodiaceae (I.C.L. Barros, A.C.P. Santiago & A.F.N. Pereira), Marattiaceae (P.H. Labiak), Marsileaceae (P.G. Windisch), Metaxyaceae (L.S. Sylvestre), Oleandraceae (L.S. Sylvestre), Ophioglossaceae (J.P.S. Condack & L.S. Sylvestre), Osmundaceae (L.S. Sylvestre), Plagiogyriaceae (J.P.S. Condack), Polypodiaceae (P.H. Labiak, R.Y. Hirai & T.E. Almeida), Psilotaceae (L.S. Sylvestre), Pteridaceae (J. Prado), Saccolomataceae (P.B. Schwartsburd), Salviniaceae (A. Salino & T.E. Almeida), Schizaeaceae (I.C.L. Barros, A.C.P. Santiago & A.F.N. Pereira), Selaginellaceae (R.Y. Hirai), Tectariaceae (J. Prado), Thelypteridaceae (A. Salino & T.E. Almeida), and Woodsiaceae (C.M. Mynssen). Results The diversity of ferns and lycophytes, in this compilation (see Brazilian List of Ferns and Lycophytes; supplementary material - DOI: xxxxxx), adds up to a total of 1,253 species, divided between 1,111 ferns and 142 lycophytes. This number is 6.5% higher than the previous one, which was 1,176 Figure 1 Comparison between the number of species and the number of endemic species of lycophytes and ferns between the 2010 and the 2015 versions of the Brazilian List. Rodriguésia 66(4):

145 1076 Prado, J. et al. species (Prado & Sylvestre 2010) (Fig. 1). Due to recent advances in the classifications of groups (e.g., Moran et al and Øllgaard 2012a, and others), 36 families and 133 genera are currently recognized, as opposed to 33 families and 121 genera listed in the 2010 Catalogue. The ten most diverse families are: Pteridaceae, with 196 species, followed by Dryopteridaceae (179), Polypodiaceae (164), Hymenophyllaceae (90), Thelypteridaceae (86), Aspleniaceae (78), Lycopodiaceae (64), Selaginellaceae (55) Anemiaceae (51), and Cyatheaceae (45) (Fig. 2). Based on these results, further changes in the 2010 data include: Hymenophyllaceae now exceed Thelypteridaceae in number of species; Lycopodiaceae exceeds Anemiaceae and Selaginellaceae, while the number of species of Anemiaceae has been reduced with relation to 2010 (Fig. 2). The 36 other families occurring in Brazil are represented by numbers ranging from 32 to one species. With respect to the information published by Prado & Sylvestre (2010), the percentage of endemic species had a small decrease, from 38.2% (450 endemic species of 1,176 species) in 2010 to 36.7% (460 of 1,253) in this compilation. However, among the ten families with the highest number of endemic taxa, Dryopteridaceae remained the richest, with 89 endemic species, followed by Polypodiaceae (72), Pteridaceae (71), Thelypteridaceae (38), Lycopodiaceae (31), Anemiaceae (30), Aspleniaceae (23), Selaginellaceae (15), and Cyatheaceae (14) (Fig. 2). There were also changes to the 2010 data regarding the Lycopodiaceae, which surpassed Anemiaceae, Selaginellaceae, and Cyatheaceae; and in Hymenophyllaceae where there was a reduction in the number of endemic taxa. On the other hand, Isoetaceae and Athyriaceae (with 18 and 11 species, respectively), which are not among the ten richest families in Brazilian flora, have a high number of endemic species (Tab. 1, genera Isoetes and Diplazium). The other families have numbers ranging from less than 10 endemic species to none. As mentioned earlier, 133 genera have been recognized in this study, with the 20 genera most diverse encompassing 804 species, 306 of them endemic (Tab. 1). As in the previous study, the three most diverse genera remain Elaphoglossum (87 species), Thelypteris (85), and Asplenium Figure 2 The ten most diverse families of ferns and lycophytes (total number of species and endemic species) in 2010 and Rodriguésia 66(4):

146 Diversity of ferns and lycophytes in Brazil Table 1 Ferns and lycophytes of the twenty richest genera. Each genus is cited with their number of species and number of endemic species in the first version of the Species List of Flora of Brazil (Prado & Sylvestre 2010) and the current list (2015). Genera N. of species 2015 N. of endemic 2015 N. of species 2010 N. of endemic 2010 Elaphoglossum Thelypteris Asplenium Adiantum Selaginella Anemia Phlegmariurus * 25* Cyathea Trichomanes Doryopteris Hymenophyllum Lindsaea Blechnum Diplazium Pteris Isoetes Campyloneurum Pecluma Megalastrum Microgramma Total *as Huperzia 1077 (74). However, it is noteworthy that the number of species in Elaphoglossum had the highest increase between the first and the second study (from 78 species in 2010 to 87 in 2015), representing an increment of 11.5%. Allied to this increase, the number of endemic species in this genus also increased from 35 to 41. Other numerical variations that stood out occurred in Anemia, which had 69 species (40 endemic) in 2010 and was reduced to 51 (30 endemic) in 2015; in Cyathea, which jumped from 30 species (19 endemic) to 39 (13 endemic); Doryopteris, previously with 27 species (16 endemic), has now 32 (20 endemic), and Ctenitis, which was among the most diverse genera in 2010 (with 17 species, 10 endemic), and was surpassed by Isoetes, now with 23 species (18 endemic). Due to changes in the classification of Lycopodiaceae, most of the species of the genus Huperzia occurring in Brazil and South America were transferred to Phlegmariurus. Currently, Huperzia is represented in Brazil by a single endemic species, Huperzia catharinae (Christ) Holub, while in 2010 this genus comprised 38 species, 25 endemic to Brazil. In the current compilation, the species previously mentioned in Huperzia appear in the genus Phlegmariurus (40 species, 23 endemic) (Tab. 1). The Brazilian biome with the highest number of species continues to be the Atlantic Rainforest, with 883 species, followed by the Amazon Rainforest (503 species), Cerrado (269), Pantanal (30), Caatinga (26), and Pampa (eight) (Fig. 3). It should be noted that in the 2010 the Caatinga had exceeded the Pantanal (25 and 18 species, respectively). The Atlantic Rainforest is the area with the largest number of endemic Rodriguésia 66(4):

147 1078 Prado, J. et al. species, followed by the Cerrado, the latter with a concentration of endemism associated to the Campo rupestre vegetation type. The greatest diversity in the Atlantic Rainforest is found in the Ombrophilous Forest, and this type of vegetation concentrates 90% of species and 58% of the endemic species found in this area. Although the Mixed Ombrophilous Forest and the Seasonally Semideciduous Forest exhibit higher species numbers than the Altitude Grassland, the latter type of vegetation accounts for a greater percentage of endemism (9%). In the Amazon Rainforest, the greatest diversity is found in the Terra Firme Forest (313 species), with six endemic species. The Cerrado latu sensu is the vegetation type with the largest number of endemic species in the Cerrado (16 species), followed by the Campo rupestre, where eight endemic species are recognized. Regarding the geopolitical regions, there was an evident increase in the number of species and the Southeast region was confirmed as the most diverse, with 841 species in this survey (vs. 789 in 2010), as well as with the higher number of endemic species (131). The other regions also increased their representation: the South showed 576 species (vs. 540 species in 2010); North with 547 species (vs. 441); Northeast 502 (vs. 385), and Central-west with 394 (vs. 245) (Fig. 4). In relation to the diversity by state, there were also some changes compared to previous data. However, Minas Gerais remained the state with the largest number of species, (657 species vs. 580, in the previous survey). The states of Rio de Janeiro, currently with 620 species (vs. 552 species in 2010) and São Paulo, with 618 species (vs. 561), corroborate the greatest amount of species in the Southeast. Compared to the previous data, the largest additions occurred in the states of Bahia (138 species) and Espírito Santo (108), probably due to increased knowledge of the flora of these states, which admittedly are among the ten richest in the country. The fewer inclusions correspond to the northeastern states of Sergipe (19 species), Piauí (17), and Rio Grande do Norte (12), the latter state with the lowest diversity, adding up to only 20 species. Nonetheless, the number of records for these states increased very significantly, especially for Tocantins (210%), Sergipe (190%), and Rio Grande do Norte (150%) (Tab. 2). Most species occurring in Brazil are herbaceous and terrestrial (684 species), followed by epiphytes (344), rupicolous (105), aquatics Figure 3 Number of species of ferns and lycophytes by biome recorded in 2010, 2015, and number of endemic species per biome in Figure 4 Number of species of ferns and lycophytes by geographic region recorded in 2010, 2015 and number of endemic species per geographic region in Figure 5 Number of species and endemic species of ferns and lycophytes by life form and substrate (data from 2015). Rodriguésia 66(4):

148 Diversity of ferns and lycophytes in Brazil 1079 Table 2 Species of ferns and lycophytes in the states of Brazil. Number of species, number of endemic species, and percentage change per state (and Federal District) in the first version of the Species List flora of Brazil (Prado & Sylvestre 2010) and current list (2015). The species endemic occur exclusively in the cited state. States + Federal District N. of species 2015 N. of endemic 2015 N. of species 2010 N. of endemic 2010 % change Minas Gerais Rio de Janeiro São Paulo Paraná Santa Catarina Bahia Espírito Santo Amazonas Rio Grande do Sul Mato Grosso Pará Goiás Acre Pernambuco Mato Grosso do Sul Roraima Rondônia Distrito Federal Ceará Alagoas Amapá Maranhão Tocantins Paraíba Piauí Sergipe Rio Grande do Norte (44), tree ferns (41), hemiepiphytes (31), and climbers (4) (Fig. 5). The arborescent habit is a hallmark in Cyatheaceae, but there are at least three species in this family that are small herbs: Hymenophyllopsis ctenitoides Lellinger, H. hymenophylloides L.D. Gómez, and Cyathea myriotricha (Baker) R.C. Moran & J. Prado. The climbing species belong to Salpichlaena (Blechnaceae) and Lygodium (Lygodiaceae), while the epiphytic species are found mainly in the families Aspleniaceae (Asplenium), Dryopteridaceae ( Elaphoglossum), Hymenophyllaceae (Didymoglossum, Hymenophyllum, Polyphlebium), Lomariopsidaceae (Nephrolepis), Lycopodiaceae (Phlegmariurus), and Polypodiaceae (in almost all genera). The hemiepiphyte species are distributed in many families, especially in the genera Mickelia and Polybotrya (Dryopteridaceae), Lomariopsis (Lomariopsidaceae), and Vandenboschia (Hymenophyllaceae). Finally, aquatic plants are Rodriguésia 66(4):

149 1080 Prado, J. et al. found mainly in the families Isoetaceae (Isoetes), Marsileaceae (Marsilea), Salviniaceae (Azolla and Salvinia), and Pteridaceae (Ceratopteris). Considering the three areas with the highest number of ferns and lycophytes, the Atlantic Rainforest is the one with the greatest diversity of life forms, followed by Amazon Rainforest, and the Cerrado, while the terrestrial herbs are most commonly found in Caatinga, Pantanal, and Pampa (Fig. 6). Most species threatened with extinction recorded in the Brazilian Red Book (Martinelli & Moraes 2013) present restricted occurrence to a single unit of the federation. Many of these species are represented by populations distributed in areas smaller than 20 km 2, and are here called restricted endemic (Fig. 7). The genera that have more than ten endemic species (Tab. 1) are also the most cited in the red book. However, no species of Cyathea and Diplazium were considered endangered, although many of them are endemic (13 and 11 species, respectively) and Cyatheaceae particularly is under pressure from the horticultural industry, and is also listed under CITES regulation worldwide. Isoetes, with 18 endemic species, is represented by a single endangered species in the Red List, indicating that more efforts may be necessary to understand these plants and their conservation requirements. As the process of evaluation of threatened species is continuous, there is an opportunity to improve the data for many families currently underestimated in the Red List. The endemic species of the Brazilian oceanic islands have a high degree of threat. Of these, we can highlight those occurring in the Trindade Island. Many of them are listed in the Brazilian Red Book (Martinelli & Moraes 2013), such as: Asplenium beckeri Brade, A. trindadense (Brade) Sylvestre, Figure 6 Representation of life forms and substrate in the three areas with the highest number of species of ferns and lycophytes (data from 2015). Figure 7 Number of species in each of the three categories of extinction threats (Vulnerable, Endangered, and Critically Endangered) appearing in the Brazilian Red Book (Martinelli & Moraes 2013), by index of endemism. Brazilian endemic species: Brazilian endemic species that occur in more than one geographical region; Regional endemic species: species with distribution restricted to a particular geographic region; Endemic restricted: species with distribution restricted to a particular unit of the Federation. Rodriguésia 66(4):

150 Diversity of ferns and lycophytes in Brazil Elaphoglossum beckeri Brade, Pecluma insularis (Brade) Salino, Pleopeltis trindadense (Brade) Salino, and Thelypteris noveana (Brade) Ponce. Discussion The recognition of 1,253 species of ferns and lycophytes in the Brazilian List of Plants and Fungi in 2015 represents an advance in the knowledge of the group and is probably more compatible to the actual values for its diversity. This number of species is closer to what was previously estimated by Prado (1998), who mentioned the existence of 1,200 1,300 species of both groups in the country, while the first survey conducted in 2010 recorded 1,176 species. The current number of species recognized in 2015 is 6.5% higher than the one recorded in The increase or decrease of species number in the current list can be attributed to recent studies that involved synopsis and taxonomic revisions. For example, in Anemia, 69 species were recognized in 2010, whereas now only 51 were accepted by Mickel (2015, in press). On the other hand, there are groups still lacking detailed studies, not only to improve species circumscription, but also to understand their geographic distribution. The main groups that lack studies are: Aspleniaceae, Blechnaceae, Cyatheaceae, Dryopteridaceae (especially Ctenitis and Elaphoglossum), Hymenophyllaceae, Isoetaceae, Lycopodiaceae (especially Phlegmariurus), Marattiaceae, Polypodiaceae (especially Campyloneurum, Microgramma, Pecluma, and Pleopeltis), Pteridaceae (mainly Adiantum), Selaginellaceae, Tectariaceae, and Thelypteridaceae (whose species were recognized in genera that have recently been segregated from Thelypteris). For these groups, no recent treatments with identification keys, descriptions, and illustrations for the taxa occurring in Brazil exist. Some of these genera are being studied, however the data are not yet available and it is likely that the number of species comprised by these groups will change as these studies are completed. The Brazilian Atlantic Rainforest was the area with the highest species, endemic and endangered counts. This region has being recognized as one of the biodiversity hotspots of the planet (Mittermeier et al. 1998, 2004), and likewise it had already been identified as a major primary centre of diversity and endemism for Neotropical ferns (Tryon 1972). This high diversity is attributed to different 1081 environmental conditions occurring in tropical mountainous areas, favouring the establishment of microenvironments with different characteristics and consequently an increase in floristic diversity (Moran 2008). Kessler (2010) stresses that the great diversity of ferns and lycophytes found in tropical rain forests is due especially to the high rates of evapotranspiration, associated with heavy rainfall and topographic complexity of the terrain. Conclusions It is believed that the information presented in the current list of ferns and lycophytes of Brazil is more accurate than those presented in However, we still need to employ more efforts to collect data in remote areas, thus improving our knowledge of the species and their distribution in Brazil, enabling us to better understand the threat they may face. References Baker, G Cyatheaceae et Polypodiaceae. In: C.F.P. Martius & A.G. Eichler (eds.). Flora Brasiliensis. Fleischer, Leipzig. Vol. 1, part 2, pp BFG Growing knowledge: an overview of Seed Plant diversity in Brazil. Rodriguésia 66: Biral, L. & Prado, J First record of Pellaea ovata (Pteridaceae) from Brazil. American Fern Journal 102: Carvalho, F.A.; Salino, A. & Zartman, C.E New country and regional records from the Brazilian side of Neblina Massif. American Fern Journal 102: Christenhusz, M.J.M.; Zhang, X.-C. & Schneider, H A linear sequence of extant families and genera of lycophytes and ferns. Phytotaxa 19: Cochran, A.T.; Prado, J. & Schuettpelz, E Tryonia, a new taenitidoid fern genus segregated from Jamesonia and Eriosorus (Pteridaceae). PhytoKeys 35: Dittrich, V.A.O.; Salino, A. & Almeida, T.E Two new species of the fern genus Blechnum with partially anastomosing veins from Northern Brazil. Systematic Botany 37: Forzza, R.C.; Baumgratz, J.F.A.; Bicudo, C.E. de M.; Canhos, D.; Carvalho Jr., A.A.; Costa, A.F.; Costa, D.P.; Hopkins, M.; Leitman, P.M.; Lohmann, L.G.; Lughadha, E.N.; Maia, L.C.; Martinelli, G.; Menezes, M.; Morim, M.P.; Nadruz, M.; Peixoto, A.L.; Pirani, J.R.; Prado, J.; Queiroz, L.P.; Souza, V.C.; Stehmann, J.R.; Sylvestre, L.S.; Walter, B.M.T. & Zappi, D.C. (orgs.) Catálogo de plantas e fungos do Brasil. Vol. 1. Jardim Botânico do Rio de Janeiro, Rio de Janeiro. 870p. Rodriguésia 66(4):

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152 Diversity of ferns and lycophytes in Brazil Martinelli, G.; Menezes, M.; Morim, M.P.; Nadruz, M.; Peixoto, A.L.; Pirani, J.R.; Prado, J.; Queiroz, L.P.; Souza, V.C.; Stehmann, J.R.; Sylvestre, L.S.; Walter, B M.T. & Zappi, D.C. (orgs.). Catálogo de plantas e fungos do Brasil. Vol. 1. Jardim Botânico do Rio de Janeiro, Rio de Janeiro. Pp Prado, J.; Hirai, R.Y. & Smith, A.R Dryopteris huberi (Dryopteridaceae), an overlooked species, and a key for the species of Dryopteris in Brazil. Brittonia 66: Salino, A.; Fernandes, R.S. & Pietrobom, M.R Thelypteris amazonica sp. nov. (Thelypteridaceae). Nordic Journal of Botany 29: 1-4. Salino, A.; Sousa, M.G.M. & Arruda, A.J Thelypteris indusiata (Thelypteridaceae), a new fern species from Amazonian Brazil. Phytotaxa 156: Schuettpelz, E.; Davila, A.; Prado, J.; Hirai, R.Y. & Yatskievych, G Molecular phylogenetic and morphological affinities of Adiantum senae (Pteridaceae). Taxon 63: Schwartsburd, P.B The identity of Polypodium gyroflexum (= Pleopeltis gyroflexa, comb. nov. - Polypodiaceae). American Fern Journal 104: Schwartsburd, P.B. & Prado, J Subspecies of Hypolepis rugosula (Dennstaedtiaceae; Pteridophyta) around the world: Morphological and biogeographic perspectives. Acta Botanica Brasilica 28: Schwartsburd, P.B. & Smith, A.R Novelties in Serpocaulon (Polypodiaceae). Journal of the Botanical Research Institute of Texas 7: Schwartsburd, P.B.; Boudrie, M. & Cremers, G Two new species of Hypolepis (Dennstaedtiaceae: Pteridophyta) from Mount Roraima region (N South America) and a revised key for the Guianan species. Fern Gazette 19: 1-9. Schwartsburd, P.B.; Moraes, P.L. & Lopes-Mattos, K.L.B Recognition of two morpho-types in eastern South American brackens (Pteridium- Dennstaedtiaceae-Polypodiopsida). Phytotaxa 170: Spring, A.F Lycopodineae. In: Martius, C.F.P. & Eichler, A.G. (eds.). Flora Brasiliensis. Fleischer, Leipzig.Vol. 1, part 2, pp Sturm, J.G Ophioglossaceae, Marattiaceae, Osmundaceae, Schizaeceae, Gleicheniaceae et Hymenophyllaceae. In: Martius, C.F.P. & Eichler, A.G.(eds.). Flora Brasiliensis. Fleischer, Leipzig. Vol. 1, part 2, pp Sundue, M.A.; Hirai, R.Y. & Prado, J Rumohra glandulosissima (Dryopteridaceae) a new species from the Atlantic Rainforest, and revision of the species occurring in Brazil. Systematic Botany 38: Tryon, R.M Endemic areas and speciation in tropical American ferns. Biotropica 4: Vasco, A Taxonomic revision of Elaphoglossum subsection Muscosa (Dryopteridaceae). Blumea 56: Vasco, A.; Mickel, J. T. & Moran, R.M Taxonomic revision of the neotropical species of Elaphoglossum sect. Squamipedia (Dryopteridaceae). Annals of the Missouri Botanical Garden 99: Artigo recebido em 06/04/2015. Aceito para publicação em 31/08/2015. Rodriguésia 66(4):

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154 Rodriguésia 66(4): DOI: / Growing knowledge: an overview of Seed Plant diversity in Brazil The Brazil Flora Group Recommended citation: BFG (2015). This paper was compiled by Daniela C. Zappi 1, Fabiana L. Ranzato Filardi, Paula Leitman, Vinícius C. Souza, Bruno M.T. Walter, José R. Pirani, Marli P. Morim, Luciano P. Queiroz, Taciana B. Cavalcanti, Vidal F. Mansano and Rafaela C. Forzza 1 With contributions by: Abreu, Maria C.; Acevedo-Rodríguez, Pedro; Agra, Maria F.; Almeida Jr., Eduardo B.; Almeida, Gracineide S.S.; Almeida, Rafael F.; Alves, Flávio M.; Alves, Marccus; Alves-Araujo, Anderson; Amaral, Maria C.E.; Amorim, André M.; Amorim, Bruno; Andrade, Ivanilza M.; Andreata, Regina H.P.; Andrino, Caroline O.; Anunciação, Elisete A.; Aona, Lidyanne Y.S.; Aranguren, Yani; Aranha Filho, João L.M.; Araújo, Andrea O.; Araújo, Ariclenes A.M.; Araújo, Diogo; Arbo, María M.; Assis, Leandro; Assis, Marta C.; Assunção, Vivian A.; Athiê- Souza, Sarah M.; Azevedo, Cecilia O.; Baitello, João B.; Barberena, Felipe F.V.A.; Barbosa, Maria R.V.; Barros, Fábio; Barros, Lucas A.V.; Barros, Michel J.F.; Baumgratz, José F.A.; Bernacci, Luis C.; Berry, Paul E.; Bigio, Narcísio C.; Biral, Leonardo; Bittrich, Volker; Borges, Rafael A.X.; Bortoluzzi, Roseli L.C.; Bove, Cláudia P.; Bovini, Massimo G.; Braga, João M.A.; Braz, Denise M.; Bringel Jr., João B.A.; Bruniera, Carla P.; Buturi, Camila V.; Cabral, Elza; Cabral, Fernanda N.; Caddah, Mayara K.; Caires, Claudenir S.; Calazans, Luana S.B.; Calió, Maria F.; Camargo, Rodrigo A.; Campbell, Lisa; Canto-Dorow, Thais S.; Carauta, Jorge P.P. ; Cardiel, José M.; Cardoso, Domingos B.O.S.; Cardoso, Leandro J.T.; Carneiro, Camila R.; Carneiro, Cláudia E.; Carneiro-Torres, Daniela S.; Carrijo, Tatiana T.; Caruzo, Maria B.R.; Carvalho, Maria L.S.; Carvalho-Silva, Micheline; Castello, Ana C.D.; Cavalheiro, Larissa; Cervi, Armando C. ; Chacon, Roberta G.; Chautems, Alain; Chiavegatto, Berenice; Chukr, Nádia S.; Coelho, Alexa A.O.P.; Coelho, Marcus A.N.; Coelho, Rubens L.G.; Cordeiro, Inês; Cordula, Elizabeth; Cornejo, Xavier; Côrtes, Ana L.A.; Costa, Andrea F.; Costa, Fabiane N.; Costa, Jorge A.S.; Costa, Leila C.; Costa-e-Silva, Maria B.; Costa-Lima, James L.; Cota, Maria R.C.; Couto, Ricardo S.; Daly, Douglas C.; De Stefano, Rodrigo D.; De Toni, Karen; Dematteis, Massimiliano; Dettke, Greta A.; Di Maio, Fernando R.; Dórea, Marcos C.; Duarte, Marília C.; Dutilh, Julie H.A.; Dutra, Valquíria F.; Echternacht, Lívia; Eggers, Lilian; Esteves, Gerleni; Ezcurra, Cecilia; Falcão Junior, Marcus J.A.; Feres, Fabíola; Fernandes, José M.; Ferreira, D.M.C.; Ferreira, Fabrício M.; Ferreira, Gabriel E.; Ferreira, Priscila P.A.; Ferreira, Silvana C.; Ferrucci, Maria S.; Fiaschi, Pedro; Filgueiras, Tarciso S.; Firens, Marcela; Flores, Andreia S.; Forero, Enrique; Forster, Wellington; Fortuna-Perez, Ana P.; Fortunato, Reneé H.; Fraga, Cláudio N.; França, Flávio; Francener, Augusto; Freitas, Joelcio; Freitas, Maria F.; Fritsch, Peter W.; Furtado, Samyra G.; Gaglioti, André L.; Garcia, Flávia C.P.; Germano Filho, Pedro; Giacomin, Leandro; Gil, André S.B.; Giulietti, Ana M.; Godoy, Silvana A.P. ; Goldenberg, Renato; Gomes da Costa, Géssica A.; Gomes, Mário; Gomes-Klein, Vera L.; Gonçalves, Eduardo Gomes; Graham, Shirley; Groppo, Milton; Guedes. Juliana S.; Guimarães, Leonardo R.S.; Guimarães, Paulo J.F.; Guimarães, Elsie F.; Gutierrez, Raul; Harley, Raymond; Hassemer, Gustavo; Hattori, Eric K.O.; Hefler, Sonia M.; Heiden, Gustavo; Henderson, Andrew; Hensold, Nancy; Hiepko, Paul; Holanda, Ana S.S.; Iganci, João R.V.; Imig, Daniela C.; Indriunas, Alexandre; Jacques, Eliane L.; Jardim, Jomar G.; Kamer, Hiltje M.; Kameyama, Cíntia; Kinoshita, Luiza S.; Kirizawa, Mizué; Klitgaard, Bente B.; Koch, Ingrid; Koschnitzke, Cristiana; Krauss, Nathália P.; Kriebel, Ricardo; Kuntz, Juliana; Larocca, João; Leal, Eduardo S.; Lewis, Gwilym P.; Lima, Carla

155 1086 The Brazil Flora Group T.; Lima, Haroldo C.; Lima, Itamar B.; Lima, Laíce F.G.; Lima, Laura C.P.; Lima, Leticia R.; Lima, Luís F.P.; Lima, Rita B.; Lírio, Elton J.; Liro, Renata M.; Lleras, Eduardo ; Lobão, Adriana; Loeuille, Benoit; Lohmann, Lúcia G.; Loiola, Maria I.B.; Lombardi, Julio A.; Longhi-Wagner, Hilda M.; Lopes, Rosana C.; Lorencini, Tiago S.; Louzada, Rafael B.; Lovo, Juliana; Lozano, Eduardo D.; Lucas, Eve; Ludtke, Raquel; Luz, Christian L.; Maas, Paul; Machado, Anderson F.P.; Macias, Leila; Maciel, Jefferson R.; Magenta, Mara A.G.; Mamede, Maria C.H.; Manoel, Evelin A.; Marchioretto, Maria S.; Marques, Juliana S.; Marquete, Nilda; Marquete, Ronaldo; Martinelli, Gustavo; Martins da Silva, Regina C.V.; Martins, Ângela B.; Martins, Erika R.; Martins, Márcio L.L.; Martins, Milena V.; Martins, Renata C.; Matias, Ligia Q.; Maya-L., Carlos A.; Mayo, Simon; Mazine, Fiorella; Medeiros, Debora; Medeiros, Erika S.; Medeiros, Herison; Medeiros, João D.; Meireles, José E.; Mello-Silva, Renato; Melo, Aline; Melo, André L.; Melo, Efigênia; Melo, José I.M.; Menezes, Cristine G.; Menini Neto, Luiz; Mentz, Lilian A.; Mezzonato, A.C.; Michelangeli, Fabián A.; Milward-de-Azevedo, Michaele A.; Miotto, Silvia T.S.; Miranda, Vitor F.O.; Mondin, Cláudio A.; Monge, Marcelo; Monteiro, Daniele; Monteiro, Raquel F.; Moraes, Marta D.; Moraes, Pedro L.R.; Mori, Scott A.; Mota, Aline C.; Mota, Nara F.O.; Moura, Tania M.; Mulgura, Maria; Nakajima, Jimi N.; Nardy, Camila; Nascimento Júnior, José E.; Noblick, Larry; Nunes, Teonildes S.; O Leary, Nataly; Oliveira, Arline S.; Oliveira, Caetano T.; Oliveira, Juliana A.; Oliveira, Luciana S.D.; Oliveira, Maria L.A.A.; Oliveira, Regina C.; Oliveira, Renata S.; Oliveira, Reyjane P.; Paixão-Souza, Bruno; Parra, Lara R.; Pasini, Eduardo; Pastore, José F.B.; Pastore, Mayara; Paula-Souza, Juliana; Pederneiras, Leandro C.; Peixoto, Ariane L.; Pelissari, Gisela; Pellegrini, Marco O.O.; Pennington, Toby; Perdiz, Ricardo O.; Pereira, Anna C.M.; Pereira, Maria S.; Pereira, Rodrigo A.S.; Pessoa, Clenia; Pessoa, Edlley M.; Pessoa, Maria C.R.; Pinto, Luiz J.S.; Pinto, Rafael B.; Pontes, Tiago A.; Prance, Ghillean T.; Proença, Carolyn; Profice, Sheila R.; Pscheidt, Allan C.; Queiroz, George A.; Queiroz, Rubens T.; Quinet, Alexandre; Rainer, Heimo; Ramos, Eliana; Rando, Juliana G.; Rapini, Alessandro; Reginato, Marcelo; Reis, Ilka P.; Reis, Priscila A.; Ribeiro, André R.O.; Ribeiro, José E.L.S.; Riina, Ricarda; Ritter, Mara R.; Rivadavia, Fernando; Rocha, Antônio E.S.; Rocha, Maria J.R.; Rodrigues, Izabella M.C.; Rodrigues, Karina F.; Rodrigues, Rodrigo S.; Rodrigues, Rodrigo S.; Rodrigues, Vinícius T.; Rodrigues, William; Romaniuc Neto, Sérgio; Romão, Gerson O.; Romero, Rosana; Roque, Nádia; Rosa, Patrícia; Rossi, Lúcia; Sá, Cyl F.C.; Saavedra, Mariana M.; Saka, Mariana; Sakuragui, Cássia M.; Salas, Roberto M.; Sales, Margareth F.; Salimena, Fatima R.G.; Sampaio, Daniela; Sancho, Gisela; Sano, Paulo T.; Santos, Alessandra; Santos, Élide P.; Santos, Juliana S.; Santos, Marianna R.; Santos-Gonçalves, Ana P.; Santos-Silva, Fernanda; São-Mateus, Wallace; Saraiva, Deisy P.; Saridakis, Dennis P.; Sartori, Ângela L.B.; Scalon, Viviane R.; Schneider, Ângelo; Sebastiani, Renata; Secco, Ricardo S.; Senna, Luisa; Senna-Valle, Luci; Shirasuna, Regina T.; Silva Filho, Pedro J.S.; Silva, Anádria S.; Silva, Christian; Silva, Genilson A.R.; Silva, Gisele O.; Silva, Márcia C.R.; Silva, Marcos J.; Silva, Marcos J.; Silva, Otávio L.M.; Silva, Rafaela A.P.; Silva, Saura R.; Silva, Tania R.S.; Silva-Gonçalves, Kelly C.; Silva-Luz, Cíntia L.; Simão-Bianchini, Rosângela; Simões, André O.; Simpson, Beryl; Siniscalchi, Carolina M.; Siqueira Filho, José A.; Siqueira, Carlos E.; Siqueira, Josafá C.; Smith, Nathan P.; Snak, Cristiane; Soares Neto, Raimundo L.; Soares, Kelen P.; Soares, Marcos V.B.; Soares, Maria L.; Soares, Polyana N.; Sobral, Marcos; Sodré, Rodolfo C.; Somner, Genise V.; Sothers, Cynthia A.; Sousa, Danilo J.L.; Souza, Elnatan B.; Souza, Élvia R.; Souza, Marcelo; Souza, Maria L.D.R.; Souza-Buturi, Fátima O.; Spina, Andréa P.; Stapf, María N.S.; Stefano, Marina V.; Stehmann, João R.; Steinmann, Victor; Takeuchi, Cátia; Taylor, Charlotte M.; Taylor, Nigel P.; Teles, Aristônio M.; Temponi, Lívia G.; Terra-Araujo, Mário H.; Thode, Veronica; Thomas, W.Wayt; Tissot-Squalli, Mara L.; Torke, Benjamin M.; Torres, Roseli B.; Tozzi, Ana M.G.A.; Trad, Rafaela J.; Trevisan, Rafael; Trovó, Marcelo; Valls, José F.M.; Vaz, Rodriguésia 66(4):

156 Seed Plant diversity in Brazil. Angela M.S.F.; Versieux, Leonardo; Viana, Pedro L.; Vianna Filho, Marcelo D.M.; Vieira, Ana O.S.; Vieira, Diego D.; Vignoli-Silva, Márcia; Vilar, Thaisa; Vinhos, Franklin; Wallnöfer, Bruno; Wanderley, Maria G.L.; Wasshausen, Dieter; Watanabe, Maurício T.C.; Weigend, Maximilian; Welker, Cassiano A.D.; Woodgyer, Elizabeth; Xifreda, Cecilia C.; Yamamoto, Kikyo; Zanin, Ana; Zenni, Rafael D.; Zickel, Carmem S Abstract An updated inventory of Brazilian seed plants is presented and offers important insights into the country s biodiversity. This work started in 2010, with the publication of the Plants and Fungi Catalogue, and has been updated since by more than 430 specialists working online. Brazil is home to 32,086 native Angiosperms and 23 native Gymnosperms, showing an increase of 3% in its species richness in relation to The Amazon Rainforest is the richest Brazilian biome for Gymnosperms, while the Atlantic Rainforest is the richest one for Angiosperms. There was a considerable increment in the number of species and endemism rates for biomes, except for the Amazon that showed a decrease of 2.5% of recorded endemics. However, well over half of Brazillian seed plant species (57.4%) is endemic to this territory. The proportion of life-forms varies among different biomes: trees are more expressive in the Amazon and Atlantic Rainforest biomes while herbs predominate in the Pampa, and lianas are more expressive in the Amazon, Atlantic Rainforest, and Pantanal. This compilation serves not only to quantify Brazilian biodiversity, but also to highlight areas where there information is lacking and to provide a framework for the challenge faced in conserving Brazil s unique and diverse flora. Key words: Angiosperms, biomes, endemism, Gymnosperms, life-forms. Resumo Um levantamento atualizado das plantas com sementes e análises relevantes acerca desta biodiversidade são apresentados. Este trabalho se iniciou em 2010 com a publicação do Catálogo de Plantas e Fungos e, desde então vem sendo atualizado por mais de 430 especialistas trabalhando online. O Brasil abriga atualmente espécies nativas de Angiospermas e 23 espécies nativas de Gimnospermas e estes novos dados mostram um aumento de 3% da riqueza em relação a A Amazônia é o Domínio Fitogeográfico com o maior número de espécies de Gimnospermas, enquanto que a Floresta Atlântica possui a maior riqueza de Angiospermas. Houve um crescimento considerável no número de espécies e nas taxas de endemismo para a maioria dos Domínios (Caatinga, Cerrado, Floresta Atlântica, Pampa e Pantanal), com exceção da Amazônia que apresentou uma diminuição de 2,5% de endemicidade. Entretanto, a maior parte das plantas com sementes que ocorrem no Brasil (57,4%) é endêmica deste território. A proporção de formas de vida varia de acordo com os diferentes Domínios: árvores são mais expressivas na Amazônia e Floresta Atlântica do que nos outros biomas, ervas são dominantes no Pampa e as lianas apresentam riqueza expressiva na Amazônia, Floresta Atlântica e Pantanal. Este trabalho não só quantifica a biodiversidade brasileira, mas também indica as lacunas de conhecimento e o desafio a ser enfrentado para a conservação desta flora. Palavras-chave: Angiospermas, Domínios, endemismo, formas de vida, Gimnospermas. Introduction The Global Strategy for Plant Conservation (GSPC), adopted by the Conference of the Parties of the Convention s signatory countries on Biological Diversity (CBD), held at The Hague, Holland, in 2002, set 16 targets to be achieved globally by The first target, vital to the completion of all others, was the development of a functional, widely accessible list of all known plant species, as a first step towards a complete World Flora. In 2008, the Botanical Garden of Rio de Janeiro was designated by the Ministry of Environment to coordinate the preparation of the Species List of the Brazilian Flora. The Botanical Garden of Rio de Janeiro invited botanists from several Brazilian institutions to constitute the project s Steering Committee. For the complete list of authors, institutions and taxonomic groups see supplementary material < DOI: / Authors for correspondence: d.zappi@kew.org, rafaela@jbrj.gov.br Rodriguésia 66(4):

157 1088 The Brazil Flora Group Target 1 of the GSPC was reached by Brazil in 2010 with the launch of the first on-line version of the List of Flora of Brazil and the publication of Plants and Fungi Catalogue. Awareness of the threats faced by the world s biodiversity has risen considerably since the 1980s (Myers et al. 2000; Mittermeier et al. 2004). Brazil is at the center of this discussion, as it includes two of the 34 recognized global hotspots, housing a larger number of plant species than any other country in the world (Mittermeier et al. 2004; Forzza et al. 2012). With an economy strongly driven by large scale agriculture, mining of its natural resources, and growing urban population, large tracts of the country s natural reserves are being depleted, even before our knowledge can ascertain what is being lost. The Flora brasiliensis (Martius et al. 1833; Urban 1906) treatment listed 19,958 species of fungi, algae, mosses and vascular plants, including 18,857 seed plants. After that, botanical knowledge expanded inordinately as new species were described and names were synonymized through floristic and monographic accounts. There have been both conservative and generous estimates for the size of the seed plant component of the Brazilian Flora (Forzza et al. 2012), and, while the 2010 list, dealing only with described plant diversity, did not reach the more conservative published estimate (35,664 species) and is very far from the exaggerated (45,015 species) ones, it still surpassed the number of plants known thus far in any other country in the world. The Brazilian Catalogue of Plants and Fungi reported 31,188 species of seed plants, distributed in 2,818 genera and 227 families of Angiosperms (Forzza et al. 2010) and 23 species in six genera and five families of Gymnosperms (Souza 2010). Five years have elapsed from the first release of the Brazilian on-line list and the addition of new facilities have enabled botanists to contribute further information regarding species habit, substrate and vegetation types, as well as continuing to refine the existing list and adding new records and taxa. Here we provide a snapshot of the changes to the Brazilian List during this period, and also analyse new data that have come to light at the end of this process, providing a fresh insight into seed plant species numbers, endemism and distribution patterns, as well as proportions of life-forms and richness throughout the main habitats of this megadiverse country. Methods The methodology followed the principles established for all plant and fungi groups catalogued by Forzza et al. (2010). The minimum Species2000 fields were adopted and increased with others (geopolitical divisions, biomes etc.) that were interesting to feature for the Brazilian context, and the design and development of an information system that allowed instant management, collaborative updating of the data and generation of the 2010 Catalogue (Forzza et al. 2010). The work took place in two phases. The first one included analysis, harmonization and importation of diverse state and regional lists and taxonomic treatments already available (see Forzza et al for a complete list), while the second phase was the revision and updating of the information relative to each taxon made directly on-line by 334 specialists. From 2011 the contributors continued with the task of including new records and modifying existent ones as more scientific data became available. Yearly releases of the system comprised new fields and dictionaries to include data regarding life-forms, substrate and vegetation type for each taxon that needed to be completed, and a new functionality to include images meant that the researchers could, from that point onwards, choose images from herbarium records and make available their own images of living plants. At the end of 2014 the Angiosperm and Gymnosperm coordinators complemented data in families where data were still missing. During the three first months of 2015 a data evaluation process generated a series of reports indicating inconsistencies aimed to help the 437 contributors to further clean the data. The corrections were added in March For more information about the final dataset see supplementary material < - DOI: / From the start of the project, the importance to distinguish between native and exotic species was highlighted, and the contributing specialists had to complete an Origin field with the following options: native, naturalized and cultivated. Moro et al. (2012) clearly states different categories for exotic species and analyses the discrepancies of treatment these species receive from different taxonomists in Forzza et al. (2010). It was never the intention comprehensively list all species Rodriguésia 66(4):

158 Seed Plant diversity in Brazil. cultivated in Brazil. Moreover, for the purpose of the statistics presented in this paper, all analyses apart from the initial overview section were based only on the numbers of native plants. However, data from Forzza et al. (2010) may take into account some non-native species, influencing the comparisons performed within this article. The term endemic, meaning that a taxon is unique to a defined geographic location, has been used at different levels (endemic to Brazil, endemic to one of Brazil s Biomes, a state or a vegetation type). The words endemic and restricted, or with restricted distribution, were used to analyse taxa in relation to where they occur, for instance, Parodia buiningii (Buxb.) N.P.Taylor (Cactaceae) is restricted to the state of Rio Grande do Sul and to the Pampa in Brazil, but it also occurs in Argentina, therefore it is not endemic of Brazil. As one of the objectives of this work was to examine the effect of the continued input by specialists into the on-line Brazilian List system (Lista de Espécies da Flora do Brasil 2015), comparative gross percentage calculations highlighting growth and decrease in species number were calculated. These were based on the species number difference between 2015 and 2010 divided by the totals found in These are presented alongside the total percentages in the results tables. The term Biome used in the project and in the results and discussion of this paper are equivalent to the six Phytogeographic Domains, or Domínios Fitogeográficos defined by IBGE (2004). Results Overall More than 110,000 names were included in the system by 2015, while the original database in 2010 had 94,144 names (Tab. 1). All 69 names of Gymnosperms, as well as the further six names added by 2015, were checked (Tab. 1). The positive balance of new Angiosperm names included in the system in five years was 9,274 (11.6% of the initial total). The number of checked names, 85,400, grew in balance by 20,033, or 30.6% of the initial total of 65,369 in 2010, thus today the Brazilian list has over 95.5% of its Angiosperm names checked by specialists (Tab. 1). Wherever possible, the checked names either marked as accepted names or as synonyms linked to an accepted name (Tab. 2). The present number of unplaced names of Angiosperms is 633, and a single name for Gymnosperms (Tab. 2). At present, seed plant diversity totals 32,109 accepted native species (23 of which are Gymnosperms) 1089 belonging to 2,746 genera and 229 families (Tab. 2). With the addition of 921 native species between 2010 and 2015, the species number figure grew 3% in the last five years (Tab. 3). Overall endemism figures have grown proportionally more than the number of accepted native species, from 17,632 in 2010 (56.5%) to 18,423 (57.4%) in 2015, with the addition of 791 endemic species, a gross increase of 4.5% (Tab. 3). The Brazilian Red List (Martinelli & Moraes 2013) includes 1,974 species (1,772 endemic and 202 not endemic to Brazil) that were currently listed under one of these threat categories: critically endangered - CR, endangered - EN, and vulnerable - VU (Tab. 4). Top 10 families The top 10 families Fabaceae, Orchidaceae, Asteraceae, Rubiaceae, Melastomataceae, Bromeliaceae, Poaceae, Myrtaceae, Euphorbiaceae and Malvaceae contribute 15,404 to the total number of Angiosperm species (47.2%). The total species contribution of these families appears to be more significant in terms of endemism, contributing 9,593 endemic species, or 62.3% of the endemism at the country level (Tab. 5). The increase found for the total species number (4.3%) is lower than the endemism difference of 6.8% found for the top 10 families. Nine of the ten largest families coincide with those found in the Catalogue (Forzza et al. 2010), the exception being Apocynaceae (down 5 species from now with 754 species, of which 403 are endemic), that was surpassed by Malvaceae in 2015 (Tab. 5). Apart from Poaceae, where there was a decrease of 120 species, the general increase found was between 2.1% (Rubiaceae) and 21.5% (Euphorbiaceae) for the largest families. The remaining 214 families that occur in Brazil have between one and 756 species. In terms of endemism, the gross increase was between 1% (Orchidaceae) and 26.6% (Euphorbiaceae), but in general endemism grew between 2.2% (Asteraceae) and 8.7% (Poaceae), with Myrtaceae (12.7%) and Bromeliaceae (14%) reaching slightly higher values. Top 30 genera The 30 largest genera of Brazilian Angiosperms add up to 6,380 species, representing 19.5% of the total diversity of the group. The contribution of these genera appears to be proportionally more significant in terms of endemism, contributing with 4,485 endemic species, or 24.3% of total endemism (Tab. 6). The order in which the top 30 largest genera appeared in Forzza et al. (2010) has changed Rodriguésia 66(4):

159 1090 The Brazil Flora Group Table 1 Total number of names from the Brazilian List system, name totals for Angiosperms and Gymnosperms and proportion between the names analysed by specialists ( checked names ) in 2010 and *Data includes all specific and infraspecific names of native, naturalized and cultivated species of the algae, fungi, and plants that occur and that does not occur in Brazil, but are recorded by the system (< Group Name total 2015* Checked names 2015 Checked names % 2015 Name total 2010* Checked names 2010 Checked names %2010 Growth rate of total names % Growth rate for checked names % Plants & Fungi 110, , ,144 78, Angiosperms 89,443 85, ,169 65, Gymnosperms Table 2 Family, genus and species diversity for Angiosperms and Gymnosperms from the Brazilian List system The lower lines indicate native Angiosperms, Gymnosperms and total (excluding naturalized and cultivated taxa). Caption: #11 naturalized and one cultivated families. ##170 naturalized and 60 cultivated genera. ###543 naturalized and 203 cultivated species. *One naturalized family. **One naturalized genus (Pinus). ***Seven naturalized species. Accepted names, synonyms, and unplaced names were analysed by specialists ( checked names see Table 1). Group Families Genera Endemic genera Species Endemic species % Endemic species Accepted names Synonyms Unplaced name Angiosperms 236 # 2,970 ## ,832 ### 18,421-36,520 48, Gymnosperms 6 * 7 ** 0 30 *** Total 242 2, ,862 18,423-36,550 48, Native species 40.8% 53.9% 0.7% Angiosperms 224 2, ,086 18, Gymnosperms Total 229 2, ,109 18, Table 3 Total and endemic number of native and naturalized Angiosperms and Gymnosperms. Group Species total 2015 Endemic species 2015 % Endemic species 2015 Species total 2010 Endemic species 2010 % Endemic species 2010 Total species growth % Endemic species growth % Angiosperms 32,629 18, ,162 17, Gymnosperms Total 32,659 18, ,188 17, Rodriguésia 66(4):

160 Seed Plant diversity in Brazil Table 4 Angiosperms and Gymnosperms* species analysed according to threat categories (Vulnerable, Endangered and Critically endangered) according to the Brazilian Red List (Martinelli & Moraes 2013). Numbers distributed according to endemism**. Caption: *Araucaria angustifolia (Bertol.) Kuntze, not endemic, Endangered. **Endemic to Brazil: species endemic to Brazil but that are distributed in more than one region; Regional endemic: species endemic to Brazil and restricted to a single region; Local endemic: endemic to Brazil and restricted to a single state; Not endemic: species not endemic to Brazil and with wide distribution; Regionally restricted: species not endemic to Brazil but with Brazilian distribution restricted to a single region; Locally restricted: species not endemic to Brazil but found in a single Brazilian state. Threat categories Endemic to Brazil Regional endemic Local endemic Not endemic and with wide distribution Not endemic but regionally restricted Not endemic but locally restricted Vulnerable Endangered Critically endangered Total 1,772 1,410 1, Table 5 Ten top diverse Angiosperms families from Brazil, showing native species total and endemic. Families Species total 2015 Endemic species 2015 % Endemic species Rank Species total 2010 Endemic species 2010 % Endemic species 2010 Total species growth % Endemic species growth % Fabaceae 2,756 1, ,694 1, Orchidaceae 2,548 1, ,419 1, Asteraceae 2,013 1, ,966 1, Rubiaceae 1, , Melastomataceae 1, , Bromeliaceae 1,343 1, ,207 1, Poaceae 1, , Myrtaceae 1, Euphorbiaceae Malvaceae Total 15,404 9, ,774 8, Rodriguésia 66(4):

161 1092 The Brazil Flora Group noticeably, with Eugenia overtaking Paepalanthus and Mimosa, and Paepalanthus falling to the third position, Croton climbing from 16 th to 4 th position, Psychotria falling from 6 th to 10 th position, Leandra from 10 th to 14 th, Vriesea moving from 15 th to 11 th and Baccharis from 24 th to 17 th position. Dyckia, Pavonia and Anthurium reached the top 30 largest genera, while Inga, Justicia and, more dramatically, Hyptis, were overtaken. While Croton was the genus with the largest increase in number of species, from 186 to 316 species (Tab. 6), other three amongst the most speciose genera have increased by around 20% or more: Baccharis (from 149 to 178 species), Dyckia (from 115 to 141 species) and Myrcia (from 215 to 260 species). The genera that were adjusted downwards are Paepalanthus (from 357 to 338 species), Psychotria (264 to 252) and Leandra (213 to 206). The proportion of endemic species within these genera followed, in general, the increases or decreases in species number. However, Croton had a more dramatic endemism increase (86.7%) compared to species richness growth of 69.9% and, in Rhynchospora, endemism increased by 45.5% for a species growth of 8.1%. In relative terms, the percentage of endemic species of Rhynchospora moved from 24.3% of the species to 32.6%. Amongst the exceptions to this trend, Philodendron species increased 7.7% from 156 to 168 species, while the endemism figure was adjusted from 127 to only 79 species, falling 37.8%, from 81.4% to 47% endemic (Tab. 6). Despite the fact that the species numbers increased for Baccharis, the proportion of endemism has slightly decreased (-2.91%). The genus Piper, with a slight increase in the number of species, had a gross endemic species decrease of 6.3%. While the ten most diverse families presented a slight increase in the level of endemism when compared to the list produced in 2010 (Tab. 5), the 30 largest genera practically maintained their proportional contribution in percentages: 70.2% in 2010 and 70.3% in 2015 (Tab. 6). Biomes Seed plant occurrence in the Brazilian biomes is illustrated in Figure 3. While for the Gymnosperms (Tab. 7) the Amazon Rainforest is the Brazilian biome with the highest number of species, for Angiosperms (Tab. 8, Fig. 1) the highest species number is found in the Atlantic Rainforest, with 15,001 native species of which 7,432 (49.5%), are endemic to this biome, followed by the Cerrado, 12,097 total species with 4,252 (35.1%) endemic; Amazon Rainforest (11,896 total species, with 1,900 (16%) endemic); Caatinga (4,657 total species, with 913 (19.7%) endemic); Pampa (1,685 total species, with 102 (6.1%) endemic); and Pantanal (1,277 total species, with 54 (4.2%) endemic species). The increment of species between 2010 and 2015 was proportionally higher for the two biomes with least species (Fig. 1, Tab. 8), totalling 44.3% (392 species) for the Pantanal and 25.3% (340 species) for the Pampa. The Caatinga had an increase of 7.8% (337 species), which was a similar increase to the Atlantic Rainforest (7.4% or 1,029 species). The Cerrado gained an additional 712 species (6.3% increase), and the Amazon Rainforest, with 547 species added since 2010, had the smallest increase (4.8%). The figures for endemism per biome followed the growth trend for the Pantanal (8 more endemic species, or 17.4% increase), Pampa (26 more endemic species, 34.2% increase), and Caatinga (169 more endemic species, 22.7% increase). Meanwhile the Atlantic Rainforest and the Cerrado had moderate growth of endemism numbers, with respectively 418 and 102, an equivalent of a 6% and 2.5% increase. The Amazon Rainforest s recorded endemism decreased, with 48 less endemic species than it had in 2010, a decrease of 2.5%. In relative terms, from 2015, the endemism in the Amazon region represents 16.1% of the total species (Tab. 8). Each biome was analysed according to its top ten Angiosperm families (Tab. 9) and, of the overall top ten families (see item Figure 1 Angiosperm species number per biome, with total numbers of species in 2010 and 2015, and endemic species in Rodriguésia 66(4):

162 Seed Plant diversity in Brazil Table 6 Comparison between the 30 most diverse Angiosperm genera. Species Genera Families total 2015 Endemic species 2015 Endemic species % Rank Species total 2010 Endemic species 2010 Endemic species % 2010 Total species growth % Endemic species growth % Eugenia Myrtaceae Mimosa Fabaceae Paepalanthus Eriocaulaceae Croton Euphorbiaceae Piper Piperaceae Miconia Melastomataceae Solanum Solanaceae Myrcia Myrtaceae Chamaecrista Fabaceae Psychotria Rubiaceae Vriesea Bromeliaceae Begonia Begoniaceae Paspalum Poaceae Leandra Melastomataceae Mikania Asteraceae Aechmea Bromeliaceae Baccharis Asteraceae Xyris Xyridaceae Ocotea Lauraceae Philodendron Araceae Tibouchina Melastomataceae Peperomia Piperaceae Habenaria Orchidaceae Rhynchospora Cyperaceae Passiflora Passifloraceae Dyckia Bromeliaceae Dioscorea Dioscoreaceae Rodriguésia 66(4):

163 1094 The Brazil Flora Group Species Genera Families total 2015 Endemic species 2015 Endemic species % Rank Species total 2010 Endemic species 2010 Endemic species % 2010 Total species growth % Endemic species growth % Pavonia Malvaceae Epidendrum Orchidaceae Anthurium Araceae Total - 6,380 4, ,932 4, Table 7 Gymnosperm species number per biome. Biomes Species total 2015 Species total 2010 Amazon Rainforest Cerrado 6 6 Atlantic Rainforest 3 4 Caatinga 2 2 Pampa 2 1 Pantanal 0 0 Table 8 Angiosperm species numbers per biome. Biome Species total 2015 Endemic species 2015 Endemic species % 2015 Species total 2010 Endemic species 2010 Endemic species % 2010 Total species growth % Endemic species growth % Atlantic Rainforest 15,001 7, ,972 7, Cerrado 12,097 4, ,384 4, Amazon Rainforest 11,896 1, ,349 1, Caatinga 4, , Pampa 1, , Pantanal 1, Rodriguésia 66(4):

164 Seed Plant diversity in Brazil Table 9 Ten top Angiosperms families per biome. Atlantic Rainforest Cerrado Amazon Rainforest Caatinga Pampa Pantanal Orchidaceae (1,574) Asteraceae (1,216) Fabaceae (1,119) Fabaceae (605) Asteraceae (299) Poaceae (162) Fabaceae (964) Fabaceae (1,207) Orchidaceae (882) Poaceae (282) Poaceae (266) Fabaceae (153) Bromeliaceae (921) Orchidaceae (727) Rubiaceae (728) Asteraceae (284) Cyperaceae (141) Malvaceae (70) Asteraceae (885) Poaceae (648) Melastomataceae (495) Euphorbiaceae (232) Fabaceae (127) Asteraceae (67) Poaceae (734) Melastomataceae (484) Poaceae (440) Rubiaceae (168) Iridaceae (44) Cyperaceae (58) Myrtaceae (710) Eriocaulaceae (461) Apocynaceae (299) Cyperaceae (163) Solanaceae (44) Bignoniaceae(45) Melastomataceae (582) Rubiaceae (406) Cyperaceae (288) Malvaceae (156) Rubiaceae (42) Sapindaceae (41) Rubiaceae (564) Euphorbiaceae (386) Annonaceae (287) Apocynaceae (131) Convolvulaceae (36) Euphorbiaceae (35) Apocynaceae (367) Malvaceae (334) Euphorbiaceae (282) Melastomataceae (129) Cactaceae (35) Rubiaceae (34) Euphorbiaceae (327) Apocynaceae (293) Araceae (270) Orchidaceae (127) Amaryllidaceae (34) Malpighiaceae (25) Table 10 Angiosperm species numbers per Region with number of endemics. Region Species total 2015 Endemic species 2015 Endemic species % 2015 Species total 2010 North 12,414 1, ,127 Northeast 10,661 2, ,220 Central-Western 9,322 1, ,516 Southeast 16,127 5, ,550 South 7,566 1, ,333 Table 11 Gymnosperm species number per Region. Region Species total 2015 Species total 2010 North Northeast 2 2 Central-Western 7 7 Southeast 4 6 South 5 7 Rodriguésia 66(4):

165 1096 The Brazil Flora Group above), only Fabaceae, Poaceae and Rubiaceae appear among the top ten throughout all six biomes. Asteraceae and Euphorbiaceae appear in five biomes, while Apocynaceae, Cyperaceae, Melastomataceae and Orchidaceae appear in four. Malvaceae appears in three biomes. Thirteen plant families appear among the 10 top families under a single biome: Amaryllidaceae, Annonaceae, Araceae, Bignoniaceae, Bromeliaceae, Cactaceae, Convolvulaceae, Eriocaulaceae, Iridaceae, Malpighiaceae, Myrtaceae, Sapindaceae and Solanaceae. Fabaceae is the most species rich family in the Amazon Rainforest and in the Caatinga, while it occupies the second position in the Atlantic Rainforest, Cerrado and Pantanal, and the fourth in the Pampa (Tab. 9). Poaceae, also included among the overall ten most diverse families, is the first in the Pantanal, and the second in the Caatinga and in the Pampa. Rubiaceae occupies the fourth position in the overall ten top families, but does not reach the five first positions in four biomes, however it is the third richest family in the Amazon Rainforest and the fifth in the Caatinga. Asteraceae, amongst the top ten families in five biomes apart from the Amazon, is the most important family in both Cerrado and Pampa. Euphorbiaceae occupies a noticeable position in the Caatinga, while it has a lower rank in the Atlantic Rainforest, Amazon Rainforest and Pantanal. Amongst the families present in four biomes, Orchidaceae figures as the most important family in the Atlantic Rainforest, while it appears in second place in the Amazon Rainforest, in third in the Cerrado and in tenth place in the Caatinga. Cyperaceae is the third most important one in the Pampa, while Malvaceae is the third most important one in the Pantanal. Among the families that appear in a single biome, Bromeliaceae is the most noticeable one as the third most important family in the Atlantic Rainforest. Pampa and Pantanal have families among their top 10 that are not in the list of the most diverse ones in other biomes (Tab. 9), with five exclusive for the Pampa (Iridaceae, Solanaceae, Convolvulaceae, Cactaceae and Amaryllidaceae) and three in the Pantanal (Bignoniaceae, Sapindaceae and Malpighiaceae). Brazilian Geopolitical divisions - Regions and States Regarding geopolitical subdivisions there was an increase in the number of Angiosperm species recorded for the Southeastern region that continues to be the most diverse (16,127 vs. 15,550 species in 2010), being the one with highest number of endemic species (5,690), representing 35.3% of endemism (Tab. 10). On the other hand, the South has the smallest number of species (7,566 vs. 7,333 in 2010) and the lowest endemism as well (1,004 species, 13.3% endemism). In relation to Gymnosperms (Tab. 11), the North is the most diverse (17 species) and the Northeast the least diverse (two species). For Southeast and South regions there was a decrease of two species in 2015 in relation to 2010 (Tab. 11). Amazonas is the state with the highest number of species for Gymnosperms (12 species), followed for other two states that have part or all their territory included in the Amazonian biome (Tab. 12): Acre (8 species), Rondônia (7), Mato Grosso (6) and Pará (6). Six states (Amapá, Maranhão, Paraíba, Piauí, Rio Grande do Norte, and Tocantins) have no Gymnosperm records so far (Tab. 12). The seven Brazilian states with the largest number of Angiosperm species remained in the same order found in 2010 (Tab. 13), with increased number of species varying from 8% to nearly 17%, as follows: Bahia (1,284 species %), Minas Gerais (849 species - 8.2%), Amazonas (733 species - 9.7%), São Paulo (604 species - 8.7%), Rio de Janeiro (586 species - 8.7%), Pará (652 species %) and Paraná (629 species - 12%). Below the seven states with the largest number of species, different addition rate of new records has caused inversions in states position in relation to what was found in Mato Grosso overtook Goiás, Espírito Santo surpassed Santa Catarina, Rio Grande do Sul overtook Acre, Rondônia passed Pernambuco, Roraima passed Maranhão and Tocantins overtook Piauí. The same trend of dramatic species number increase found within the biomes with the least species was true of the states with least number of species, such as Sergipe, that more than doubled its species count from 770 to 1,608 (103.8% increase), while Rio Grande do Norte (72.8%), Tocantins (70.3%) and Alagoas (63.3%) increased their numbers by more than 50%. In terms of Brazilian states (Tab. 13), there is a trend of sharp drop in endemism for Rio de Janeiro (less 105 endemic species, or -8.9% than in 2010), Acre (less 50 species, or -46.3%), São Paulo (25 less species, -7.6%), Goiás (24 less species, -4.6%), Mato Grosso (15 less species, Rodriguésia 66(4):

166 Seed Plant diversity in Brazil. -5.7%), Maranhão (10 less species, -20.8%), Amazonas (9 less species, -1.3%), Pará (8 less species, -2.8%), Pernambuco (7 less species, -9.2%), Rondônia (6 less species, -12.8%) and others states with 5 or less species. There was an increase in the number of endemic species registered for Espírito Santo, with 92 more endemic species, showing an increase of 19.5%, Rio Grande do Sul (41 species, 23.2%), Mato Grosso do Sul (14 species, 23.7%) and Roraima (5 species, 17.2%). Table 12 Gymnosperm species number per Brazilian State and Federal District. States and Federal District Species number 2015 Species number 2010 Amazonas Acre 8 8 Rondônia 7 4 Mato Grosso 6 5 Pará 6 4 Paraná 4 6 Minas Gerais 4 4 Goiás 4 5 Rio Grande do Sul 4 7 São Paulo 3 6 Santa Catarina 3 6 Rio de Janeiro 3 5 Roraima 3 0 Bahia 2 2 Distrito Federal 2 3 Espírito Santo 2 3 Alagoas 1 1 Ceará 1 0 Mato Grosso do Sul 1 1 Pernambuco 1 1 Sergipe 1 1 Amapá 0 1 Maranhão 0 0 Paraíba 0 0 Piauí 0 0 Rio Grande do Norte 0 0 Tocantins Life-forms The proportion of life-forms varies within the different biomes, with trees appearing in larger proportion in the Amazon and Atlantic Rainforests, while herbs predominate in the Pampa (Tab. 14, Fig. 2). Other biomes, such as the Pantanal, Cerrado, Caatinga and, partly, the Atlantic Rainforest also have strong presence of herbs, but this is less extreme than what was found for the Pampa. Lianas are more expressive in the Amazon and Atlantic Rainforests, followed by the Pantanal. When comparing the profiles of the different biomes (Fig. 2), the Atlantic Rainforest and the Cerrado appear to be most comparable, also resembling what was found for the Caatinga, with a balanced presence of shrubs and trees, and 30% or more of herbs, with moderate numbers of lianas. The proportion found between tree and shrub versus herbaceous species, including lianas, varies amongst the biomes (Tab. 14). The lowest proportion was recorded for the Amazon Rainforest, where for each tree species there are two shrubs or herbs, i.e. a ration of 1:2; while the Atlantic Rainforest presents 1:4 proportion. For the Caatinga, it was found to be 1:6, the Cerrado has 1:7 and the Pantanal, 1:8. The grassland dominated Pampa has a proportion of 1:31, i.e. one tree species for over 30 species of shrub, subshrub, herb or liana. Substrate The majority of plant species throughout Brazilian biomes are terrestrial (Fig. 4), ranging from 83.7% in the Amazon Rainforest through to 75.6 in the Atlantic Rainforest (Tab. 15). Figure 2 Angiosperm life-form species number per biome. The exact values are found in table 14. Rodriguésia 66(4):

167 1098 The Brazil Flora Group Table 13 Angiosperm species number per Brazilian State and Federal District. States and Federal District Species number 2015 Endemic species 2015 Endemic species % 2015 Species number 2010 Endemic species 2010 Endemic species % 2010 Total species growth % Endemic species growth % Minas Gerais 11,239 2, ,039 2, Bahia 8,970 1, ,686 1, Amazonas 8, , São Paulo 7, , Rio de Janeiro 7,354 1, ,768 1, Pará 6, , Paraná 5, , Mato Grosso 5, , Goiás 5, , Espírito Santo 5, , Santa Catarina 4, , Rio Grande do Sul 4, , Acre 4, , Mato Grosso do Sul 3, , Rondônia 3, , Distrito Federal 3, , Pernambuco 3, , Roraima 3, , Maranhão 2, ,3 2, Amapá 2, , Ceará 2, , Tocantins 2, , Piauí 1, , Paraíba 1, , Alagoas 1, , Sergipe 1, Rio Grande do Norte 1, The Pantanal has the largest proportion of plants inhabiting aquatic substrates (14.2%), followed by Pampa (7.3%), Caatinga (4.5%), Amazon Rainforest (3.1%), Cerrado (3.0) and Atlantic Rainforest (2.4%). Caatinga (10.3%), Cerrado (9%) and Pampa (8.5%) have a high representativity of rupicolous plant species (Tab. 15). The Atlantic Rainforest is the biome with the highest number of epiphytic species, with 12.2%, followed by the Amazon Rainforest (7%). If taking into account epiphytes plus hemiepiphytes, the number is even higher for the Atlantic Rainforest (13%), followed by the Amazon Rainforest (9%), Cerrado (3.7%), Caatinga (3%), Pampa (0.9%), and Pantanal (0.8%). Parasitic plants, despite the fact that the Amazon Rainforest (133 species), Cerrado (125) and Atlantic Rainforest (122) present the highest values of hemiparasites (Tab. 15), the largest proportional representation is found in the Pantanal (1.6%), followed by the Caatinga (1.3%). Saprophytes are present in the Amazon Rainforest (0.3%), Atlantic Rainforest (0.2%) and Cerrado (0.1%), but were not recorded so far for Caatinga, Pampa and Pantanal (Tab. 15). Rodriguésia 66(4):

168 Seed Plant diversity in Brazil. Vegetation types Within the 24 vegetation types recognized by the Brazilian List (Lista do Brasil 2015), including Anthropized areas (Tab. 16), the highest Angiosperm number of species is found in Ombrophilous forest (12,000 species). Analysing only the species total, this is followed by Cerrado lato sensu (7,210 species), Terra firme forest (5,722) and Campo rupestre (4,928), Gallery Forest (4,259 species) and Semideciduous Seasonal Forest (3,384 species). Grassland reaches almost 3,000 species while Figure 3 Map of Brazilian biomes showing the total number of seed plant species (top number), the number of endemic species (middle number), and the percentage endemism for each biome. *Podocarpus barretoi Laubenf. & Silba is the only endemic species of Gymnosperm. Figure 4 Substrate types in the Angiosperms Restinga, Altitude Grassland and Caatinga stricto sensu, as well as the Anthropized areas, have around 2,000 species records each. All other vegetation types were scored for less than 1,900 species, and the smallest number of species was found in the Mangrove (81 species) (Tab. 16). Species restricted to a single vegetation type show a slightly different pattern from that found for total species richness (Tab. 16), with Ombrophilous forest (5,210), Terra firme forest (2,014), Campo rupestre (1,994), Cerrado lato sensu (1,799) and Grassland (638) topping the list. Lower richness of endemic species was found in comparison to restricted species, also with differences in relation to their distribution in different vegetation types. The largest number of endemic species per vegetation type was, again, recorded for Ombrophilous forest (4,159), followed by Campo rupestre (1,951), Cerrado lato sensu (1,357), Terra firme forest (598) and Altitude grassland (479). Mangrove had no endemic species, while Palm grove had only six endemic out of 240 species (Tab. 16). When analysing the relative endemism, the Campo rupestre has the highest percentage of endemism (1,951 endemic out of 4,928 species, 39.6%), followed by the Ombrophilous Forest (34.7%), Altitude grassland (21.8%) and Cerrado lato sensu (18.8%). Aquatic vegetation has shown the highest percentage of restricted species (45.1%) but much lower endemism, as was also reported for the Terra firme forest (35.2%). Vegetation types where the number of restricted species is similar to the number of endemics were Carrasco, Campo rupestre, Caatinga stricto sensu, Palm grove, Mixed ombrophilous forest, Restinga and Semideciduous seasonal forest. While the Angiosperms are present in all vegetation types (Tab. 16), the Gymnosperms occur only in seven of those (Tab. 17), predominantly in the Terra firme forest (eight species), Gallery Forest (six) and Amazon savanna (four). The latter, together with the Terra firme forest present the highest number of species restricted to these vegetation types, with four each (Tab. 17). Rodriguésia 66(4):

169 1100 The Brazil Flora Group Table 14 Life-form of Angiosperm species analysed by biome. The proportion tree:herbaceous was calculated by dividing the tree species from the sum of all other life-forms (shrubs, subshrubs, herbs, lianas). Proportion indicates the real value of trees divided by herbaceous. Life forms Atlantic Rainforest Amazon Rainforest Cerrado Caatinga Pampa Pantanal Tree 3,343 4,725 1, Shrub 3,491 2,821 3,380 1, Subshrub 2,102 8,02 3,185 1, Herb 6,663 3,648 5,189 1,812 1, Liana or climber 1,900 1,901 1, Proportion tree:herbaceous Proportion 1:4 1:2 1:7 1:6 1:31 1:8 Table 15 Substrate of Angiosperm species analysed by biome. Numbers between parentheses represent the percentage of the substrate in relation to the total species number found in the biome. Substrate Atlantic Rainforest Total and (%) Amazon Rainforest Total and (%) Cerrado Total and (%) Caatinga Total and (%) Pampa Total and (%) Pantanal Total and (%) Aquatic 395 (2.4) 382 (3.1) 387 (3.0) 230 (4.5) 137 (7.3) 201 (14.2) Epiphyte 1,980 (12.2) 876 (7.0) 419 (3.2) 127 (2.5) 16 (0.9) 7 (0.5) Hemiepiphyte 135 (0.8) 244 (2.0) 62 (0.5) 26 (0.5) 0 (0.0) 4 (0.3) Hemiparasite 122 (0.8) 133 (1.1) 125 (1.0) 65 (1.3) 21 (1.1) 23 (1.6) Parasite 18 (0.1) 9 (0.1) 23 (0.2) 7 (0.1) 4 (0.2) 2 (0.1) Rupicolous 1,273 (7.8) 354 (2.8) 1,177 (9.0) 529 (10.3) 159 (8.5) 48 (3.4) Saprophyte 31 (0.2) 38 (0.3) 16 (0.1) 0 (0.0) 0 (0.0) 0 (0.0) Terrestrial 12,284 (75.6) 10,448 (83.7) 10,864 (83.1) 4,175 (80.9) 1,540 (82.0) 1,135 (79.9) Rodriguésia 66(4):

170 Seed Plant diversity in Brazil Table 16 Angiosperm species number scored for each vegetation type. Vegetation type Species total Restricted to the vegetation type Endemic and restricted to the vegetation type Restricted species % Anthropic area 2, Caatinga stricto sensu 2, Campinarana 1, Altitude grassland 2, Várzea grassland Grassland 2, Campo rupestre 4,928 1,994 1, Carrasco Cerrado lato sensu 7,210 1,799 1, Gallery forest 4, Igapó forest 1, Terra Firme forest 5,722 2, Várzea forest 1, Deciduous seasonal forest Evergreen seasonal forest Semideciduous seasonal forest 3, Ombrophilous forest (Pluvial forest) 12,000 5,210 4, Mixed ombrophilous forest 1, Mangrove Palm grove Restinga 2, Amazonian savanna 1, Aquatic vegetation Rock outcrop vegetatiom 1, Endemic species % Rodriguésia 66(4):

171 1102 The Brazil Flora Group Figure 5 Above: Distribution map provided to the contributors by the on-line system, showing herbarium records distributed throughout Brazil and neighbouring countries. Below: Table provided to the contributors by the on-line system, relating the crosses to the herbarium data and enabling them to see provenance and determiner of each specimen. Rodriguésia 66(4):

172 Seed Plant diversity in Brazil Table 17 Gymnosperm species number scored for each vegetation type. Vegetation type Species total Restricted to the vegetation type Campinarana 3 1 Gallery forest 6 3 Terra firme forest 8 4 Ombrophilous forest (Pluvial forest) 1 0 Mixed ombrophilous forest 1 0 Restinga 1 0 Amazonian savanna 4 4 Discussion Overall The continued refinement of the Brazilian List over five years involved intensive review of the data already entered, with the addition of new records for Brazil, synonymization of names into already accepted names, updating names following new taxonomic framework, and specialist checking and accepting newly described species. Species number grew through checking and accepting newly described species and adding new records for Brazil. While for the Gymnosperms all the included names were checked both in 2010 and 2015, Angiosperm names had around 82% of their names checked in A considerable number of names (both accepted and synonyms) were added to the system and checked by the specialists. The number of new Angiosperm names included in the system between 2010 and 2015 was 9,274 (11.6%), growing less dramatically in relation to the total numbers, where 16,623 (17.7%) were added, reflecting mostly the activity of the phycology and mycology specialist groups, for which the state of knowledge was always known to be less developed (Maia & Carvalho Jr 2010; Bicudo & Menezes 2010). In relation to 2010, the number of checked names of Angiosperms grew 30.6%, addressing both the newly included names and the extant, but not yet checked, names from the 2010 edition that were then estimated at 18.5% (calculated from Table 1). The list is now in a much better situation, with almost 94% of its names checked and, wherever possible, either marked as accepted names or as synonyms linked to an accepted name. As an indication, there were 1,191 new names of Angiosperms published for Brazil in the period between 2010 and 2014 (IPNI 2015). Not all these new taxa were added as accepted species to the list as some of them were considered synonyms by the specialists, but it is likely that a large part of them would have been added. According to Sobral & Stehmann (2009), Brazilian new species represented an average of 7.8% of the plants described for the world between 1990 and In the years prior to this project ( ), we found that the average percentage of Brazilian plants described compared to the rest of the world was around 8.5%, while from 2010 this figure appears bigger, at 12.2%. An acceleration in the number of species described for Brazil during the period of the project can also be noticed, as from 2004 to 2009 the average species described for Brazil per year was /-36 while from 2010 to 2014 the figure increased to / New records for Brazil were spotted through the collection, study and naming of new material, much of which was promptly digitalized by herbaria and made available on-line through the portal of the Brazilian List. New records were also highlighted by catalogues, such as Zuloaga et al. (2008) and Jørgensen et al. (2014), floras and monographs (i.e. Medeiros et al. 2014). Included in the present figures is also the decrease of species numbers resulting from synonymization and changes of circumscription of accepted species, and exclusion of species that were cited for but not effectively located within the national territory thus far. Better knowledge is becoming available regarding increase in species range beyond the Brazilian borders and prior errors originated by the default setting as endemic during the successive revisions of the Brazilian List were corrected. Even taking these readjustments into account, we saw an overall steady increase both for accepted species Rodriguésia 66(4):

173 1104 The Brazil Flora Group totals (3%) and endemic species (4.5%, or 4.1% if we eliminate 62 indications for Anthropized area from Table 16) between 2010 and The number of threatened plant species currently found in the Brazilian Red List (Tab. 4) is slightly above 6% the total number of species of seed plant accepted for Brazil. When considering only Brazilian endemic species, the number of threatened plants represents 9.6% of the species. According to Brummitt et al. (2015), the expected number of species endangered worldwide represents around 20-26%. For specialist groups that were globally assessed, such as Cactaceae (Zappi et al. 2012), almost 32% of the species belong to one of the three categories of threat. The smaller proportion currently recorded for Brazil as a whole reflects the fact that the red-listing process in Brazil did not depart from a complete list of taxa, but focused in former lists of endangered species suggested by botanists. In order to get to a realistic picture of how endangered the Brazilian flora really is, a blanket approach studying family by family, species by species is urgently needed. Top 10 families This suite of families represents 47.2% of the species richness of Angiosperms in Brazil, having been increased by 68.5% (630 out of the 921 species) in relation to the number of species recorded in 2010 (Forzza et al. 2010). In terms of endemism, the increase was more expressive, at 77%, or 609 of the total of 791 endemic species newly recorded for Angiosperms between 2010 and 2015 belonging to the top 10 families. The Apocynaceae, formerly the tenth largest family (Forzza et al. 2010), was the only family to be surpassed in species richness by the Malvaceae, number 10 in Apart from the Poaceae, which has decreased in size by 120 species (-8.6%), the increase in species numbers was between 1 and 26.6% for the largest families, with Rubiaceae (2.1%), Fabaceae (2.3%) and Asteraceae (2.4%) showing the slowest increase while Euphorbiaceae (21.5%) has been the fastest growing family between 2010 and 2015, with a difference well above the 3% found for the Angiosperms as a whole. Two other families that have also markedly increased in size during this period were the Bromeliaceae (11.3%) and the Myrtaceae (11%). Sobral & Stehmann (2009) pointed out Bromeliaceae as one of the five fastest growing families together with the Orchidaceae, Fabaceae, Asteraceae and Poaceae. By 2015, while Bromeliaceae continues to grow at a high rate, increase in new species in other families has slowed down, with Orchidaceae at medium pace, Fabaceae and Asteraceae presenting slow increase, and the Poaceae decreased in size through the elimination of over 200 cultivated and introduced species from the present calculations. The increase in the growth speed of the species richness in Euphorbiaceae and Myrtaceae reflects a possible change in the balance of the activities of researchers describing biodiversity in Brazil during the last few years. It is possible that, for less popular and more complex, speciose plant families such as these ones, this acceleration may be a direct result of the present project. In terms of endemism, the three families where the endemism figures grew faster are Euphorbiaceae (26.6%), Bromeliaceae (14%) and Myrtaceae (12.7%), all well above the growth found for Angiosperms as a whole (4.5%). On the other hand, Orchidaceae has only 16 more endemic species than five years ago, increasing its percentage of endemism by only 1%, while Asteraceae had a modest increase of 2.2%. 30 largest genera Amongst the 30 top genera, 19 belong to the 10 top families, thus influencing the numbers exposed above. The largest species increase took place in Croton, where the increase of the number of endemic species was also the highest. Following the creation of an international group to study this mega-genus in 2001 (Berry 2015), around 15 new species of Croton were described between 2010 and 2015, and more new discoveries are being published. However, the 130 species difference cannot be explained by the discovery and description of new species, and it stems from a delay in including existing information (checking all accepted names) that some groups faced during the implementation of the Brazilian List online. This is the most high profile example of instances that occurred in other, smaller groups, and by no means the rule within this mainly successful project. In most cases where there was a significant decrease in species numbers within a genus, the adjustment could be tracked back to a new generic circumscription; for example several species formerly included in Hyptis were transferred to five recently recognized genera (Cantinoa, Cyanocephalus, Eplingiella, Medusantha, Mesosphaerum, and Oocephalus; see Pastore et al. 2011; Harley & Pastore 2012). Rodriguésia 66(4):

174 Seed Plant diversity in Brazil. Endemism reduction that was linked to a decrease of species number was seen in Paepalanthus (-16 species) and Leandra (-one species). Adjustments regarding endemism that were not always linked to a reduction in species number were seen in Philodendron (-48 species), Piper (-12 species), Peperomia (-seven species) and Epidendrum (-one species). The more extreme case, that of Philodendron, can be explained by indepth research in the Amazonian species, with the study of additional specimens of species previously thought to be endemic to Brazil collected in bordering countries, such as Colombia, combined with the fact that the initial default setting of the Brazilian List was marked ON for endemism. Readjustments made within Philodendron show that the endemic species of the genus are, in their majority, from the Atlantic Rainforest. The revision process during the last five years was supported by growing evidence from collections of a vast number of herbaria that went on-line during this period, thus enabling the botanists to visualize species dot maps and perform important corrections for the endemism in their groups (Fig. 5). The top 10 families have shown a slightly higher proportional growth in endemism than the 30 largest genera that remained collectively similar in terms of their endemism between 2010 and It is possible that the numbers for the group of 30 top genera were influenced by the dramatic drop in endemism caused by further studies in Paepalanthus, Piper and Philodendron. Biomes The overall trend of growing species numbers was maintained, to a certain extent, in data concerning the Brazilian biomes. The almost exclusively Brazilian Atlantic Rainforest and the Cerrado continue to lead in terms of biodiversity and endemism amongst seed plants (Fig. 3), coinciding with the hotspots highlighted by Mittermeier et al. (1998, 2004) and Myers et al. (2000). They are followed by the Amazon Rainforest and the Caatinga; and the Cerrado, a biome currently under severe threat of rapid destruction due to the expansion of the agricultural frontier, has now a wider gap in relation with the Amazon Rainforest through increased species and endemism records added during the last five years. While the number of species now recorded for the Atlantic Rainforest has increased 7.4% (1,029 species) over the number presented by 1105 Forzza et al. (2010), the records of endemic species has been less expressive, with only 418 extra endemic species, or 6%, added to this biome between 2010 and 2015, causing a fall in endemism percentage from 50.2% (Forzza et al. 2010) to 49.5%. For the Cerrado, the increase in species number was smaller, at 6.3% (713 species), while the endemism increase was also proportionally smaller, at 2.5%, with 102 endemic species record added to this biome, also a fall in the percentage of endemic species from 36.5% (Forzza et al. 2010) to 35.1%. It is possible that this rise in species numbers, yet not followed closely by the increase of endemic species, results from refinement of the knowledge of plant distributions within and beyond each biome, for instance the presence of Atlantic Rainforest species in other biomes by the way of gallery forests in the Cerrado or of enclaves of humid highland forest in the Caatinga, or by species being recorded beyond the Brazilian borders within the same biome (for instance in the Bolivian Cerrado or in the extension of the Atlantic Rainforest into Paraguay and Argentina). Meanwhile the Caatinga, the only exclusive Brazilian biome, saw an increase of 7.8% (337 species) with 169 records of endemic species added, or an increase of 22.7% from the 2010 data, bringing the percentage of endemic species from 17.2% (Forzza et al. 2010) to 19.7%. This is the only biome where the percentage of endemic species has risen during the period and this is connected to the local botanical activity in the universities, and may also be related to the compilation of lists and floras that include the Caatinga biome (i.e. Siqueira Filho 2012; Prata et al. 2013). Both Pampa and Pantanal, with respectively 340 species (25.3%) and 392 species (44.3%) added, have seen dramatic increases in the biodiversity recorded. In the Pantanal this may be a reflection of recent efforts employed to produce an up-to-date checklist for Mato Grosso do Sul (Farinaccio et al no prelo). However, even if the addition of eight endemic species to the 2015 list meant an increase of 17.4%, the overall percentage of endemic species in this biome has decreased from 5.2% to 4.3%, no doubt because a great many species from the Pantanal are also found in Argentina, Bolivia and Paraguay (Sarmiento 1983, Haase & Beck 1989, Larrea-Alcázar et al. 2010). The recent surveys that inform the Bolivian Catalogue (Jørgensen et al. 2014) probably Rodriguésia 66(4):

175 1106 The Brazil Flora Group contributed with additional evidence of new examples of such taxa. In the Pampa, the addition of 26 endemic species (34.2%) did not change the overall percentage of endemic species, that remains almost unchanged, at 5.7%. It is possible that the increase in species number for the Pampa is a consequence of comparisons with the Catalogue of the Cono Sur (Zuloaga et al. 2008) and the concentration of local effort to catalogue the Flora of Rio Grande do Sul. In the case of the Amazon Rainforest, the increase of 4.8% (547 species) was the smallest increase in terms of species total. Moreover, the percentage of endemic species for this biome has decreased by 48 (-2.5%) species from 1,948 in 2010 (Forzza et al. 2010) to 1,900 in 2015, with an overall endemic percentage drop from 17.2% to 16.1%. Considering the large expanse covered by this biome, it is surprising that it now falls behind the Cerrado in terms of species richness, but, in terms of species endemism, this result is justified. The Cerrado has more diverse and abundant vegetation types than the Amazon Rainforest, however, and includes large sections spanning over much wider altitudinal and latitudinal ranges. On the other hand, the fact that the Cerrado is more accessible and better studied, thus better represented in herbaria, may play some role in the recording of known species number. Apart from the survey work developed for the whole state of Acre by Daly & Silveira (2009), research in the Amazon Rainforest has focused on isolated locations. Survey work such as the Flora of the Reserva Ducke, by Ribeiro et al. (1999), looking at all life-forms rather than focussing on woody species, has shown that systematic and complete collections may add large quantities of known and not yet described species to each locality studied within this biome. Comparing the two studies cited above, the proportions between woody and herbaceous life-forms are different. According data from Daly & Silveira (2009), the proportion between trees and other life-forms found for all vegetation types in Acre was 1:2; while the data from Ribeiro et al. (1999) indicated a proportion of 1:1 between trees and herbaceous plants for an area of Terra Firme forest. The research carried out on the Amazon basin by ter Steege et al. (2013) suggests that there are possibly between thousand tree species. Moreover, from our findings we now understand that there are 2 or more shrubby or herbaceous species for each Amazonian tree species. Considering these proportions, Hopkins (2003) projection that the Brazilian Amazon Rainforest might harbour between 40 and 50 thousand species of seed plants, many of them yet undescribed, seems to be more appropriate for the whole basin. Hopkins (2007) and ter Steege et al. (2013), point at the poor state of floristic knowledge of the Amazon basin. In addition, botanical research in the Amazon Rainforest has been biased towards studying its tree flora, but there is much else to be discovered besides this if more inclusive surveys of all life-forms were to be carried out. Analysing the number of 0,18 exsiccate per km 2 (including duplicates) in the Northern region (Specieslink 2015) the need of intensive and systematic work in this area to cover the gaps of knowledge about its biodiversity becomes evident. When analysing the different biomes according to their more abundant families in terms of species, it was found that their shared ten top families were only Fabaceae, Poaceae and Rubiaceae, all extremely species rich and not exclusively woody (in fact, Poaceae can be considered marginally woody when bamboos are taken into account). Asteraceae and Euphorbiaceae appear in five biomes, while Melastomataceae and Orchidaceae appear in four, together with Apocynaceae and Cyperaceae, which are not amongst the overall top ten families. Malvaceae appears in three biomes. Families that appear amongst the top ten within a single biome are seldom amongst the overall top ten (Bromeliaceae, Myrtaceae), more often making a single apparition (Amaryllidaceae, Annonaceae, Araceae, Bignoniaceae, Cactaceae, Convolvulaceae, Eriocaulaceae, Iridaceae, Malpighiaceae, Sapindaceae and Solanaceae). In the case of Myrtaceae and Bromeliaceae, both possessing a center of diversity in the Atlantic Rainforest, their sheer number of species is so large within that biome that it supports their presence within the top ten. The Eriocaulaceae is a point in case, with huge biodiversity within the Campos Rupestres that are included within the Cerrado biome, thus occupying the sixth position within this very rich biome, despite not being one of the overall top ten families. In biomes with less species, specific characteristics pertaining these individual families, or one offs, that may be favouring their presence amongst the top ten, are the woodiness of Annonaceae in the Amazon Rainforest, the climbing habit of Bignoniaceae, Malpighiaceae and Rodriguésia 66(4):

176 Seed Plant diversity in Brazil. Sapindaceae in the Pantanal and the corm and bulbbearing, herbaceous Iridaceae and Amaryllidaceae found to be expressive in the Pampa. Also in the Pampa, the expressivity of rupicolous plants may have led to the position occupied by the Cactaceae. Brazilian Geopolitical divisions - Regions and States While a balance of 921 new native seed plant records included for Brazil, the number of additional state records is much larger, in excess of 19 thousand new state records entered in the system during the last five years. The release of a facility that generates a provisional map by plotting the underlying herbarium collections made available to the specialists in 2011 has guided them towards including many records by observing the map and checking the specimen identification data. As some of these herbarium collections included specimens from outside Brazil, the map was also useful to double-check endemism of some taxa, encouraging further research and adjustments (Fig. 4). States with floras and lists partially or totally published by 2010 have shown a smaller increase in the number of species than states where the floristic knowledge is still incomplete. The list of plants from the Catalogue of the Flora of Acre (Daly & Silveira 2009) was one of the databases added to the system when it was created, and this was reflected in a modest increase of species for this state (7.7%), while dramatic adjustments to the number of endemic species for the state were made between 2010 and 2015, with a fall in the number of endemic species of the order of -46.3%. The recent growth of knowledge regarding the flora of Acre is described by Medeiros et al. (2014). Also in the process of organizing its Catalogue, the state of Rio de Janeiro (Baumgratz et al. 2014) saw moderate growth in the species number (586 species, growth of 8.7%) and a fall on the number of endemic species (105 species, decreasing -8.9%). On the other hand, states such as Paraná, whose list was recently published (Kaehler et al. 2014), using Forzza et al. (2010) as its starting point, saw an increase of 629 species, with 12% growth in the species number and a slight increase in the number of endemic species. Another ongoing project that may probably be contributing to an accelerated increase of the species count is the checklist of Espírito Santo (Dutra et al. in press), focussing on this still poorly known and extremely biodiverse state where 1,333 new species records, or a percentual growth 1107 of 33.6%, took place during the last five years. It is possible that an intensification of work within the state of Espírito Santo has resulted in range extension of many species formerly thought to be endemic to Rio de Janeiro, resulting in the decrease of endemism seen for the latter state. Through the publication of the Catalogue (Forzza et al. 2010), the states of the Northeastern region of Brazil were stimulated to record their flora at an intense pace, with increases of more than 28% on the species numbers for all states (Sergipe 103.8%, Rio Grande do Norte 72.8%, Alagoas 63.3%, Paraíba 44.4%, Piauí 40.7%, Pernambuco 29.4% and Ceará 28.5%), with exception of Bahia, the second most species rich state, which saw a respectable increase of 1,284 species (16.7%), and Maranhão with an increase of 441 species (18.3%). Finally, the state with the highest growth percentage in terms of species numbers was Sergipe, probably due to the publication of its own Flora project (Prata et al. 2013). Some Amazonian states show that, despite the publication of the list, the knowledge seems to have stalled and did not improve much during the 2010 to 2015 period, namely in the Amazonas, with less than 10% increase, and Pará, with 11.6%. The state of Tocantins is a relatively recent split of a former larger state of Goiás and many of the species historically referred to Goiás have wider distribution, occurring also in Tocantins. The growth of the number of species from this state has reached an increase of 70.3%, occupying the third place in growth, after Sergipe and Rio Grande do Norte. The mapping facility has made this situation more obvious and hence there was a drop in the number of endemic species from Goiás (-4.6%) due to their newly recorded occurrence in Tocantins. Life-forms When comparing the proportion of tree species with the other life-forms, such as shrubs, herbs and lianas, there is wide variation among the Brazilian biomes. Five of the biomes (Atlantic Rainforest, Cerrado, Caatinga, Pampa and Pantanal) show a majority of herbs amongst their life-forms, while tree species are more expressive in the Amazon Rainforest, but even so in a proportion of 1:2. Even in biomes composed mostly by forested vegetation, such as the Amazon and the Atlantic Rainforests, different proportions have been found, with the Atlantic Rainforest showing a proportion of 1:4. This diversity of life-forms in Rodriguésia 66(4):

177 1108 The Brazil Flora Group the Atlantic Rainforest biome is justified by the presence of many different vegetation types in this biome, including Restinga, Rock outcrops, Ombrophilous, Semideciduous and Mixed forests, Altitude grassland and even outlying Campo rupestre. The Atlantic Rainforest is also a centre of diversity for Bromeliaceae and Orchidaceae, both herbaceous families with expressive numbers of epiphytes. Amongst biomes where the vegetation is mostly composed by different savanna types, the Caatinga has a proportion of 1:6, and the Cerrado has 1:7. At the other end of the spectrum from the Amazon Rainforest, open habitats such as the Pampa and Pantanal count with less tree species, being rated at respectively 1:31 and 1:9. The Pampa is the only biome characterized by large extensions of grassland, and the diversity of tree species is very reduced under its subtropical conditions. While the lianas are less well represented in the Atlantic Rainforest, Cerrado and Caatinga, subshrub species are very few in the Amazon Rainforest, and trees, as already mentioned, are the least represented life-form in the Pampa and the Pantanal. Such proportions represent a useful snapshot of the overall different types of biomes in Brazil: forests, savannas and grasslands. The presence of some exclusive families amongst the top ten families in the different biomes provides, in some cases, a link with this information. The abundance of species of Malpighiaceae, Sapindaceae and Bignoniaceae in the Pantanal may be linked to the presence of habitats suitable for climbers, even if the biome possesses low diversity of tree species. The high position in the top ten rank for herbaceous families such as Iridaceae and Amaryllidaceae in the Pampa is linked to the grassland habitats, together with the expressive numbers of species of Poaceae, Cyperaceae and Asteraceae. The presence of Fabaceae and Rubiaceae amongst the top 10 families in all six biomes help to understand the landscape diversity found within the Brazilian biomes, as they are families with huge amplitude in terms of life-forms, from small herbs to tall trees. Substrate While terrestrial plants represent constantly more than three quarters of the biodiversity in terms of substrate in all biomes, epiphytes were the second major component of the Atlantic Rainforest biome. The first and third position amongst the top ten families in the Atlantic Rainforest was occupied by Orchids and Bromeliads, both well represented in terms of epiphytic species. Other speciose group of epiphytes is Peperomia (Piperaceae), the 22 nd largest genus in Brazil. The Araceae, with almost 50% of the species occurring as epiphytes or hemiepiphytes, are prominent in the Atlantic Rainforest and also appear amongst the top ten families within the Amazon Rainforest biome. Despite their comparative low diversity in relation to the Atlantic Rainforest, epiphytism was also the second substrate occupied by plants in the Amazon Rainforest. While aquatic plants were, as expected, expressive in the Pantanal and Pampa, it could have been expected that they would occupy a larger proportion of the vegetation types available in the Amazon (Igapó and Várzea forests, Várzea fields, Aquatic Vegetation), however this was not conveniently highlighted by our data. However, in terms of proportion within the Amazon Rainforest biome, the aquatic plants occupy the third, following terrestrial and epiphytic substrate. The Caatinga, despite of its xeric conditions, was the third biome to display an expressive aquatic vegetation, due to the seasonal floods that create temporary lagoons teeming with life during the rainy season. The high number of rupicolous species found in the Cerrado and Caatinga biomes can be justified by the presence of the Campo rupestre vegetation type across both. This substrate is the second most relevant in the Pampa, where many low rock outcrops are found almost at the same level as the grass. Upon land conversion for agriculture or into pasture, these outcrops are quickly deteriorated and lose their originally restricted flora. In the Atlantic Rainforest the rupicolous substrate is the third most relevant following terrestrial and epiphytic, and this can be explained by the Altitude grassland (Martinelli 2007) as well as the Rock outcrop vegetation type, a term used for non-crystalline outcrops generally at lower altitude that do not fall within the definition of Campo rupestre. These Rock outcrops, or inselbergs, are also abundant within the Caatinga and the Cerrado. The ecotone between Campo rupestre and Caatinga stricto sensu, known as Carrasco, is also stony and harbours rupicolous species. Parasites and saprophytes are proportionally less expressive in the Brazilian flora, and occur basically in the Amazon Rainforest, Atlantic Rainforest, and Cerrado. Rodriguésia 66(4):

178 Seed Plant diversity in Brazil. Vegetation types The inclusion of the 24 vegetation types represented a huge advance in terms of the refinement of the knowledge Brazilian plant diversity, and it was not without difficulties. Similarly to what we experienced at the start of the project in 2009, contributors engaged differently with the input of this new request for information. These concepts were clearer for the botanists that are active in the field and have experience of collecting in different environments, however, due to the diversity of professionals involved in the project there were some obvious shortcomings in the data. An example of this was seen for Anthropised areas that were considered as an independent vegetation type in the list. These areas, however, can be originated from any of the 23 vegetation types that were submitted to drastic human actions, such as deforestation, degradation, urbanization, mining, etc. There were 162 native species listed as restricted to disturbed environments, of which 62 are supposedly endemic to Brazil. Other problems arose from the initial difficulty faced by the committee to reach an agreement for the vegetation types list. Ideally, those should be comparable from one biome to another, but in reality we settled for a list that recognizes a considerable number of local formations as unique. One good example is Terra firme forest, basically an Ombrophilous forest type from the Amazon basin, and Deciduous seasonal forest, a forest type that might be included by some in Caatinga stricto sensu. Likewise, it is possible that confusion generated between Campo rupestre and Rock outcrop vegetation. The analysis in those cases has to take into account that the conflicts may be handled differently and variable levels of consistency were achieved in the different groups. Mirroring what was found for the Atlantic Rainforest biome, its most expressive vegetation type, the Ombrophilous forest, has not only the highest number of species but a large percentage of species restricted to this type of forest in Brazil, and a large proportion of endemic species to this vegetation type. The Atlantic Rainforest is not totally endemic to Brazil, with outlying areas in Argentina and Paraguay, which account for the difference between the restricted species and endemism to this vegetation type in Brazil. Similarly, the Cerrado lato sensu is the most extensive vegetation type of a largely Brazilian 1109 biome that extends into Bolivia and Paraguay, and also in the Venezuelan llanos (Sarmiento 1983), thus the percentage of species restricted within Brazil is higher than the percentage of endemic Cerrado lato sensu species found in the country. The Terra firme forest is undoubtedly the most species rich vegetation type of the Amazon Rainforest biome, however the percentage of both restricted and endemic species are lower than for the former two, a fact that is fully justified by the vast sections of Amazon Rainforest found in the lowland areas of neighbouring countries such as Peru, Colombia, Venezuela and the Guianas, with many species widely distributed through many of these and beyond. The Campos rupestres, with their shared distribution in the Cerrado (Minas Gerais, Goiás, Tocantins) and Caatinga biomes (Bahia) occupy the fourth place in number of species, with 40.5% of the 4,928 being restricted to this vegetation type, with nearly all of those (39.6%) being also endemic to this vegetation type in Brazil. It is possible now to point to the fact that this extremely biodiverse vegetation type contributes towards the relatively high endemism percentages found at the biome level both for the Caatinga and Cerrado biomes. Recent data from the Serra do Cipó reveal that, within a single of mountain range, nearly three thousand vascular plants can be found (Pirani et al. 2015). In comparison, the Altitude Grassland in the Atlantic Rainforest contributes with a small number of species, reflecting its restricted geographic spread, nonetheless with 21.8% of endemic species. In contrast, for the Aquatic vegetation, where the specialized life-forms and requirement of the species have led to the highest percentage of restricted species (45.1%), the percentage of endemism is low because these species are widely distributed throughout Brazil and also into neighbouring countries. Concluding remarks The overall position of Brazil as the country harbouring the highest plant diversity (Forzza et al. 2012) continues to be accurate. Added to this, the endemism levels found in Brazilian seed plants continue to be outstanding, surpassed only by large island floras (such as Australia, Papua New Guinea, Madagascar), archipelagos (such as New Caledonia, New Zealand, French Polynesia) and South Africa (65%), the only other continental country where levels of endemic flora surpass 50%. Rodriguésia 66(4):

179 1110 The Brazil Flora Group There was marked improvement in the list as a realistic reference for species found in Brazil, apart from the somewhat disappointing slow growing records regarding the Amazon Rainforest biome, partly reflected both in the numbers shown for the biome, the Brazilian Northern region and some states that are included in it (Acre, Amapá, Amazonas, Pará, Roraima and Rondônia). On the other hand, the results obtained reinforce the important role of the Atlantic Rainforest and Cerrado on the diversity of Angiosperms in Brazil, and also of the Caatinga, as an exclusive Brazilian biome. Considering their smaller surface area, both Pampa and Pantanal contribute much towards the Brazilian plant diversity, even if they share many species with neighbouring countries. When considering the richest vegetation types in terms of their overall, restricted and endemic species number, it is possible to see that the country displays a balance between the forests and the more open habitats. Amongst the six richest types, four are forests (Ombrophilous, Terra firme, Gallery and Semideciduous Seasonal forests) and two are open vegetation types ( Cerrado lato sensu and Campo rupestre ). Amongst the restricted, two forests (Ombrophilous and Terra firme ) and four open vegetation types ( Campo rupestre, Cerrado lato sensu, Grassland and Altitude grassland). When focusing on endemism, two are forests (Ombrophilous and Terra firme ) while four are open habitats ( Campo rupestre, Cerrado lato sensu, Altitude grassland and Rock outcrops). The elevated total number of native species recorded for Anthropized areas (2,167) may still partly reflect different concepts used by contributors. The importance of this data, which includes a large number of possible pioneer plants with potential for reforestation and re-establishment of natural habitats cannot be underestimated (Elliott et al. 2013). Extremely interesting data have been compiled during the second phase of this endeavour, enabling more refined analysis of biomes according to habit, substrate and vegetation types. While the Brazilian forested biomes, namely the Amazon and Atlantic Rainforests, have respectively 1:2 (one tree to two shrubby/herbaceous species) and 1:4 (one tree to four shrubby/herbaceous species), savannic biomes, such as Cerrado, Caatinga and Pantanal, have ratios of 1:6, 1:7 and 1:8. In the grass dominated Pampa, the ratio is 1:31. After the predominant terrestrial habit, the second highest group are the epiphytes, more expressive in the Atlantic Rainforest and pushing up the numbers of Orchidaceae and Bromeliaceae into the higher positions in terms of species rich families. Rupicolous plants are very expressive in the Caatinga, Cerrado and Pampa. Doubtlessly more interesting links can be found by further refining the data made available in this study. Less well-known areas, such as the states of Tocantins, southern Maranhão, Piauí and northwestern Bahia, have shown increased volume of data. A large proportion of this biodiversity corresponds to the portions of these states covered by the Cerrado biome (2,130 of the 2,306 species found in Tocantins; 1,977 of the 2,855 species from Maranhão; 1,621 of the 1,992 from Piauí; and 1,977 of the 8,970 from Bahia). However, the future of the Cerrado natural resources is threatened by concerted efforts from the Brazilian government and national and international investors to further expand the arable land for soybean, cotton, sugarcane, and rice production (MaToPiBa 2015). The Cerrado in these states is already responsible for 10% of the country s grain production, however the recent growth of the knowledge regarding the plant species found in these states between 2010 and 2015, with Piauí s plant list growing by 40.7% and Tocantins by a staggering 70.3% suggests that such knowledge may still be far from complete. During the last five years, the relatively slow increase of the knowledge regarding the species found in the Amazon Rainforest is concerning, especially when we take into account the rampant deforestation and habitat change (large scale agriculture including soybean and cattle farming, construction of hydroelectric dams with huge immediate and long term impact on the environment, large-scale mining), all this supported by government sectors that are increasingly averse to conservation and hostile towards Brazil s need to preserve its natural habitats through protected areas. Acknowledgements Ministry of Science and Technology - MCTI; Brazilian National Research Council - CNPq; The Alberto Luiz Coimbra Institute for Graduate Studies and Research in Engineering at the Federal University of Rio de Janeiro - COPPE/UFRJ; Information Technology Department of the Rio de Janeiro Botanical Gardens - TI JBRJ; to Alexandre Antonelli and two anonymous reviewers that contributed valuable comments to the manuscript; Rodriguésia 66(4):

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182 Seed Plant diversity in Brazil. F.; Di Fiore, A.; Jimenez, E.M.; Penuela Mora, M.C.; Fernando Phillips, J.; Rivas, G.; van Andel, T.R.; von Hildebrand, P.; Hoffman, B.; Zent, E.L.; Malhi, Y.; Prieto, A.; Rudas, A.; Ruschell, A.R.; Silva, N.; Vos, V.; Zent, S.; Oliveira, A.A.; Cano Schutz, A.; Gonzales, T.; Nascimento, M.T.; Ramirez-Angulo, H.; Sierra, R.; Tirado, M.; Umana Medina, M.N.; van der Heijden, G.; Vela, C.I.A.; Vilanova Torre, E.; Vriesendorp, C.; Wang, O.; Young, K.R.; Baider, C.; Balslev, H.; Ferreira, C.; Mesones, I.; Torres-Lezama, A.; Urrego Giraldo, L.E.; Zagt, R.; Alexiades, M.N.; Hernandez, L.; Huamantupa-Chuquimaco, I.; Milliken, W.; Palacios Cuenca, W.; Pauletto, D.; Valderrama Sandoval, E.; Valenzuela Gamarra, L.; Dexter, K.G. & Feeley, K Hyperdominance in the Amazonian tree flora. Science 342: 325. DOI: /science Specieslink Indicadores. Available at < splink.cria.org.br/indicators/index?setlang=pt.>. Acess on 27 May Urban, I Index Familiarum. In: Martius, C.P.F. von; Eichler, A.W; Urban, I. & Oldenbourg, R. Flora brasiliensis. Monachii. Vol 1, pars 1, pp Zappi, D.C.; Taylor, N.P. & Larocca, J A riqueza das Cactaceae no Brasil. In: Ribeiro-Silva, S.; Zappi, D.C.; Taylor, N.P. & Machado, M.C. (eds.) Plano de ação nacional para a conservação das cactáceas. ICMBio, Brasília. Pp Zuloaga, F.O.; Morrone, O. & Belgrano, M.J Catálogo de las plantas vasculares del ConoSur (Argentina, Sur de Brasil, Chile, Paraguay y Uruguay). 3 vols. Monographs in Systematic Botany from the Missouri Botanical Garden 107: Artigo recebido em 28/05/2015. Aceito para publicação em 19/08/2015. Rodriguésia 66(4):

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184 Rodriguésia 66(4): DOI: / Flora of Brazil Online: Can Brazil s botanists achieve their 2020 vision? Marli Pires Morim 1 & Eimear M. Nic Lughadha 2 Abstract This paper seeks to provide an assessment of the prospects of delivering an online Flora of Brazil by Our approach is to evaluate the nature and extent of documentation of the Brazilian flora over the past 15 years before exploring whether existing botanical documentation and capacity is sufficient to support the production of a complete Flora over a five-year period. We address the following headline questions: Has a high proportion of Brazilian species been described? Are collection densities sufficient to underpin a national Flora? Is there sufficient botanical expertise available to execute such a big project over a five-year period? Are there current taxonomic treatments that support the development of a national Flora? The results obtained show that the proportion of the flora estimated to have been described is high enough to be confident that those preparing Flora treatments will not be overwhelmed by the numbers of species new to science requiring description; the national average of 1.08 specimens per km 2 is significant; relevant taxonomic treatments with a variety of different scopes and geographic scales are available; and, finally the botanical expertise available is demonstrated by the results presented in the List of Species of the Flora of Brasil. Key words: Angiosperms, herbaria, Reflora, sampling, Taxonomic. Resumo Este trabalho visa fornecer uma avaliação das perspectivas de concluir a Flora do Brasil on-line até Nossa abordagem é avaliar a natureza e a extensão da documentação da Flora Brasileira ao longo dos últimos 15 anos, antes de explorar se a documentação e capacidade botânica existentes são suficientes para apoiar a produção de uma Flora completa no período de cinco anos. Nós abordamos as seguintes questões temáticas: há uma proporção elevada de espécies brasileiras já descritas? A densidade de coleta é suficiente para sustentar uma Flora Nacional? Há expertise botânica disponível o bastante para executar um projeto tão grande ao longo de um período de cinco anos? Existem tratamentos taxonômicos atuais que apoiem o desenvolvimento da Flora Nacional? Os resultados obtidos mostram que a proporção da flora que se estima já ter sido descrita, é alta o suficiente para se ter a certeza de que os tratamentos destinados à elaboração da Flora não serão subjugados por números de novas espécies para a ciência necessitando de descrição; a média nacional de 1,08 espécimes por km 2 é significativa; tratamentos taxonômicos relevantes com uma variedade de diferentes escopos e escalas geográficas estão disponíveis; e, finalmente, a expertise botânica disponível é demonstrada pelos resultados apresentados na Lista de Espécies da Flora do Brasil. Palavras-chave: Amostragem, Angiospermas, herbários, Reflora, Taxonomia. Introduction Early impressions of and interest in Brazilian plant species and vegetation formations are reported by european visitors at the time of the discovery of Brazil (Andrade-Lima 1984; Filgueiras & Peixoto 2002). In his botanical analysis of the letter of Pero Vaz de Caminha to the Portuguese king D. Manoel I, Andrade-Lima (1984) highlighted the observation..the plants were seen before the land.... Reports, iconographic records, specimens and scientific expeditions to various regions of Brazil documenting the Brazilian flora all evidence the extensive research undertaken by naturalists from the 17 th to 19 th Centuries (Giulietti & Forero 1990; Peixoto 1999). Some 340 years after the discovery of the country by Europeans, publication 1 Instituto de Pesquisas Jardim Botânico do Rio de Janeiro, R. Pacheco Leão 915, , Rio de Janeiro, RJ, Brasil. mpires@jbrj.gov.br 2 Conservation Science Department, Royal Botanic Gardens, Kew, Richmond, United Kingdom. el02kg@kew.org

185 1116 Morim, M.P. & Nic Lughadha, E.M. of the first national Flora began: Flora Brasiliensis described 19,958 species including algae, fungi and plants (Martius et al. 1833; Urban 1906). Completing a national Flora of any country is an audacious objective in which many years of work are often invested in order to achieve the goal. The probability of success with such a task, and the complications and obstacles likely to be encountered while completing a national Flora depend on diverse factors, but three key considerations come to mind immediately which must form part of any such evaluation: (i) the geographical extent of the area to be treated by the Flora, (ii) the levels of richness and diversity of the species occurring therein and (iii) the available knowledge about these species. Considering these three primary parameters, the preparation and completion of a Flora of a large and megadiverse country in which many plant species remain to be discovered can be seen as an even greater challenge. Brazil is located in the Neotropical region which encompasses c. 40% of the terrestrial surface of the earth and is home to a surprisingly large number of living organisms (Forero & Mori 1995). Brazil s vast area (8,515,767 km 2, IBGE 2015) makes it the largest country in the Neotropics. Of the six phytogeographic domains or biomes occurring within Brazil: Amazônia, Cerrado, Caatinga, Mata Atlântica, Pampa and Pantanal, two are recognised as global biodiversity hotspots: Cerrado and Mata Atlântica (Mittermeier et al. 1998; Myers et al. 2000). These six phytogeographic domains represent a broad diversity of vegetation types, resulting from biotic and abiotic factors which influenced the history and evolution of the neotropics (Antonelli & Sanmartín 2011). In this context, it can readily be understood why the Brazilian flora has the greatest diversity of vascular plants of any country in the world (Forzza et al. 2010, 2012; BFG 2015). This unrivalled plant diversity encompasses an extraordinary variety of habits, life-forms and biological associations ranging from species confined to particular environments on different geographical scales to more generalist taxa with broader distributions, which often exhibit great morphological plasticity across their range. Authoritative, reliable taxonomic identification of species is fundamental to understanding species diversity and richness of life on earth, as well as providing the basis for defining conservation strategies consistent with rational use of natural resources and sustainable development (Trias-Blasi & Vorontosova, 2015). Efforts invested in floristic surveys, Flora treatments and monographs dealing with a whole family, genus or infra-generic group, and the biological collections associated with these activities are the primary sources for our knowledge of plant biodiversity (Thomas et al. 2012). The Convention on Biological Diversity (CBD 1992) had a marked impact on the perceived importance of biodiversity and on the regulation of access to biodiversity, as well as on the treatment of scientific questions relating to biodiversity. The scientific quest of researchers to estimate the number of plant species on earth, a concern shared by naturalists since the time of Linnaeus (1753 apud Forzza et al. 2010), was combined with the international committments made by the signatories to the CBD in addressing targets envisaged to reach the primary objectives of the convention the conservation of biological diversity, the sustainable use of its components and the fair and equitable sharing of the benefits arising from the utilization of genetic resources. The Global Strategy for Plant Conservation, designed to accelerate knowledge and conservation of plants within the framework of the CBD, defined 16 targets to be attained by The first of these targets was the preparation of a widely accessible working list of all known plant species, as a step towards a world flora (Nic Lughadha 2004). Fulfilling the committment assumed by Brazil through the GSPC in relation to the national list of species certainly provided a major motivating incentive for Brazilian taxonomists. To meet this challenge c. 413 taxonomists worked in a network from October 2008, and 21 May 2010 saw the online launch of Lista de espécies da flora do Brasil and the subsequent publication of Catálogo de fungos e plantas do Brasil (Forzza et al. 2010): covering 40,989 species of plants and fungi, of which 18,932 (46.2%) were reported to be species endemic to Brazil. Details on the development of the Brazilian List and comparative analyses of the different major taxa, levels of richness and endemism in Brazil and globally, are provided in Forzza et al. (2010, 2012). Having met the GSPC Target 1 in 2010, the team of taxonomists did not consider their census of Brazilian plant species complete. They envisaged the Brazilian List as a dynamic resource, bearing in mind that, in taxonomic science, new information Rodriguésia 66(4):

186 Flora of Brazil OnLine 2020 is discovered on a daily basis concerning species already described, not to mention the many species new to science still being discovered and described. The Brazilian List process continued from 2010 through to March 2015 with later versions of the online resource being greatly enriched with ecological data as well as updates on taxonomy and distribution. Results to date for all the groups of plants and fungi treated are available in BFG (2015), Costa & Peralta (2015), Maia et al. (2015), Menezes et al. (2015) and Prado et al. (2015). The same period saw the establishment by Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) of the Reflora Programme and the creation of the Reflora Virtual Herbarium (< The Brazilian List became an integral part of the Reflora programme and there was a large increase in the numbers of specimen images linked to vouchers for names. As well as the images originating from the databases of the herbarium of the Rio de Janeiro Botanical Garden (RB) and the Virtual Herbarium of Flora and Fungi (INCT), hundreds of thousands of images from the herbaria of the Royal Botanic Gardens, Kew (K) and the Muséum National d Histoire Naturelle in Paris (P) were also made available within the framework of Reflora. Subsequently, further new partnerships resulted in contributions from the following Brazilian and overseas herbaria, all of which are now incorporated in the Reflora framework: UFRN, EAC, ASE, ALCB, CEPEC, VIES, HUFU, ESA, CEN, UPCB, MBM, FLOR, HBR, HDCF, RON, MG, S, NY, MO and W. Encouraged by the success of the target setting approach, of which the GSPC was a pioneering example (Paton & Nic Lughadha, 2011), at its tenth meeting, in Nagoya, Japan in October 2010, the Conference of the Parties to the CBD adopted a revised and updated version of the GSPC, with new targets to be achieved by Where the original Target 1 had called for a widely accessible working list of all known plant species, as a step towards a world Flora, the revised Target 1 called for An online Flora of all known plants to be delivered by The technical rationale suggested that important steps towards this objective might include developing more complete synonymy; updating geographic distributions, inclusion of basic identification tools (keys, picture and basic descriptions) and inclusion of local and vernacular names where possible (Executive Secretary to the Convention on Biological Diversity 2010) It is by no means coincidental that several of the tasks undertaken between by the network of botanists committed to enhancing the Brazilian List are exactly those required to convert a Checklist into a Flora. As we write, in May 2015, levels of confidence and enthusiasm among Brazilian botanists at the prospect of completing a national Flora are manifestly high and groups of specialists are preparing plans to work together to tackle particular groups. While confidence and enthusiasm are valuable commodities at the outset of any project, they need to be balanced with a critical assessment of the resources available versus those that may be required and an evaluation of the lessons learned from similar endeavours in Brazil and elsewhere. This paper is intended as a contribution towards the enormous challenge of completing a Flora of Brazil by 2020 by considering (i) the experiences of the past seven years since the Brazilian List was initiated, (ii) the knowledge acquired and the resources mobilised in the preparation of the Brazilian List and (iii) the state of documentation of the flora and the human capital and taxonomic infrastructure required for the next phase. We do not attempt a comprehensive treatment of the sort that might be included in a feasibility study but rather address the following broad headline questions: Has a sufficiently high proportion of Brazilian species been described to justify attempting a national Flora? Are collection densities sufficient to underpin a national Flora? Is there sufficient botanical expertise available to develop such a big project over a five-year period? Are there current taxonomic treatments which support our understanding of the morphological variation of Brazilian plant species? Methods For each of the headline questions in our review we framed a series of more specific, narrowly-focused questions designed to address particular facets of the headline question. We then sought quantitative and/or qualitative data to address each of these specific questions. We focused primarily on angiosperms as they account for the vast majority of the known species in the Brazilian flora. For concepts and boundaries of biomes we followed IBGE Our analyses of Rodriguésia 66(4):

187 1118 Morim, M.P. & Nic Lughadha, E.M. angiosperm species in the Brazilian List were based on the results presented in BFG (2015). Proportion of flora already known Underlying our headline question Has a sufficiently high proportion of Brazilian species been described to justify attempting a national Flora?, we identified the following quantitative and qualitative questions: What proportion of Brazilian angiosperms is estimated to have been described/discovered to date? How many remain to be described? How many are already known to science but yet to be discovered within Brazilian borders? Will the ongoing description of new species disrupt or assist the development of the national Flora? We first derived estimates of the proportions of Brazilian angiosperm species described to date and awaiting discovery/description. We adapted the approach adopted by Pimm et al. (2010) which was based on partial lists of angiosperm species endemic to a single region of Brazil derived from the World Checklist of Selected Plant Families. We extended their approach to the more up-to-date and comprehensive species listing in the Brazilian List (BFG 2015) to obtain estimates of the number of angiosperms endemic to a single region awaiting discovery/description. To check the validity of our extrapolation we compared the relative numbers of species analysed by Pimm et al. (2010) for different regions of Brazil to equivalent numbers for angiosperms as a whole and for Apocynaceae, recently shown to have species richness distribution exceptionally highly correlated with the species richness distribution of Brazilian angiosperms as a whole (Pugliesi & Rapini 2015). We used the estimates of single-region endemic angiosperms awaiting discovery to predict the number of other additions to the flora, of species already known to science but not previously reported from Brazil. In light of results of parallel analyses (BFG 2015), we assumed that the proportion of endemic angiosperm species would remain relatively stable as knowledge of the flora increases i.e. that for every 100 species added to the flora, c. 57 would be Brazilian endemics while c. 43 would be species already known from other countries. If this proportion remains stable then the number of non-endemic additions to the flora could be predicted from the number of newly described endemics by applying the ratio 1:0.75. We used a sample of data from the International Plants Names Index (IPNI) to quantify recent rates of addition of species new to science described from Brazil from 2010 to 2014 inclusive and evaluate the likely impact of future additions on the preparation of the Flora over a five-year time-scale. Sampling sufficiency To address the headline question Are collection densities sufficient to underpin a national Flora?, we identified the following underlying questions: What collection density is considered sufficient? What is the current collection density? How does collection density vary across Brazil? We consulted 155 entries for Brazilian herbaria in Index Herbariorum (Thiers [continuously updated]). We also contacted curators of herbaria located beyond the borders of Brazil which are known to hold important collections of Brazilian material and obtained estimates of their total Brazilian holdings (BM, BR, K, M, MO, MPU, NY, US) as well as those of several smaller German herbaria (Hajo Esser pers comm.). We collated these data to obtain approximations of the numbers of specimens of Brazilian plants in the world s herbaria and the mean number of specimens per square kilometre. We compared these estimates with thresholds of collection density (numbers of herbarium specimens per 100 km 2 ) commonly cited for sampling sufficiency in tropical countries (Campbell 1989) and (Shepherd 2003), and with equivalent figures at global level and for other tropical countries where national flora initiatives are already in progress. We also compared the totals for Brazilian herbaria to equivalent figures reported by Sobral & Stehmann (2009) for 1990 and 2006 to provide an estimate of rates of growth in specimen numbers over the past 25 years. We analysed the totals for Brazilian herbaria by region to estimate collection densities for areas of scale more comparable to those of other national Flora projects and compared the resulting totals to equivalent data compiled from Index Herbariorum by Barbosa & Peixoto (2003). It is important to note that these regional analyses report numbers of specimens deposited in each major region, not all of which will have been collected in that region. Finally, to facilitate consideration of the results for Brazil in a global context we calculated the mean number of herbarium specimens per km 2 at global level using the estimate of 350 million specimens reported in Index Herbariorum and the Rodriguésia 66(4):

188 Flora of Brazil OnLine 2020 terrestrial surface of the earth at c. 149 million square kilometres. Taxonomic capacity Unpacking our headline question: Is there sufficient botanical expertise available to develop such a big project over a five-year period?, we identified the following underlying questions: Does sufficient relevant botanical expertise exist? Where is it located? What will determine its availability for this project? What competing priorities are there for botanists? To gauge the scale of existing botanical expertise of direct relevance to the Flora of Brazil we relied on the experience gained during the delivery of the Brazilian List in 2010 and subsequent enhancements (BFG 2015). We obtained data on the composition of the team that delivered this project and compared them with those analysed by Sobral & Stehmann (2009) on the publication of new species from Brazil over the period Finally, we undertook a more detailed analysis of some of the new species published from Brazil over the period to detect changes over time in the geographical location of taxonomic effort applied to the Brazilian flora. The sample used was developed for a larger study (Cowling et al. in prep.) and comprised 50% of the names reported in the IPNI as being published during each of two three-year intervals: and Within this sample we considered authors of new species described from Brazil over two 3-year periods: and For each of these names we determined the country in which the author was based when the work was undertaken (as reported in the protologue). We then compared the numbers of Brazil-based authors for the two time-periods. To evaluate these data in a global context we considered numbers of species and numbers of botanists for three major tropical floras, two in progress (Flora of Cameroon, Flora of Thailand) and one recently completed (Flora of Tropical East Africa). We calculated the number of species per botanist and, where applicable, number of species per botanist per year. Building on existing treatments To evaluate the extent to which the new national Flora can be developed based on existing treatments we first framed the rather broad 1119 question: Are there current taxonomic treatments, produced in the 21 st Century, which support our understanding of the morphological variation of Brazilian plant species? The specific underlying questions here are: How many taxonomic revisions of Brazilian plant families and/or genera have been produced? What is the coverage of Floras treating Brazilian species? What proportion of the species in the Brazilian List have been treated in a Brazilian state Flora? The Bibliographic Reference field of the Brazilian list (BFG 2015) was our main source of evidence for the production of taxonomic revisions and synopses and of Floras. For this first survey we focused on the years from 2000 to the present, a period during which major changes in the circumscription of taxa were frequent as a result of molecular and phylogenetic studies. We counted complete taxonomic revisions of families, partial revisions (e.g. of sections of genera, tribes and subfamilies) and synopses. Based on the compiled information we estimated the number of species treated in these revisions by consulting the published article or the author. Where neither of these approaches was feasible (chiefly in the case of doctoral theses and masters dissertations as yet unpublished) we assumed that the number of species treated in the revision was the same as the number of species recognised in Brazilian List. Based on the survey we chose five Floras to illustrate the extent of knowledge of Brazilian plants at different geographical scales and in different Brazilian biomes. Quantitative data on the species covered to date by each of these Floras were obtained as follows. For the monographic series Flora Neotropica, we consulted monographs published for angiosperm taxa known to occur in Brazil and checked the Numerical List of Taxa to obtain the total number of species treated (Organization for Flora Neotropica Monograph ). Quantitative data on Flora Neotropica published from 2010 to 2014 inclusive were supplied by the editor (Thomas pers. comm.). For each taxon that had been monographed, the corresponding number of Brazilian species was obtained from the BFG (2015). For the Flora Ilustrada Catarinense (Reitz ; Reis ) we consulted the CONSPETO GERAL (General Conspectus) for each of the volumes online at Enciclopédia Flora Ilustrada Catarinense (< com.br/news/fic/>) and we selected the families Rodriguésia 66(4):

189 1120 Morim, M.P. & Nic Lughadha, E.M. of angiosperms native to Brazil. In the case of the Flora Fanerogâmica de São Paulo we obtained quantitative data on families, genera and species from Wanderley et al. (2011). Similarly, for Flora of Sergipe, quantitative information were obtained from Prata et al. (2013). Finally for the Flora of the Distrito Federal, quantitative information on treatments published to date were obtained from the editors (Cavalcanti & Ramos 2001; Andrielli C. A. Lopes pers.comm.). In order to evaluate the extent of overlap in coverage of species between these State Floras, we conducted pairwise comparisons of species lists for three plant families each of which had been treated in more than one of the Floras. For this purpose we chose two families which we expected to represent the extremes of variation in overlap and one which we expected might be broadly representative of angiosperms more generally. We selected Myrtaceae, a family well known for having many species with restricted distributions in Mata Atlântica (Murray-Smith et al. 2009), Poaceae which includes many widespread species (Longhi-Wagner et al. 2001), and Apocynaceae which was recently shown to have species richness distribution exceptionally highly correlated with the species richness distribution of Brazilian angiosperms as a whole (Pugliesi & Rapini 2015). For each comparison, we summed the number of species reported for each state (gross total) and subtracted the number of species reported as occurring in both states (net total). Dividing net by gross totals gives an indication of the amount of overlap in species between state Flora treatments of the same family, with a value close to 1 indicating little overlap while values close to 0.5 would reflect susbtantial overlap. Results Proportion of flora already known Consulting the Brazilian List we noted almost 12,000 species (11,973) which were reported to be endemic to one of the five major administrative regions of Brazil. Numbers of single-region endemics ranged from just over 1,000 in the South to 5,690 in the South East (Tab. 1). The numbers of single region endemics considered by Pimm et al. (2010) were very highly correlated with those reported by the Brazilian List for Apocynaceae (r 2 = 0.99) and for angiosperms (r 2 = 0.99). Applying the percentage increases per region predicted by the Pimm et al. (2010) model to these more comprehensive numbers for regional endemics yielded estimates for single-region endemics awaiting discovery/description ranging from 246 species for Centre-West and South regions to 1,179 for Northeast Brazil. Summing these regional estimates gave a national total of 2,618 angiosperm species endemic to a single region and still awaiting discovery/description (Tab.1). Assuming the proportions of endemic and non-endemic elements of the flora remain stable over time, an assumption broadly in line with results reported in parallel analyses (BFG 2015), then we can predict that the number of non-endemic species awaiting discovery in Brazil is c. 1,963 species. Summing predicted additions of singleregion endemics and non-endemic species gives a total predicted growth in the flora of 4,581 species, equivalent to a 14% increase in species numbers against the baseline total of 32,086 angiosperm species reported in BFG (2015). In other words our analysis suggests a total angiosperm diversity in Brazil of 36,666 species of which 88% are already known to science and recorded from Brazil. Records of species described as new to science with types from Brazil totalled 1,191 over the five-year period , thus averaging 238 per annum (see BFG 2015 for further details). Sampling sufficiency Of the 155 Brazilian herbaria for which entries were consulted, 137 are current repositories for Brazilian specimens. The total number of specimens reported to Index Herbariorum by these herbaria was 6,697,000 (hereafter rounded to 6.7 million, Fig.1). This figure includes material collected in countries other than Brazil but deposited in Brazilian herbaria. A breakdown by country was not possible, but our experience is that non-brazilian material contributes only a very small proportion of the total material. For example, in RB, one of the most international of Brazil s herbaria, only 6% of the material is from countries other than Brazil (Luis Alexandre Estevão da Silva pers. comm.) and much lower proportions of non-brazilian material would be typical of most Brazilian herbaria (Forzza pers. comm.) While a figure for total holdings in Brazilian herbaria certainly overestimates holdings collected in Brazil, this may be offset in part by the fact that many Brazilian herbaria do not update their entries in Index Herbariorum with great regularity, thus published figures for herbarium holdings inevitably Rodriguésia 66(4):

190 Flora of Brazil OnLine 2020 lag behind actual figures. In the case of the rapidly growing herbarium RB, some 130,000 specimens had been added to the collection since the last update, a figure we added to our estimate, giving a total of 6.8 million for use in collection density estimates (Tab. 3). Analysis of these Brazilian repositories by region showed a very skewed distribution (Tab. 2), with the vast majority of the collections deposited in the South and Southeast and little change between 2003 and 2015 in terms of the regional distribution of the collections. The combined estimates of the Brazilian holdings in European herbaria for which we could obtain data exceeded 1.3 million while the collections of just three major US herbaria together account for a further million specimens (Tab. 3). Combining these estimates we obtained a total of 9.2 million Brazilian specimens, equivalent to 1.08 specimens per square kilometer of Brazil s 8,515,767 km 2 extent. The calculated global mean of 2.4 specimens per km 2, exceeds the threshold of one specimen per km 2 proposed by Campbell (1989) and approaches that of three specimens per km 2 suggested by Shepherd (2003). Equivalent values for other tropical countries with Flora projects in progress were estimated at 0.82 for Cameroon and calculated for Thailand at 0.96 (See Tab. 4 for sources). It is important to note that all of the above estimates are for numbers of specimens (typically comprising a single herbarium sheet) rather than for numbers of collections (which typically comprise several specimens, usually distributed to several different herbaria and thereafter counted as separate specimens). This approach is necessary in Figure 1 Change over time in total numbers of specimens reported in Index Herbariorum as deposited in Brazilian Herbaria order to facilitate comparison with earlier studies and due to the limitations of the data available. Taxonomic capacity A total of 413 collaborators were involved in the preparation of the Brazilian List (Thomas et al. 2012). Of these, 413 scientists, the vast majority (86%), were reported to be Brazilian. A similar number of scientists, 483 in total, were reported by Stehmann & Sobral (2009) as having been involved in the description of new species from Brazil over the period , but just 43% (209) of these authors were reported to be Brazil-based (Tab. 5). More detailed analysis of 50% samples of IPNI data for and showed that four times as many Brazilian species new to science were described by Brazil-based botanists over the later time period than in the earlier one. There was also a marked increase in the number of Brazil-based botanists participating in the process of description of new species, with more than three times as many individuals involved in the publication of at least one species new to Brazil during as were involved in (Fig. 2). Comparing species numbers, timescales and human resources for the Brazilian flora with those for other tropical flora projects in progress or completed (Tab. 6) indicated that the number of species to be treated each year would be several times higher than the rate recently proposed for another project considered ambitious in timescale (Flora of Thailand) and many times higher than the annual rates achieved for the only tropical Flora of > 10,000 species completed this century (Flora of Tropical East Africa). However, when the scale of the trained taxonomic capacity potentially available is factored in we used the figure for the collaborators involved in the most recent round of enhancements to the Brazilian List (BFG 2015) - the numbers of species treatments to be completed per person per year are just twice those projected for Flora of Thailand. Building on existing treatments For the period under review (2000-early 2015) authors of the Brazilian List cited c. 168 taxonomic revisions at diverse taxonomic ranks. Notably, these include seven revisions at family level, treating all known Brazilian species of, respectively, Apodanthaceae, Balanophoraceae, Canellaceae, Cannaceae, Caryophyllaceae, Heliconiaceae and Rhamnaceae. Included in this Rodriguésia 66(4):

191 1122 Morim, M.P. & Nic Lughadha, E.M. Table 1 Estimating proportions of Brazilian angiosperms described vs yet to be discovered/described. Columns A-D show results presented by Pimm et al Columns E-H show equivalent figures based on LEFB Columns J-M provide projections for non-endemic additions to the angiosperm flora and total additions to angiosperm flora based on assumptions outlined in text. Original results based on WCSPF (as presented in Pimm et al. 2010) Results of reanalysis based on current Brazilian List (BFG) 2015) Brazilian region Single-region Endemic species in WCSP 2008 fide Pimm Predicted species totals (incl. those yet to be discovered) fide Pimm Regional endemics awaiting description fide Pimm-WCP Predicted % increase Endemic angiosperms fide Brazilian List Endemics as a proportion of WCP 2008 endemics Predicted regional totals for single-region endemics Regional endemics awaiting discovery/ description extrapolating WCP 2008 to Brazilian List data Predicted non-endemic additions to Flora Predicted total additions to Flora (endemic and not) Predicted total angiosperms % of Flora described to date A B C=B-A D = B-A)/A E F = E/A G =(D/100+1) E H =G-E J =H*0.75 K =H+J L =K M =32086/L*100 North , , Northeast 1,016 1, , ,610 1,194 Central-West , , Southeast 3,485 3, , , South , , Totals 6,382 7,645 1, ,973 14,591 2,618 1,963 4,581 36, Table 2 Numbers of specimens deposited in herbaria in each region of Brazil as reported in Index Herbariorum. IH data from 2015* IH data from 2003** Brazilian region Number of specimens (1000s) % of specimens Collection density (specimens/km 2 ) Number of specimens (1000s) % of specimens Collection density (specimens/km 2 ) North Northeast Central-West Southeast 3, , South 1, Totals 6,930 5,138 *Thiers, continuously updated, consulted May 2015 **Barbosa & Peixoto 2003 Rodriguésia 66(4):

192 Flora of Brazil OnLine 2020 large set of recent revisions are several theses with submission dates of 2005 or later. Collectively the revisions treat c. 2,186 Brazilian species. The authors included 1,341 references to Flora treatments (including multiple citations of the same Flora treatment). Although the data are not available in a suitable format to derive numbers of different Flora treatments cited, nor the number of species they treat, they do provide a good overview of the differing geographic scopes, of the diversity of Floras and of the qualitative information they offer about Brazilian species, which we address further in our Discussion, along with the quantitative results presented below. Among the Floras cited by the Brazilian List taxonomists, one of the most cited is that with the greatest geographical scope, the Flora Neotropica monograph series, which has been published since 1967, and documents the great richness and diversity of the plants of the different vegetation formations of the Neotropics. Of the 114 Flora Neotropica volumes published to date, 84 treat angiosperm families or genera and collectively these cover a total of c. 7,532 species. The vast majority (81 of the 84 monographs) include angiosperm species which occur in Brazil. Over the period , a total of 23 Flora Neotropica monographs were published, treating 1,555 species (Tab. 7). These monographic treatments cover families and genera for which c. 831 species are now reported from Brazil. Although some of these species will have been described as new or reported for Brazil after the publication of the monograph, for the vast majority of the 831 Brazilian species in these groups a high quality description will be available, either in the monograph or in subsequent papers, often by members of the same research team (e.g. Knapp et al. 2015). The range of Brazilian Floras cited as supporting references in the Brazilian List ranges from the classic Flora Brasiliensis (Martius et al. 1833; Urban 1906) to Floras at state and local level which were prepared over the course of the 20th Century and continue to be produced to the present day. The results of our partial evaluation of Floras for three Brazilian states and for the Distrito Federal are summarised below. Flora Ilustrada Catarinense (Illustrated Flora of Santa Catarina) Initiated and edited by Raulino Reitz and subsequently Ademir Reis (Reitz ; Reis ), this work comprises 142 fascicles, 1123 each treating a single family (or part of a larger family) with the most recent being published in Of the 132 angiosperm families treated, eight are represented only by species in cultivation. Ten families of pteridophytes s.l. have been published. A total of 4,228 species have been treated, of which 3915 are native species of angiosperm (Reitz ; Reis ). The Flora of Santa Catarina is the most complete Flora for a Brazilian State, the number of species treated to date being equivalent to c. 82% of the angiosperm species recorded for Santa Catarina (BFG 2015). In Santa Catarina the predominant vegetation formations are Dense Ombrophilous Forest, Araucaria Forest and Campos. Flora Fanerogâmica de São Paulo (Phanerogamic Flora of São Paulo) Initiated in 2001, this state Flora comprises seven volumes to date, with the most recent having been published in In total, 151 families have been treated, of which two are gymnosperm families. A total of 3,237 species have been published (Wanderley et al. 2011), equivalent to c. 43% of the number of angiosperm species reported for São Paulo (BFG 2015). The vegetation of the State of São Paulo is quite diverse, with the dominant vegetation formations being Dense Ombrophilous Forest, Deciduous or Semi-deciduous Forest, High Altitude Grassland and Cerrado (Wanderley et al. 2011). Flora do Distrito Federal The Flora of the Distrito Federal was also initiated in 2001 (Cavalcanti & Ramos 2001) with the most recent volume having been published in To date, this work has treated 88 plant families: 70 Seed Plant families and 18 Pteridophyte families and a total of 974 species, of which 860 are Seed Plants (Andrielli C. A. Lopes pers. comm.). This coverage equates to c. 27% of Seed Plants reported for Distrito Federal (BFG 2015) The vegetation of the Distrito Federal is predominantly Cerrado s.l. (Cavalcanti & Ramos 2001). Flora de Sergipe This state Flora was initiated in 2013 and, to date, has published one volume treating 37 angiosperm families in which 494 species are reported and described (Prata et al. 2013), equivalent to c. 31% of those reported for Sergipe (BFG 2015). The most prevalent vegetation types in Sergipe are Humid Forest, Atlantic Forest and Caatinga (Prata et al. 2013). Rodriguésia 66(4):

193 1124 Morim, M.P. & Nic Lughadha, E.M. Table 3 Estimated numbers of Brazilian specimens deposited in Brazil and in major herbaria in Europe and the United States. Brazilian Herbaria (fide IH) 6,697,300 Additions to RB since last update to IH* 130,000 Total for Brazilian herbaria 6,827,300 Major European Herbaria NHM London (BM) 300,000 RBG Kew (K) 300,000 MNHN Paris (P) 420,000 Munich (M) 70,000 Other German herbaria 60,000 Montpellier (MPU) 50,000 Brussels (BR) 110,000 Total for European Herbaria 1,310,000 Major USA herbaria New York Botanical Garden (NYBG) 650,000 Missouri Botanical Garden (MO) 208,097 Smithsonian Institution (US) 200,000 Total for USA Herbaria 1,058,097 Grand Total 9,195,397 Area of Brazil in sq km 8,514,877 Specimens per sq km 1.08 * Forzza pers comm. Table 4 Comparison of sampling sufficiency estimates for Brazil with those of other tropical Flora projects in progress. Area (km 2 ) Specimen numbers Collection Density (Specimens/km 2 ) Source of specimen numbers Earth 149,000, ,000, Thiers (continuously updated, consulted May 2015) Brazil 8,514,877 9,195, This paper Cameroon 427, , G. Gosline pers. comm. Thailand 515, , Thiers (continuously updated, consulted May 2015) Table 5 Scale and distribution of taxonomic capacity relevant to Brazil. Description of taxonomic activity Based in Brazil Based elsewhere Total Brazilian List Collaborators* Authors of new species with types from Brazil ** * Thomas et al ** Sobral & Stehmann 2009 Rodriguésia 66(4):

194 Flora of Brazil OnLine Table 6 Comparisons of scale of task, timescale and human resources for tropical flora projects. Flora project (proposed, in progress or completed) Species Botanists involved Years since start (or total years in the case of completed project) Species treated Species treated per project year Species yet to be treated Years projected to completion Flora of Brazil 36, N/A N/A N/A 36, , Flore du Cameroun 7,850 20? Flora of Thailand 10 12, , , ,071 8 Flora of Tropical East Africa 12, , N/A N/A N/A 2* Species to treat per year Species per botanist per year Table 7 Numbers of Flora Neotropica monographs and angiosperm species treated therein over two time periods Monographs number Species number Monographs number Species number 84 7, ,555 Table 8 State Floras and species number. State Flora Species number Flora Ilustrada Catarinense 3,915 Flora Fanerogâmica de São Paulo 3,237 Flora Distrito Federal 860 Flora de Sergipe 494 Total 8,506 Rodriguésia 66(4):

195 1126 Morim, M.P. & Nic Lughadha, E.M. Table 9 Taxonomic treatments and species number. Treatments of species Species number Taxonomic revisions 2,282 Brazilian Floras 6,700 Total 8,982 Figure 2 Growth in publication of new species by Brazilbased authors. (Based on data from Cowling et al. in prep.). Angiosperms and Gymnosperms (BFG 2015) 32,109 % Treated species 28% Collectively, the four Floras analysed include c. 8,506 species treatments (Tab. 8), a total equivalent to c. 33.5% of the angiosperm and gymnosperms species reported for Brazil (BFG 2015). It must be borne in mind however that this figure does not represent 8,506 different species, since a significant number of these species are not confined to a single state and thus may have been treated in more than one Flora. Our pairwise comparisons for selected families showed that the extent of overlap in species between Floras varied widely, with unique species as a proportion of total species treatments ranging from 0.95 for Myrtaceae in Santa Catarina and Sergipe to 0.68 for Poaceae in Santa Catarina and São Paulo. Apocynaceae yielded an intermediate value of 0.72 for Santa Catarina and São Paulo. To extrapolate directly from these results would overstate the degree of overlap in species between these Flora treatments since the pairwise comparisons could only be made between families which had been treated for two Floras. For example there are c. 42 families which have been treated for the Flora of Santa Catarina but not yet for the Flora of São Paulo and collectively these account for 2,056 species. For purposes of estimation we assumed that these 2,056 species were not treated elsewhere and that all the other species treatments in state Floras had levels of overlap equivalent to those reported here for Apocynaceae, so that 6,450 species treatments might represent 4,644 different species. Summing these Flora totals (2, ,644 ) with those reported above for monographs (2,282 species) gives a total of 8,982 species (Tab.9). Therefore, for at least 28% of angiosperms and gymnosperms (8,982 of the 32,109 species native to Brazil) modern descriptions are available which might form the basis for an online Flora treatment. Discussion Proportion of flora already known Our results suggest that as many as 88% of all Brazilian angiosperm species may already have been discovered and described. This figure is much higher than estimated for the neotropics as a whole (Thomas 2005) but broadly consistent with global level analyses (Joppa et al a, b) which is scarcely surprising, since it is derived from the same model applied in a Brazilian context (Pimm et al. 2010). The credibility of this result depends to a large extent on whether the assumptions inherent in the model and analyses are valid in a Brazilian context. Hence, we evaluate the assumptions underlying the original analyses and the extrapolation presented above before attempting to discuss the broader significance of the result. Apart from rates of description of species over time, the two key factors involved in the model presented by Joppa et al. (2011a) are (i) taxonomic effort, defined as the number of taxonomists involved in describing species, and (ii) taxonomic efficiency, defined as the number of species described per taxonomist adjusted for the continually diminishing pool of unknown species. The logic underpinning the model is that taxonomic effort is a powerful predictor of the number of species described and if numbers of species described per individual scientist involved in species description are declining over time, that decline must be attributable to a shrinking pool of species yet to be discovered/described. Joppa et al. (2011a) found that at global level, for many plant taxa there was an increase in taxonomic efficiency over the course of a century or more, followed by a decline which they attributed to the declining pool of species as-yet-unknown. In a Brazilian context, Rodriguésia 66(4):

196 Flora of Brazil OnLine 2020 Pimm et al. (2010) considered species endemic to each region of Brazil and reported similar patterns of increase then decrease in taxonomic efficiency, with the shape of the curve differing from region to region, resulting in widely varying predictions of proportions of plant species yet to be described from each region (from 9% in Southeast Brazil to 49% in Northeast Brazil). To what extent are these model assumptions defensible in light of the data available today? The results reported under Taxonomic Capacity above are certainly compatible with the assumption that, in broad terms, taxonomic effort is a powerful predictor of number of species described, with a four-fold increase in species described associated with a > three-fold increase in botanists involved. However, the data do not provide support for the idea of a decline in taxonomic efficiency, rather this limited dataset is suggestive of increased taxonomic efficiency when the later period studied is compared to the earlier, an anomaly which has prompted a more detailed treatment elsewhere (Nic Lughadha, Morim in prep.). Beyond the assumptions built into the Pimm et al. (2010) model, it is important to consider assumptions implicit in the selection of the datasets to which the model is applied. For both global and Brazilian analyses, the dataset comprised species lists for all monocots and selected non-monocot families (sourced from the World Checklist of Selected Plant Families). Thus monocots were over-represented, accounting for 58% of all the species in the global analysis although only one in five angiosperms are monocots (R. Govaerts, pers. comm. 2015). Exact figures were not supplied for the Brazilian analysis but since c. 27% of Brazilian angiosperms are monocots (Forzza et al. 2010) and all known monocots were included in the list from which the sample was drawn they were certainly over-represented among the 6,382 angiosperm species analysed by Pimm et al (Tab. 1). There was an implicit assumption in both global and Brazilian studies that, despite being dominated by monocots, the datasets analysed were broadly representative of angiosperm diversity at global and national level respectively. This latter assumption is arguably the most important, since the inferences to be drawn from our extrapolation depend directly on it. A robust test of the validity of extrapolating from the monocot-dominated dataset to angiosperms more broadly is beyond the scope of this review 1127 and the fact that Brazilian monocot species have slightly higher levels of endemism than Brazilian angiosperms as a whole (60% vs 57%, fide Forzza et al. 2010) might be considered cause for concern. However, the strong correlations we found between numbers of endemic species analysed by Pimm et al. (2010) and numbers of Apocynaceae and angiosperms respectively endemic to these regions lend some support to the idea that patterns of species richness may be similar. In summary, although not all model assumptions can be fully explored, most appear credible enough to suggest that our results extrapolated from the Pimm et al. (2010) modelled results merit consideration. What are the implications of these results for the preparation of a Flora of Brazil? Our overall estimate that c. 88% of Brazilian species have already been discovered and described is encouraging, as is the result that almost half the of the predicted additions are likely to be already known from other countries suggesting that those undertaking treatments of particular groups for the Flora will not be overwhelmed by an endless stream of novelties to be described as new to science. However, the simple fact that the great majority of Brazilian plant species are already known to science should not lead us to underestimate the work involved in treating them effectively in an online Flora, including, among others, documenting their morphological variation throughout their known range, addressing gaps in knowledge of their distribution (e.g. through new collections) and their circumscription (e.g. through phylogenetic and phylogeographic studies). Our indicative results on the regional distribution of species new to science are arguably more useful for those planning the Flora: they suggest that of the 2,618 species yet to be discovered and described as new to science, close to half (1,194=46%) are confined to Northeast Brazil. Conversely, we can predict that most of the projected 1,963 additions to the Flora of species already known from other countries will not be found in Northeast Brazil but in regions bordering other South American countries. For example, in their analysis of five years of additions to the flora of Acre, Medeiros et al. (2014) reported 34 additions to the flora of Brazil of which most (23 = 67%) had previously been described from other countries, while just 11 (33%) were new to science (including some species still awaiting publication). Rodriguésia 66(4):

197 1128 Morim, M.P. & Nic Lughadha, E.M. Sampling sufficiency Comparing our 2015 estimate of 6.7 million specimens reported by Index Herbariorum as deposited in Brazilian herbaria to equivalent figures of 3 million and 4.9 million for 1990 and 2006 respectively (Sobral & Stehmann 2009), indicates significant growth over the past 25 years in numbers of specimens deposited in Brazilian herbaria (Fig 1). The data also suggest an acceleration in growth rate in the past decade compared to the previous 15 years. However, since the intervals at which herbaria provide updates to Index Herbariorum on their accession numbers are variable and often long, caution must be exercised in interpreting the data in terms of trends over time. Combining our collation of published information for Brazilian herbarium holdings with estimates from curators of herbaria outside Brazil which hold important collections of Brazilian material yielded an estimate of 9.2 million specimens. We consider this estimate to be conservative for several reasons: (i) delayed reporting of Brazilian herbarium holdings to Index Herbariorum (as illustrated under Results); (ii) we approached only a very limited selection of the non-brazilian herbaria with important deposits of material collected in Brazil, so holdings outside Brazil are certainly underrepresented in our sample. Balancing these factors tending to under-estimation is the fact that the figures supplied (i) include non-brazilian material deposited in Brazilian herbaria, (as outlined under Results) and (ii) are generally not confined to angiosperm holdings but may include other groups including non-vascular plants, algae and fungi. While specimens other than of vascular plants are in a minority in most herbaria they represent an important proportion in some cases (e.g. NY) or even the majority of the collection in several instances (e.g. URM at the Federal University of Pernambuco specialises in tropical and temperate fungi and lichens) so that a total restricted to angiosperms might sum to < 8 million specimens. Our estimates suggest that numbers of specimens exceed the arbitrary threshold value of 100 herbarium specimens per 100 km 2 suggested by Campbell (1989) as being the minimum necessary to ensure that a botanical inventory has sampled most of the taxa and gathered sufficient distributional information. Applying this criterion, Brazil considered as a whole would be deemed to have reached sampling sufficiency, a situation which was not envisaged by Campbell (1989) who, based on reported increase in collections over the period , projected a 51 year interval before Brazil would reach such collection densities. Collecting densities in Brazil are now broadly comparable to or slightly in excess of those estimated for Cameroon and Thailand, where Flora projects are ongoing. Shepherd (2003) deemed Campbell s arbitrary threshold conservative and suggested that three collections per km 2 would be more appropriate. While many tropical botanists (including the present authors) might agree with Shepherd in principle, it would be unrealistic in practice to refrain from embarking on a Flora project until this ambitious target was met, as our results suggest that it has not yet been met for most parts of the world, given our calculated global mean of 2.4 specimens per km 2. In fact, the global mean is not very meaningful as it masks the extreme disparities between the extraordinarily high collection densities in many temperate areas and the many areas of the world which would still be considered poorly collected in Campbell s terms (see below for definition). The same can be said for Brazil, given the vast extent of the territory to be considered and the distributions of herbaria and of collecting effort, both strongly skewed to the South and Southeast regions. Our results indicate that despite an apparent increase in the rate of growth of the collections, coupled with increased awareness of the need for additional collections in the North and Northeast (e.g. Sobral and Stehmann 2009; Medeiros et al. 2014; Moro et al. 2014) there has been little or no change in the proportions of the collections deposited in the major regions of Brazil over the past decade or more (Tab. 2). Values at regional level are perhaps the most relevant for addressing the question as to whether collection densities are sufficient to support the development of a Flora, though they need to be interpreted carefully in light of the fact that they only represent material deposited in Brazil. At this scale the contrast between the South and Southeast and the remainder of Brazil is stark. Collection densities for the South are far in excess of Campbell s (1989) threshold and equal the global mean of 2.4 specimens per km 2. Herbaria in the Southeast hold c. 47% of all the herbarium specimens deposited in Brazil and include greater numbers of specimens collected from other regions of Brazil than is the case for herbaria based in other regions. Therefore the calculated collection density of 3.4 specimens per km 2 probably over- Rodriguésia 66(4):

198 Flora of Brazil OnLine 2020 represents the level of sampling for the Southeast. Nonetheless, there can be no doubt that sampling in the South and Southeast is sufficient to support production of state Floras, since significant progress has been made in the publication of Floras of São Paulo and Santa Catarina, as described earlier in this review. For the remainder of Brazil, Shepherd s threshold of 3 specimens per km 2 is far above the reality and our discussion is probably best framed in terms of the additional criteria proposed by Campbell (1989) who introduced the terms relatively well-collected for areas with more than 50 specimens per 100 km 2 and poorly collected for those with fewer than 50 specimens per 100 km 2. Thus the Centre-West and North regions,which have collection densities of 0.4 and 0.2 specimens per km 2 respectively, judged only on collections deposited within their own region, would qualify as poorly collected in Campbell s terms. When collections deposited outside the region are considered, the Centre-West would likely qualify as relatively wellcollected. In contrast, even allowing for the major holdings from this region deposited in other regions of Brazil (e.g. > 60,000 at RB), and overseas (e.g c. 60,000 at K and > 80,000 at NY) the North is barely halfway to the threshold for being deemed relatively well-collected. The historical, economic and sociological factors underpinning this sampling deficit are discussed in detail by Sobral & Stehmann (2009), as are some of the implications in terms of basic understanding of Amazonian plant diversity and its conservation status. While there are compelling arguments for increased investment in documenting the Amazon flora the immediate question to be addressed is: should the sampling deficit be considered so severe as to render attempts at a Flora unviable? We think not. Utility must be the primary consideration in evaluating whether a project is worthwhile. A Flora that facilitated identification of the great majority of the species most likely to be encountered in the Brazilian Amazon would surely be recognised as an important and useful contribution, even if many rarer species received only tentative or provisional treatments. Recent insights into the distribution of Amazon tree diversity has shown that just 5,000 tree species account for > 99% of the trees in the Amazon forests as a whole (ter Steege et al. 2013). A Flora that treated these in some detail, complemented by preliminary treatments for other species less frequently encountered, would 1129 provide a very useful foundation which could be improved over time. In fact, this pragmatic approach highlights an important difference between Flora projects initiated in the 21 st Century and those with longer histories. The discussions in the literature as to levels of collection density which are sufficient to support a Flora date, for the most part, from a time when hard copy publication was the norm, with Floras being published in fascicles as families were completed or in large volumes encompassing several families (e.g. Prance 1977). The primary disadvantage of publishing a Flora treatment based on insufficient collections was that the publication would soon be rendered out of date by the discovery of new species and range extensions for known species, necessitating the production of supplements or second editions at significant cost in time and money. In contrast, the online approach adopted in the development of the Flora of Brazil renders low collection densities a much less troubling problem. As new collections become available, revealing new distributions or even new species, the family treatments can be updated with the latest information. The collection density of 0.6 per km 2 estimated for NE Brazil indicates that it qualifies as relatively well collected in Campbell s terms. At first glance, this finding may appear surprising in light of results presented earlier (Pimm et al and this paper) suggesting that the number of endemic species yet to be discovered/described from NE Brazil is similar to the number already known. However, the two conclusions are not necessarily contradictory. Even relatively high overall collection densities can result in poor representation of the flora when the distribution of collections is very uneven, as is the case for NE Brazil. The effects of uneven collection may be amplified by the fact many NE Brazilian species may have more restricted distributions than species of the cerrado, for example (Moro et al. 2014). We found no evidence to suggest that inadequate sampling sufficiency would provide such a severe impediment that it would be inadvisable to initiate a national Flora at this stage. However, it is important to note that while Campbell s approach and threshold values provide a useful starting point for discussion and for comparison between countries they are very much a product of their time and a more sophisticated and multi-faceted consideration of the availability of specimens for the production of a Flora will Rodriguésia 66(4):

199 1130 Morim, M.P. & Nic Lughadha, E.M. likely prove more useful going forward. Two aspects of Campbell s metric appear to have been driven by pragmatism rather than best practice. Firstly, he considered specimens (typically single sheets), rather than collections which typically comprise multiple sheets prepared from single gathering bearing the same collector number, which are later separated and distributed to different herbaria, so that the number of sheets deposited in herbaria may exceed the number of collections made by five-fold or more. Secondly, Campbell considered only those sheets deposited in-country because these were the data readily available via Index Herbariorum. Sobral & Stehmann (2009) followed the same approach for similar reasons, reporting that numbers of specimens of Brazilian plants in foreign herbaria were difficult to evaluate. The advent of the Reflora Programme, and its current focus on digital repatriation of Brazilian herbarium specimens has resulted in rapid change in this landscape, as scientists in major European and US herbaria prepared estimates of the size of their Brazilian holdings in preparation for their digitisation. Thus, estimating numbers of sheets of Brazilian collections deposited outside Brazil is a much easier task in 2015 than in Once these specimen data are digitised and accessible via a single interface, true estimates of collection densities (as distinct from specimen densities) should be achievable. These estimates, combined with the tools now available within Lacunas (Canhos et al. 2013) to evaluate numbers of collections per species, will provide much more meaningful and refined insights into the areas and the taxa for which lack of collections represents a major impediment to the production of the Flora of Brazil. Taxonomic capacity Is there sufficient botanical expertise available to develop such a big project over a five-year period? Our results showed that the annual rate of production of species treatments required to complete a Brazilian flora by 2020 was ambitious, when compared to tropical Flora projects which were ongoing or recently completed, but that the taxonomic capacity recently engaged on the enhancement of the Brazilian List which could potentially be mobilised for future work, was proportionate to the scale of the challenge. In other words, the relevant expertise clearly exists in sufficient quantity, 437 collaborators fide BFG (2015), but there remains a question as to whether it can be mobilised and focused sufficiently on the delivery of the Flora. Twenty or even ten years ago, few taxonomists might have declined the opportunity to make a substantial contribution to a Flora of this importance. It remains to be seen whether this is still the case, despite increasing focus by agencies co-ordinating scientific research and higher education on metrics based on impact factor of journal publications, citations, and grantfunding secured. The fact is that many of those who have the knowledge and experience to produce authoritative, high-quality taxonomic treatments for the Flora at the pace and scale required to meet the 2020 Target may also be considering the potential impact of this commitment on their career prospects. The ageing profile of the taxonomic specialists willing to tackle large groups has been noted by Medeiros et al. (2014) who also reported a shortage of younger taxonomists willing to take their places. Increased availability of data in digital form coupled with user-friendly interfaces to streamline the preparation of species treatments may lighten the burden of the task for taxonomists but it may prove equally important to ensure that the importance of their contributions to science and conservation in Brazil are recognised at the highest level and that this recognition is incorporated into the systems and metrics by which scientists are evaluated and rewarded. Building on existing treatments The numbers of species treatments reported in our (partial) survey of taxonomic revisions, State Floras and Distrito Federal Flora and the proportion of Brazilian angiosperm species which we estimate that they encompass (28% Tab.9) might not, in principle, seem very encouraging, since they immediately prompt further questions, perhaps the most important being: Are there taxonomic treatments available for the remainder (the majority) of the angiosperm species known from Brazil, other than in their original place of publication? Although we do not have a complete survey of all taxonomic treatments, with totals for different species treated therein, our review of the data for the period in the Bibliographic Reference field of the Brazilian List provides a qualitative overview of the taxonomic treatments available for the angiosperm flora of Brazil. Beyond the classic Rodriguésia 66(4):

200 Flora of Brazil OnLine th Century works, various taxonomic revisions were produced during the 20 th Century, both by Brazilian botanists and others. Despite changes in nomenclature and taxonomic circumscription in the interim, as well as the addition of newly described species, works such as those compiled for Orchidaceae (Pabst & Dungs 1977) or for the genus Mimosa (Barneby 1991) represent invaluable syntheses of information on the morphology of species which will be important resources for the preparation of Flora do Brasil There are also many important treatments of groups above the level of species focused on a particular ecosystem or politically-defined area which have not been incorporated into a formal state Flora (e.g. Alves- Araújo et al. 2009; Barros & Morim 2014; Esteves 2001; Queiroz 2009; Soares et al. 2014). Such treatments were not sampled quantitatively in this review, though there are many (BFG 2015). Thus our estimate of the numbers of relevant species treatments available is very conservative. We have emphasized the importance of existing taxonomic treatments for the Brazilian List but of course, such benefits accrue in the other direction also: the Brazilian List has played an important role in the updating of taxonomic revisions. Knapp et al. (2015) comments that following the revision of the Solanum havanense species group (section Geminata (G. Don), in which 35 species were reported for Brazil, nomenclatural updates and information made available about the Geminata clade of Solanum in the Brazilian List gave rise to the recognition of a further nine species records for Brazil, as well as updating the information on the diversity and distribution of species of this clade. Beyond the Flora Neotropica series and the four Brazilian Floras included in our results, there is a considerable number of studies focused on local Floras, some with a focus on Protected Areas. In general, neither state Floras nor local Floras have a regular publication schedule and many remain unfinished. Nonetheless, as for the other sources cited, they provide collated information on Brazilian species, at more restricted geographical scales. For regions such as the Amazon, these Floras are all the more important because they provide some of the few contemporary sources on the flora of the region. The series of articles concerning Flora of the Reserva Ducke (Manaus, Amazonas) represent a good example of this kind. This series made its mark with the treatment of taxa (pteridophytes and angiosperms) published in Rodriguesia (Freitas & Forzza 2005 in editorial). Existing knowledge which documents the morphological variation of the Brazilian species and their adpatations to the widely differing environmental conditions in the areas where they occur can be evaluated qualitatively across the broad spectrum of taxonomic treatments (revisions and Floras) already available for the Brazilian flora. Thus we concur with the conclusions of Thomas et al. (2012) in which, among other information sources mentioned by the authors, small-scale Floras are recognised as the basis for the preparation of more wide-ranging Floras, and monographs which treat all the species of a single group can be as important as partial treatments and regional revisions. Summarising, our review suggests that while levels of taxonomic coverage, collection densities and availability of taxonomic capacity may not yet have reached optimal levels in Brazil they need not represent insurmountable barriers to the creation of an Online Flora of Brazil by The proportion of the Flora which is already known, estimated here at 88%, is such that new discoveries will be the exception rather than the rule in most regions. Overall collection densities have improved significantly and compare favourably to those for other tropical countries with Floras in preparation, though the long-standing problem of undersampling in the North continues, and more intensive collection in the Northeast may also be required in order to detect the many undescribed species likely to occur there. Recent species descriptions exist for many of the species to be treated for the Flora, and will often serve as a useful starting point, although they vary greatly in scope, level of detail and degree of accessibility. And, perhaps most importantly, the trained taxonomic workforce of scientists who have already demonstrated significant commitment to enhancing the Brazilian List represents a key asset, which, if harnessed effectively, has the potential to deliver another outstanding success in the form of a comprehensive online resource covering the most species-rich flora in the world. Conclusions From the classic works produced in the 19 th Century to the present, a rich and diverse knowledge base on the species of the Brazilian flora is available in a variety of forms and scopes in taxonomic treatments (revisions and Flora), Rodriguésia 66(4):

201 1132 Morim, M.P. & Nic Lughadha, E.M. most notably the national Flora completed in the early years of the twentieth century (Martius et al.1833; Urban 1906). The taxonomic identities of the 32,086 angiosperm species then known to occur in the country, along with a suite of detailed information about each species, were compiled and checked by a network of 437 specialists over the past seven years, as part of the different stages in the finalisation of the Brazilian List. This synthesis has in turn provided a basis for a series of novel analyses of the diversity of plants and fungi of Brazil (BFG 2015; Costa & Peralta 2015; Maia et al. 2015; Menezes et al. 2015; Prado et al. 2015). The body of knowledge of Brazilian plants generated over the decades of taxonomic endeavour in Brazil is intimately linked to and, in large part, based on the enormous number of specimens deposited in herbaria within Brazil and in other countries, which day-by-day are being digitised and made available online, in the form of Virtual Herbaria. Initiatives such as the Reflora Virtual Herbarium illustrate the progress which has been made. To date more than one million high resolution images of specimens from different herbaria, in Brazil and elsewhere, are available not only for reference purposes but also so that taxonomists can add new determinations, nomenclatural updates and information of other kinds, where appropriate. In this context, we consider that planning an online Flora of Brazil for 2020 is a realistic idea, based on a solid foundation. Our awareness of the limitations and lacunas in our knowledge of the flora should not deter us from our efforts to document it. On the contrary, this awareness should form the basis for a strategic plan that envisages proceeding with the Flora, while at the same time directing efforts and resources to build taxonomic capacity and address the numerous knowledge gaps. Furthermore, it must always be borne in mind that the preparation of an Online Flora allows for continuous updating, as our knowledge of the flora improves. This incremental approach has had notable success over the past seven years. Indeed, the present moment seems to offer a unique opportunity to capture the momentum created by the finalisation of the Brazilian List and the major milestones reached by Reflora, building on these achievements to reach the new, still more challenging target. To meet Target 1 of the GSPC by 2020 is, without doubt, a huge project which will require consideration of aspects beyond the capacity and motivation of the taxonomic community: interinstitutional linkages, financial investment and the support of decision-makers will be fundamental to the success of the project. Providing the conditions and resources required for the development of the Flora of Brazil Online 2020 is a responsibility which Brazil, as a party to the CBD, must accept and incorporate as an integral part of government policies and plans. Acknowledgements The authors are grateful to the following, without whom preparation of this review would have been difficult, if not impossible: taxonomists Cassia M.Sakuragui, Rafaela C.Forzza, Thiago E.C. Meneguzzo and Wayt Thomas who shared data on the number of species treated in: Philodendron section Macrobelium, Aechmea subg.chevalliera, Orchidaceae and in Flora Neotropica Monographs ( ) respectively; Andrielle C. A. Lopes who shared data on the Flora of the Distrito Federal; Lucas S. B. Jordan who collaborated on the quantitative survey of species of Flora Ilustrada Catarinense; Catia Canteiro who assisted in the preparation of the Figures; Martin Cheek and Daniela Zappi who provided useful comments which helped improve an earlier version of the manuscript; two anonymous referees who provided helpful suggestions to improve the final manuscript; Rafael Govaerts, Rafaela Forzza and Luis Alexandre Estevão da Silva who provided prompt responses to last-minute requests for missing facts; Fabiana L. Ranzato Filardi and Paula Leitman who responded patiently to queries throughout the preparation of the Lista de Espécies da Flora do Brasil and continue to do so on a daily basis; all taxonomists who work with plants in Brazil. References Alves-Araújo, A; Dutilh, J.H.A. & Alves, M Amaryllidaceae s.s. e Alliaceae s.s. no Nordeste Brasileiro. Rodriguésia 60: Andrade-Lima, D A botânica da Carta de Pero Vaz de Caminha. Rodriguésia 36: 5-8. Antonelli, A. & Sanmartín, I Why are there so many plant species in the Neotropics? Taxon 60: Barbosa, M.V de & Peixoto, A Coleções botânicas brasileiras: situação atual e perspectivas. In: Peixoto, A. (org.). Coleções biológicas de apoio ao inventário, uso sustentável e conservação de biodiversidade. Instituto de Pesquisas Jardim Rodriguésia 66(4):

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204 Flora of Brazil OnLine 2020 Reitz, R (ed.). Flora Ilustrada Catarinense. Herbário Barbosa Rodrigues, Itajaí. Avaiable at < enciclopedia-flora-ilustrada-catarinense-fic/>. Access on 15 May Shepherd, G.J Avaliação do estado do conhecimento da diversidade biológica do Brasil: plantas terrestres versão preliminar. Ministério do Meio Ambiente, Brasília. 60p. Soares, K.P.; Longhi, S.J.; Neto, L.W. & Assis, L.C Palmeiras (Arecaceae) Rio Grande do Sul, Brasil. Rodriguésia 65: Sobral, M. & Stehmann, J.R An analysis of new angiosperm species discoveries in Brazil ( ). Taxon 58: ter Steege, H.; Pitman, N.C.A.; Sabatier, D.; Baraloto, C.; Salomao, R.P.; Guevara, J.E.; Phillips, O.L.; Castilho, C.V.; Magnusson, W.E.; Molino, J.-F.; Monteagudo, A.; Nunez Vargas, P.; Carlos Montero, J.; Feldpausch, T.R.; Coronado, E.N.H.; Killeen, T.J.; Mostacedo, B.; Vasquez, R.; Assis, R.L.; Terborgh, J.; Wittmann, F.; Andrade, A.; Laurance, W.F.; Laurance, S.G.W.; Marimon, B.S.; Marimon, B.-H.; Guimaraes Vieira, I.C.; Amaral, I.L.; Brienen, R.; Castellanos, H.; Cardenas Lopez, D.; Duivenvoorden, J.F.; Mogollon, H.F.; de Almeida Matos, F.D.; Davila, N.; Garcia-Villacorta, R.; Stevenson Diaz, P.R.; Costa, F.; Emilio, T.; Levis, C.; Schietti, J.; Souza, P.; Alonso, A.; Dallmeier, F.; Duque Montoya, A.J.; Fernandez Piedade, M.T.; Araujo-Murakami, A.; Arroyo, L.; Gribel, R.; Fine, P.V.A.; Peres, C.A.; Toledo, M.; Gerardo, A.A.C.; Baker, T.R.; Ceron, C.; Engel, J.; Henkel, T.W.; Maas, P.; Petronelli, P.; Stropp, J.; Eugene Zartman, C.; Daly, D.; Neill, D.; Silveira, M.; Rios Paredes, M.; Chave, J., de Andrade Lima, D.; Jorgensen, P.M.; Fuentes, A.; Schoengart, J.; Cornejo Valverde, F.; Di Fiore, A.; Jimenez, E.M.; Penuela Mora, M.C.; Fernando Phillips, J.; Rivas, G.; van Andel, T.R.; von Hildebrand, P.; Hoffman, B.; Zent, E.L.; Malhi, Y.; Prieto, A.; Rudas, A.; Ruschell, A.R.; Silva, N.; Vos, V.; Zent, S.; Oliveira, A.A.; Cano Schutz, A.; Gonzales, T.; 1135 Nascimento, M.T.; Ramirez-Angulo, H.; Sierra, R.; Tirado, M.; Umana Medina, M.N.; van der Heijden, G.; Vela, C.I.A.; Vilanova Torre, E.; Vriesendorp, C.; Wang, O.; Young, K.R.; Baider, C.; Balslev, H.; Ferreira, C.; Mesones, I.; Torres-Lezama, A.; Urrego Giraldo, L.E.; Zagt, R.; Alexiades, M.N.; Hernandez, L.; Huamantupa-Chuquimaco, I.; Milliken, W.; Palacios Cuenca, W.; Pauletto, D.; Valderrama Sandoval, E.; Valenzuela Gamarra, L.; Dexter, K.G. & Feeley, K Hyperdominance in the Amazonian tree flora. Science 342: 325. DOI: /science Thiers, B. [continuously updated]. Index Herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden s Virtual Herbarium. Available at < org/ih/>. Access on 15 May Thomas, W.W Flora Neotropica-Monographs as Inventories. In: Species plantarum 250 years: proceedings of the species plantarum symposium held in Uppsala August 22-24, Acta Universitatis Upsaliensis, Symbolae Botanicae Upsalienses 33: Thomas, W.W.; Forzza, R.C; Michelangeli, F.A.; Giulietti, A.M. & Leitman, P.M Large-scale monographs and floras: the sum of local floristic research. Plant Ecology and Diversity: 5: Trias-Blasi, A. & Vorontosova, M Botany: plant identification is key to conservation. Nature 521: 161. Urban, I Index Familiarum. In: Martius, C.P.F. von. Flora Brasiliensis. Karl W. Hiersemann, Munich & Leipzig. Vol. 1, pars 1, pp Wanderley, M.G.L.; Shepherd, G.J.; Martins, S.E., Estrada, T.E.M.D; Romanini, R.P.; Koch,I.; Pirani, J.R.; Melhem, T.S.; Harley, A.M.G.G.; Kinoshita, L.S.; Magenta,M.A.G.; Longhi Wagner, H.M.; Barros, F.; Lohmann, L.G.; Amaral, M.C.E.; Cordeiro, I.; Aragaki, S.; Bianchini, R.S. & Esteves, G.L Checklist das Spermatophyta do Estado de São Paulo, Brasil. Biota Neotropica 11: Available at < Access on 5 June Artigo recebido em 09/06/2015. Aceito para publicação em 19/08/2015. Rodriguésia 66(4):

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206 Rodriguésia 66(4): DOI: / The naturalized flora of Brazil: a step towards identifying future invasive non-native species Rafael Dudeque Zenni 1 Abstract Human-mediated species introductions are one of the defining factors of the Anthropocene. Non-native species can form invasive populations that affect biodiversity, ecosystem services, or farming. Here I summarized data on naturalized vascular species from the Brazilian list of seed plants and tested the hypotheses that more populated regions and biomes with less remaining natural vegetation harbor more naturalized species. The Atlantic Forest had the largest number of naturalized species, whereas the Pampa had the highest proportion of naturalized species in relation to the biome s total richness. The number of naturalized species was affected both by human population size and proportion of remaining natural vegetation. Family Poaceae had the highest numbers of naturalized species in all biomes, and, together with Asteraceae and Fabaceae, forms the top three families in number of naturalized species in five of the biomes. There were no records of species naturalized in all six biomes. However, of the 46 species naturalized in five out of six biomes, half were Poaceae. The results indicate that the Brazilian flora is currently composed both by native and naturalized species. Assessments of invasion risks and of impacts by naturalized species could help set management priorities and resource allocation towards control. Key words: Alien species, early detection, invasion biology, nature conservation. Resumo Introduções de espécies mediadas por humanos são um dos fatores determinantes do Antropoceno. Espécies exóticas podem formar populações invasoras que afetam a biodiversidade, serviços ecossistêmicos e agropecuária. Neste trabalho, eu avalio os dados sobre espécies naturalizadas no Brasil da lista de plantas com semente da flora do Brasil e testo as hipóteses de que regiões mais populosas e biomas com menor área natural remanescente contêm mais espécies naturalizadas. A Mata Atlântica teve o maior número de espécies naturalizadas, enquanto o Pampa teve a maior proporção de espécies naturalizadas em relação ao total de espécies no bioma. O número de espécies naturalizadas foi afetado tanto pelo número de habitantes quanto pela proporção de vegetação remanescente nos biomas. A família Poaceae conteve o maior número de espécies naturalizadas em todos os biomas e, junto com as famílias Asteraceae e Fabaceae, foram as três famílias com maior número de espécies naturalizadas em cinco biomas. Não houve registro de espécies naturalizadas em todos os seis biomas. Entretanto, das 46 espécies naturalizadas em cinco biomas, metade são Poaceae. Os resultados indicam que a flora brasileira é atualmente composta por espécies nativas e naturalizadas. Análises formais dos riscos de invasão biológica e potenciais impactos negativos causados por espécies naturalizadas devem ajudar na definição de prioridades de manejo e na alocação de recurso para controle. Palavras-chave: Biologia de invasões, conservação da natureza, detecção precoce, espécies exóticas. Introduction Humans have been transporting species around the globe for centuries. Among the reasons for introducing species to regions where they did not occur previously are agriculture, agroforestry, forestry, forage, and horticulture (Zenni 2014). As a consequence, virtually all ecosystems in the world currently host non-native species (van Kleunen et al. 2015). Together with other drivers of global environmental change (i.e., climate change and land use transformation), human-mediated species introductions are one of the defining factors of the Anthropocene (Lewis & Maslin 2015). A small proportion of the species introduced either with 1 Universidade de Brasília, Inst. Ciências Biológicas, Campus Darcy Ribeiro, Asa Norte, bl. E, , Brasilia, DF. rafaeldz@gmail.com

207 1138 Zenni, R.D. intentional or accidental human assistance form invasive populations that threaten biodiversity, ecosystem services, and farming (Pyšek et al. 2012; Yelenik & D Antonio 2013). Some invasive species are known to have caused species extinctions, immense economic losses, and losses of important crops around the world (Simberloff 2013). Thus, it is pivotal to understand the patterns and drivers of species naturalizations and invasions in order to prevent and reduce the negative impacts caused by biological invasions. A biological invasion is a populationlevel process often defined as the introductionnaturalization-invasion continuum (Blackburn et al. 2011). Briefly, an organism which survives transport is successfully introduced into a region where it did not occur previously (i.e., non-native species), reaches reproductive age, produces descendants to form a viable self-sustaining population (i.e., naturalized species), and spreads over considerable distances from the point of introduction in a short period of time, often achieving high population densities (i.e., invasive species). For instance, for woody plants it was suggested that a population spreading 100 m away from the point of introduction in less than 50 years could be considered a biological invasion (Richardson et al. 2000). Only a fraction of the non-native species form naturalized populations, and only a small subset of these result in invasions. Communities containing a combination of native and naturalized species are the new norm in most ecosystems (van Kleunen et al. 2015) and some of these naturalized populations will eventually invade. Thus, although biological invasions are considered an improbable result of species introductions, when it does happen it may cause dramatic changes in native communities and ecosystems and reach massive proportions (Wardle et al. 2011). From a management point of view, it is always preferable to prevent invasions from happening or controlling them no later than just after the naturalization stage, when populations are often restricted to well-delimited areas. The Brazilian List of seed plants (BFG 2015) identified 32,634 vascular species in Brazil, of which 525 were naturalized non-native species. The inventory revealed that the Atlantic Forest is the richest biome with 15,001 native species, followed by the Cerrado (12,097 native species), the Amazon (11,896), the Caatinga (4,657), the Pampa (1,685), and the Pantanal (1,277). However, analyses of numbers, types, and proportions of naturalized species in Brazil were lacking. In this study, I aimed to (i) summarize the data on naturalized species available in the inventory of Brazilian seed plants and (ii) to test the hypotheses that biomes with more people and less remaining natural vegetation cover of Brazil harbor more naturalized species. Material and Methods Data were gathered from BFG (2015), Costa & Peralta (2015), Maia et al. (2015), Menezes et al. (2015) and Prado et al. (2015). A detailed description of how the list was compiled and curated is available in BFG (2015). I kept only vascular plant species with accepted names, known to be present in Brazil, and with known origin (i.e., native, naturalized, or cultivated). Avascular plants were removed because data on these groups were limited to a few regions and knowledge regarding the actual origin of many avascular plants is not available. From those species, I prepared a list of States, regions, terrestrial biomes (Atlantic Forest, Amazon, Caatinga, Cerrado, Pampa, and Pantanal), and vegetation types (phytophysiognomies) where they were reported to occur. I also obtained the percentage of remaining natural vegetation cover for each biome from the Brazilian Biomes Project (Projeto biomas do Brasil 2015). Finally, I gathered data on the States total areas, States urban areas, and human population per biome from the Brazilian Institute of Geography and Statistics (IBGE 2015). All the IBGE data are for the year 2009 and were the most recent available. I calculated the proportion of naturalized species in each biome by dividing the number of naturalized species by the sum of all species known to occur in the biome (native, cultivated, and naturalized). To test the associations between number of naturalized species per biome and human population size, percentage of remaining natural vegetation cover, and number of native species, I used generalized linear models (GLM) with Poisson error distribution (α = 0.05). I also tested the associations between number of naturalized species and the proportion of urban area per State. State or biome total area was added to all models as a covariate to check whether results were scale dependent. I tested both biome and State level metrics in order to obtain two independent confirmations of the hypotheses that naturalized species richness is associated to human population and to conversion of natural areas to areas of Rodriguésia 66(4):

208 The naturalized flora of Brazil intensive use (i.e., urban, agriculture, and pastures). Lastly, I used a chi-square test to see if native and naturalized species were equally frequent in anthropogenic areas (e.g., degraded areas). Results There were a total 525 non-native naturalized vascular plant species in the inventory of Brazilian plants (Appendix 1). The Atlantic Forest biome had the highest number of naturalized species (n = 416), followed by the Cerrado (n = 223), the Amazon (n = 184), the Caatinga (n = 155), the Pampa (n = 114), and the Pantanal (n = 39) (Fig. 1a). In terms of proportional representation, the Pampa had the highest number of non-native naturalized species per total species richness (6.8%), followed by the Caatinga (3.4%) and the Pantanal (3.6%), the Atlantic Forest (2.7%), the Cerrado (1.9%), and the Amazon (1.5%) (Fig. 1b). The number of naturalized species in a biome was positively associated with human population size (z=2.3, p=0.02; Fig. 2a) and was not affected by biome total area (z=1.8, p=0.08). Also, the number of naturalized species in a given biome was negatively associated with the proportion of remaining natural vegetation cover (z=-9.2, p<0.001; Fig. 2b). Less remaining natural vegetation meant more naturalized species. Further, the number of native species in a biome predicted the number of non-native naturalized species (z=11.7, p<0.001; Fig. 3); more native species meant more naturalized species. However, in these two previous cases, the interaction term with biome total area was also significant (z=-3.7 and z=-3.8, respectively, p<0.001). As a consequence, it is not possible to evaluate the sole role of remaining natural vegetation in naturalized species richness and biome size must be taken into consideration. Removing outliers did not change the results of the tests. When looking at the State level, I found patterns similar to those of the analyses with biomes. States with larger urban areas harbour more naturalized species (z=10.6, p<0.001) and this effect was independent of State size (z=0.3, p=0.76). Likewise, States with larger population sizes also had more naturalized species (z=9.74, p<0.001). Again, the effect was independent of State size (z=-0.9, p=0.2). São Paulo had the largest number of naturalized species (n=332), followed by Paraná (n=270), Minas Gerais (n=263), Santa Catarina (n=260), and Rio Grande do Sul (n=255) Non-native species were more likely to be recorded occurring in disturbed areas than native species (c 2 =4.39, p<0.001). While 71.05% (n=373) of the naturalized species were reported to occur in disturbed areas, only 6.46% (n=2,042) of the native species occurred in these habitats. The remaining naturalized species (n=124) were only recorded occurring in natural areas or had no specific habitat associated to them (n=28). By definition Figure 1 a. Number of non-native naturalized plant species; b. proportion of non-native naturalized plant species present across six terrestrial biomes and two aquatic regions in Brazil. Lighter shades represent lower numbers and proportions, whereas darker shades depict higher numbers and proportions. Rodriguésia 66(4):

209 1140 Zenni, R.D. Table 1 The three families with the highest numbers of non-native naturalized species per biome, the number of species in each family that are naturalized in each biome and the proportion of naturalized species in each biome that belong to the family. Biome Family Number of naturalized species % Atlantic Forest Poaceae Asteraceae Fabaceae Cerrado Poaceae Asteraceae Fabaceae Amazon Poaceae Fabaceae Asteraceae Caatinga Poaceae Fabaceae Amaranthaceae Pampa Poaceae Caryophyllaceae Asteraceae Pantanal Poaceae Fabaceae Asteraceae (i.e., Blackburn et al. 2011), it is probable the 124 records of non-native species in natural habitats were records of invasion rather than naturalization. The family Poaceae had the largest number of naturalized species in all six biomes (Tab. 1). The families Asteraceae and Fabaceae were among the top three families in number of naturalized species in five out of the six terrestrial biomes, albeit not in the same ones. Those three families were the three richer ones in the Amazon, the Atlantic Forest, the Cerrado, and the Pantanal biomes. In the Caatinga, the family Amaranthaceae was the third in species richness after Poaceae and Fabaceae. And in the Pampa biome the family Caryophyllaceae was the second richest after Poaceae, before Asteraceae (Tab. 1). None of the 525 species were naturalized in all six biomes. However, of the 46 non-native species that were naturalized in five biomes, half were grasses (Poaceae). The remaining species belonged to 14 different families. Another 52 species were present across four terrestrial biomes, of which 19 species belong to the Poaceae family and the remaining species are contained in 18 different families. Thus, a single family (Poaceae) made up to 43% of all non-native species naturalized in more than half of the Brazilian terrestrial biomes. Discussion Based on the data gathered by the Brazilian Flora Group (BFG 2015), the hypotheses that more populated and more deforested biomes of Brazil harbour more naturalized species were supported. The results also supported the idea that habitats that support a high diversity of native species also tend to support a high diversity of non-native species (Fig. 3). The Poaceae family was disproportionally overrepresented in the non-native naturalized flora, having the highest overall richness among all naturalized species (n=142) and the most widespread species (23 species naturalized across 5 biomes). The Poaceae family was also overrepresented among invasive non-native species in Brazil (Zenni & Ziller 2011), especially the African C-4 grasses (Zenni 2014). African grasses are known to benefit from Rodriguésia 66(4):

210 The naturalized flora of Brazil Figure 2 Relationship between number of non-native naturalized plant species in the Brazilian Biomes and a. human population in each biome; b. percentage of the native biome habitats remaining (areas not converted for human intensive use). Solid lines represent generalized linear model fittings and grey shades are ±95% confidence intervals. The outlier in (a) did not change model fitting. anthropogenic disturbances, thus it comes as no surprise that grasses were the most abundant group of naturalized plants in a scenario where the naturalization of non-native species is both directly and indirectly related to human population, urbanization, and habitat degradation. The results make it clear that a significant part of the Brazilian flora is now composed 1141 of introduced species forming self-sustaining populations. While the Atlantic Forest had the richest non-native naturalized flora, the Pampa had the highest proportion of naturalized species. Given that native and naturalized species richness were correlated (Fig. 3), it is unclear which one of the two metrics better reflect the current magnitude of species naturalization in the biomes. I suggest considering both (richness and proportion) in subsequent analyses. Although naturalized species richness was associated with human-induced disturbances, previous work has shown that nonnative species are also widespread in Brazilian natural and protected areas (Sampaio & Schmidt 2013; Ziller & Dechoum 2013). Ecologists, managers, and policy makers need a better understanding of the role non-native species have in natural ecosystems. The inventory of the Brazilian Flora did not point out which of the naturalized species are known to have invasive populations, and the available data on invasive alien plants in Brazil were gathered using different criteria. Thus, it is unclear which naturalized species became invasive in the different biomes and regions. Previous research identified at least 180 invasive alien plants in Brazil (Sampaio & Schmidt 2013; Ziller & Dechoum 2013; Zenni & Ziller 2011), but only 40% (n=73) of these invasive non-native species were also listed in the inventory of the Brazilian flora. The remaining 107 non-native species considered invasive somewhere in Brazil were absent from the inventory. For instance, the inventory of Brazilian seed plants used in this study did not include any of the five Acacia Mill. species (Fabaceae) which are currently being cultivated and shown to be naturalized (Attias et al. 2013). When considering only the invasive non-native species that were included in the inventory, 13.4% of the naturalized species in Brazil have already crossed the naturalization stage and become invasive. Because of these discrepancies, it is unclear how underestimated the inventory of the Brazilian flora is regarding non-native species and if datasets on invasive species are overestimated. Future updates of the Brazilian List should try to address the current gap in Brazilian non-native flora. Naturalization is a required intermediate stage towards biological invasion (Blackburn et al. 2011). While some naturalized populations may never turn into invasive populations, others will. Thus, non-native naturalized populations Rodriguésia 66(4):

211 1142 Zenni, R.D. Figure 3 Relationship between number of native species and number of naturalized species in Brazilian Biomes. The solid line represents the GLM fit and grey shades represent ±95% confidence intervals. and species that are not invading should be prioritized for research and management efforts regarding potential invasion and impact on native ecosystems. For instance, the Amazon biome has 80% of its natural area remaining, has the third highest number of naturalized species, but has the lowest number of invasive species (Zenni & Dechoum 2013). Inversely, the Atlantic Forest is the most degraded biome, and has both the highest numbers of naturalized and invasive species. These two opposite scenarios provide important insights for dealing with biological invasions. First, the current condition of the Atlantic Forest suggests what may happen in the Amazon and other biomes if managers and policy makers do not take action. Second, different regions of Brazil may require different strategies regarding non-native species; for instance, while in the Amazon the focus should be on prevention and early detection of invasive non-native species, in the Atlantic Forest the focus should be on containment and reduction of invaded areas, and on ecological restoration. Whenever possible, especially regarding protected areas, naturalized populations posing high invasion risk should be excluded before they invade. By understanding the current status of naturalized species in different habitats and the drivers of naturalization (i.e., which species are naturalized, and where the naturalized populations are), managers and policy makers can identify priorities and allocate resources wisely to prevent spread and potential negative impacts. The focus of the Brazilian list of seed plants was to catalogue the native flora of Brazil (BFG 2015), and the initiative to include nonnative species (cultivated and naturalized) could be considered a parallel effort. Also, inclusion of species in the list depended on herbarium vouchers, but not all botanists include non-native plants in their samplings and many ecologists working on non-native plants do not make herbarium deposits of study species. The lack of herbarium vouchers seems to be especially true for non-native species that are neither naturalized nor invasive (e.g., cultivated). The existing gap between the list of Brazilian seed plants and several published lists of invasive plants attests to that fact. Consequently, the current list of naturalized species in Brazil and the estimates I obtained for numbers and proportions of species may be considered conservative estimates. Another limitation of this study is that species that are native to some Brazilian biome or vegetation type, but are naturalized in another biome or vegetation type were not considered by the group of researchers that compiled the Brazilian list. For instance, Schyzolobium parahyba (Vell.) S.F.Blake is native to the Atlantic Coastal Rainforest, but became invasive in Seasonally Semi-deciduous Forests (both are part of the Atlantic Forest biome) after human-mediated introduction (Abreu et al. 2014). Given that species ranges are a biogeographical concept rather than a geopolitical one, care should be taken when interpreting statements of species as native or non-native to Brazil. Also, botanists should be considerate of inter-habitat, but intranational, species introductions. In conclusion, the current study presents the first nationwide assessment of the naturalized flora of Brazil and conveys important knowledge for research and conservation prioritization. It was made evident that non-native species are widespread in all Brazilian biomes and regions. Human presence, and human actions and activities, are an important reason for the observed patterns of non-native species naturalization. Research and conservation actions could greatly benefit from the integration among databases of native and nonnative species in Brazil regardless of the statuses of populations. To improve the record of non-native Rodriguésia 66(4):

212 The naturalized flora of Brazil 1143 Figure 4 Relationship between number of naturalized plant species in Brazilian States and (a) State urban area (km 2 ), and (b) human population size. Solid lines represent generalized linear model fittings and grey shades are ±95% confidence intervals. The outliers did not change model statistical significance. species, researchers and managers working on invasive non-native species should make an effort to improve the herbarium record on non-native species. Finally, formal assessments of invasion risks and of negative impacts caused by the species identified as already naturalized could help setting management priorities and resource allocation. Acknowledgements I thank Dr. Michele Dechoum and Daniela Zappi for comments that helped to improve the manuscript. This work would not have been possible without the collective effort of the Brazilian Flora Group in gathering, curating, and sharing the data on plant species in Brazil. RDZ is supported by CNPq-Brazil (Grant /2014-0). References Abreu, R.C.R.; Santos, F.F.M. & Durigan, G Changes in plant community of seasonally semideciduous forest after invasion by Schizolobium parahyba at southeastern Brazil. Acta Oecologica 54: Attias, N.; Siqueira, M.F. & Bergallo, H.G Acácias australianas no Brasil: histórico, formas de uso e potencial de invasão. Biodiversidade Brasileira 3: BFG Growing knowledge: an overview of Seed Plant diversity in Brazil. Rodriguésia 66: Blackburn, T.M.; Pyšek, P.; Bacher, S.; Carlton, J.T.; Duncan, R.P.; Jarošík, V.; Wilson J.R.U. & Richardson D.M A proposed unified framework for biological invasions. Trends in Ecology & Evolution 26: Costa, D.P. & Peralta, D.F Bryophytes diversity in Brazil Rodriguésia 66: IBGE Instituto Brasileiro de Geografia e Estatística. Availeble at < home/>. Access on 3 September Lewis, SL & Maslin, MA Defining the anthropocene. Nature 519: Projeto Biomas do Brasil Biomas do Brasil. Available at < >. Access on 3 September Maia, L.C.; Carvalho Júnior, A.A.; Cavalcanti, L.H.; Gugliotta, A.M.; Drechsler-Santos, E.R.; Santiago, A.L.M.A.; Cáceres, M.E.S.; Gibertoni, T.B.; Aptroot, A.; Giachini, A.J.; Soares, A.M.S.; Silva, A.C.G.; Magnago, A.C.; Goto, B.T.; Lira, C.R.S.; Montoya, C.A.S.; Pires-Zottarelli, C.L.A.; Silva, D.K.A.; Soares, D.J.; Rezende, D.H.C.; Luz, E.D.M.N.; Gumboski, E.L.; Wartchow, F.; Karstedt, F.; Freire, F.M.; Coutinho, F.P.; Melo, G.S.N.; Sotão, H.M.P.; Baseia, I.G.; Pereira, J.; Oliveira, J.J.S.; Souza, J.F.; Bezerra, J.L.; Araujo Neta, L.S.; Pfenning, L.H.; Gusmão, L.F.P.; Neves, M.A.; Capelari, M.; Jaeger, M.C.W.; Pulgarín, M.P.; Menolli Junior, N.; Medeiros, P.S.; Friedrich, R.C.S.; Chikowski, R.S.; Pires, R.M.; Melo, R.F.; Silveira, R.M.B.; Urrea-Valencia, S.; Cortez, V.G. & Silva, V.F Diversity of Brazilian Fungi. Rodriguésia 66: Rodriguésia 66(4):

213 1144 Zenni, R.D. Menezes, M.; Bicudo, C.E.M.; Moura, C.W.N.; Alves, A.M.; Santos, A.A.; Pedrini, A.G.; Araújo, A.; Tucci, A.; Fajar, A.; Malone, C.; Kano, C.H.; Sant Anna, C.L.; Branco, C.Z.; Odebrecht, C.; Peres, C.K.; Neuhaus, E.B.; Eskinazi-Leça, E.; Aquino, E.; Nauer, F.; Santos, G.N.; Amado Filho, G.M.; Lyra, G.M.; Borges, G.C.P.; Costa, I.O.; Nogueira, I.S.; Oliveira, I.B.; Paula, J.C.; Nunes, J.M.C.; Lima, J.C.; Santos, K.R.S.; Ferreira, L.C.; Gestinari, L.M.S.; Cardoso, L.S.; Figueiredo, M.A.O.; Silva, M.H.; Barreto, M.B.B.B.; Henriques, M.C.O.; Cunha, M.G.G.S.; Bandeira-Pedrosa, M.E.; Oliveira-Carvalho, M.F.; Széchy, M.T.M.; Azevedo, M.T.P.; Oliveira, M.C.; Cabezudo, M.M.; Santiago, M.F.; Bergesh, M.; Fujii, M.T.; Bueno, N.C.; Necchi Jr., O.; Jesus, P.B.; Bahia, R.G.; Khader, S.; Alves-da- Silva, S.M.; Guimarães, S.M.P.B.; Pereira, S.M.B.; Caires, T.A.; Meurer, T.; Cassano, V.; Werner, V.R.; Gama Jr., W.A. & da Silva, W.J. Update of the Brazilian floristic list of Algae and Cyanobacteria. Rodriguésia 66: Prado, J.; Sylvestre, L.S.; Labiak, P.H.; Windisch, P.G.; Salino, A.; Barros, I.C.L.; Hirai, R.Y.; Almeida, T.E.; Santiago, A.C.P.; Kieling-Rubio, M.A.; Pereira, A.F.N.; Øllgaard, B.; Ramos, C.G.V.; Mickel, J.T.; Dittrich, V.A.O.; Mynssen, C.M.; Schwartsburd, P.B.; Condack, J.P.S.; Pereira, J.B.S.& Matos, F.B. Diversity of ferns and lycophytes in Brazil. Rodriguésia 66: Pyšek, P.; Jarošík, V.; Hulme, P.E.; Pergl, J.; Hejda, M.; Schaffner, U. & Vilà, M A global assessment of invasive plant impacts on resident species, communities and ecosystems: the interaction of impact measures, invading species traits and environment. Global Change Biology 18: Richardson, D.M.; Pyšek, P.; Rejmánek, M.; Barbour, M.G.; Panetta, F. D. & West, C.J Naturalization and invasion of alien plants: concepts and definitions. Diversity and Distributions 6: Sampaio, A.B. & Schmidt, I.B Espécies exóticas invasoras em unidades de conservação federais do Brasil. Biodiversidade Brasileira 3: Simberloff, D Invasive species: what everyone needs to know. 1ed. Oxford University Press, New York. 329p. van Kleunen, M.; Dawson, W.; Essl, F.; Pergl, J.; Winter, M.; Weber, E.; Kreft, H.; Weigelt, P.; Kartesz, J.; Nishino, M.; Antonova, L.A.; Barcelona, J.F.; Cabezas, F.J.; Cardenas, D.; Cardenas-Toro, J.; Castano, N.; Chacon, E.; Chatelain, C.; Ebel, A.L.; Figueiredo, E.; Fuentes, N.; Groom, Q.J.; Henderson, L.; Inderjit; Kupriyanov, A.; Masciadri, S.; Meerman, J.; Morozova, O.; Moser, D.; Nickrent, D.L.; Patzelt, A.; Pelser, P.B.; Baptiste, M.P.; Poopath, M.; Schulze, M.; Seebens, H.; Shu, W.- s.; Thomas, J.; Velayos, M.; Wieringa, J. J. & Pysek, P Global exchange and accumulation of non-native plants. Nature 525: Wardle, D.A.; Bardgett, R.D.; Callaway, R.M. & Van der Putten, W.H Terrestrial ecosystem responses to species gains and losses. Science 332: Yelenik, S.G. & D Antonio, C.M Self-reinforcing impacts of plant invasions change over time. Nature 503: Zenni, R.D. & Ziller, S.R An overview of invasive plants in Brazil. Brazilian Journal of Botany 34: Zenni, R.D. & Dechoum, M.S Paisagens antropizadas e invasão por plantas exóticas. In: Peres, C.A; Barlow, J.; Gardner, T.A. & Vieira, I.C.G. Conservação da biodiversidade em paisagens antropizadas do Brasil. Editora UFPR, Curitiba. Pp Zenni, R.D Analysis of introduction history of invasive plants in Brazil reveals patterns of association between biogeographical origin and reason for introduction. Austral Ecology 39: Ziller, S.R. & Dechoum, M.S Plantas e vertebrados exóticos invasores em unidades de conservação no Brasil. Biodiversidade Brasileira 3:4-31. Artigo recebido em 04/09/2015. Aceito para publicação em 12/11/2015. Rodriguésia 66(4):

214 Rodriguésia 66(4): DOI: / Angiosperm Checklist of Espírito Santo: using electronic tools to improve the knowledge of an Atlantic Forest biodiversity hotspot Valquíria Ferreira Dutra 1, Anderson Alves-Araújo 2 & Tatiana Tavares Carrijo 3 Abstract A checklist of angiosperm species for the state of Espírito Santo, Brazil is presented. A total of 6,204 native species was recorded, representing an increase of 16% in species richness for the area. Espírito Santo shelters 32% of the native species of Brazil s Atlantic Forest and holds 516 endemic species (8.3% of the total registered taxa for the State). Bromeliaceae and Orchidaceae are the families with highest number of endemic species (142 spp. and 80 spp., respectively), followed by Myrtaceae (33 spp.), Melastomataceae (30 spp.), and Araceae (23 spp.). This paper represents an important landmark for future research in plant diversity in Espírito Santo, and highlights the importance of consulting online databases in order to update the knowledge presented by the Brazilian flora checklist. Key words: Brazil, endemism, floristics, species richness, virtual herbarium. Resumo Uma listagem de angiospermas do estado do Espírito Santo é apresentada, incluindo um total de espécies nativas, representando um aumento de 16% na riqueza de espécies registrada para o Estado. O Espírito Santo abriga 32% das espécies nativas da Floresta Atlântica e apresenta 516 espécies endêmicas (8.3% do total registrado para o Estado). As famílias Bromeliaceae e Orchidaceae apresentam o maior número de espécies endêmicas (142 spp. e 80 spp., respectivamente), seguidas por Myrtaceae (33 spp.), Melastomataceae (30 spp.) e Araceae (23 spp.). Os resultados representam um importante marco para futuras pesquisas em diversidade de plantas no Espírito Santo e destacam a importância de consultar as plataformas virtuais para atualização da Lista de Espécies da Flora do Brasil. Palavras-chave: Brasil, endemismo, flora, herbários virtuais, riqueza de espécies. Introduction The Brazilian Atlantic Forest is one of the richest biogeographic zones for plant species and endemism in the world (Prance 1982; Mittermeier et al. 1999; Tabarelli et al. 2004). The high number of endemic species, together with the accelerated habitat loss, places this biome among the 35 biodiversity hotspots of the world (Myers et al. 2000; Mittermeier et al. 2005), justifying a higher conservation priority. Centuries of deforestation have led to the reduction of this biome s original cover to %, resulting in forest fragmentation, with most fragment sizes smaller than 50 ha (Ribeiro et al. 1999). This is of major concern, because species richness can be strongly affected by the size and shape of remnants, the degree of connectivity among them and by land use history (Metzger 2003). Espírito Santo is one of Brazil s richest states in terms of angiosperm species. The state has an area of approximately 45,600 km 2, which was originally completely covered by Atlantic Forest (MMA 2000). The different vegetation types comprise ombrophyllous and semideciduous forests, swamps, restingas (coastal lowland formations), mangroves, and vegetation refuges (MMA 2000; Pereira 2007). There are two main geological zones in the state: a central-southern 1 Universidade Federal do Espírito Santo, Depto. Ciências Biológicas, CCHN, Av. Fernando Ferrari 514, Goiabeiras, , Vitória, ES, Brazil. valquiriafdutra@gmail.com 2 Universidade Federal do Espírito Santo, Depto. Ciências Agrárias e Biológicas, CEUNES, Rodovia BR 101 Norte, Km 60, Litorâneo, , São Mateus, ES, Brazil. sapotae@gmail.com 3 Universidade Federal do Espírito Santo, Depto. Biologia, CCA, Alto Universitário s.n., , Alegre, ES, Brazil. tcarrijo@gmail.com

215 1146 Dutra, V.F., Alves-Araújo, A. & Carrijo, T.T. montane region reaching 2,000 m above sea level, comprising deep valleys with open ombrophyllous and upper montane forests, and a tabuleiro (tabletop mountain) zone in the central-northern region. The vegetation of the latter ranges from restinga to dense ombrophyllous forests (IPEMA 2005). Espírito Santo is included in the Corredor Central da Mata Atlântica (Aguiar et al. 2005), one of the main regions of plant endemism in Brazil (Prance 1982; IPEMA 2005). According to the Brazil Flora Group (BFG 2015) the angiosperm diversity estimated for Espírito Santo comprises about 16.5% of the total species richness in Brazil, with an estimated 32,109 taxa. Amongst these species, many are restricted and currently assigned as threatened (BFG 2015; Martinelli & Moraes 2013). Despite the existence of some floristic studies carried out in the last three decades (e.g. Peixoto & Gentry 1990; Fabris & César 1996; Pereira & Zambom 1998; Pereira et al. 1998; Pereira & Araújo 2000; Assis et al. 2004; Fraga et al. 2007; Fabris & Peixoto 2013; Chagas et al. 2014; Saiter & Thomaz 2014; Sarnaglia Junior et al. 2014; Pertele et al. 2015), the species richness recorded to this state continues to be less well known than what is found for its neighboring states, whose floras are published in print or on the internet: Rio de Janeiro (< gov.br/consulta.php>), São Paulo (Wanderley et al. 2011), and some regional floras within Minas Gerais (Giulietti et al. 1987; Pirani et al. 2003). To date, the best compilation of plant species richness in ES is supplied by the Brazilian List of Plants and Fungi, henceforth Brazilian List (BFG 2015). The Brazilian List was continuously updated until early 2015 through a digital online platform. This database lists species information such as correct and accepted names and synonyms documented through vouchers or bibliographical references, habit, substrate and vegetation type, geographic distribution and endemism searchable fields. The INCT - Virtual Herbarium of Flora and Fungi (HVFF) launched through the SpeciesLink platform in 2009 (CRIA 2014), provides free and open access data and information regarding Brazilian plant collections. Together, these databases have become one of the most important tools for Brazilian researchers in plant taxonomy, and an important source of information for the development of the Flora do Espírito Santo project. This project aims to provide support for the establishment of regional plant family monographs in accordance with the Updated Global Strategies of Plant Conservation (GSPC-CDB). Considering the inconsistencies found between the Brazilian List and the HVFF databases: 1. voucher specimens (identified by experts) in herbaria bearing species names not found in the Brazilian List; 2. species listed as occurring in Espírito Santo in the Brazilian List without vouchers in the HVFF database; this study aimed to find and to correct inconsistencies between the two databases, in order to provide a more complete list and an accurate estimate of angiosperm species number in Espírito Santo. Methods The Espírito Santo State (ES) is located in Southeastern Brazil, bordering with Minas Gerais, Bahia, and Rio de Janeiro states (Fig. 1). A combination of two main databases, the Brazilian List (BFG 2015) and the INCT - Virtual Herbarium (HVFF, < was used to prepare the checklist. In the first instance, a filtered list of angiosperm species that occur in ES was requested from the Brazilian List project coordination. This initial list consisted of 5,304 records (subfamilies, infraspecific taxa, and hybrids were not included). Each record was checked against the INCT Virtual Herbarium database, where herbarium specimen data can be searched by using relevant parameters such as taxon name, geographical distribution, collector name and number. Therefore, a list of species (taxon name) that occur in a particular geographic location (a Brazilian state or municipality - ES in this case) can easily be accessed, and includes information about the hosting herbaria, identification authorship Figure 1 Location of Espírito Santo State, Brazil. Rodriguésia 66(4):

216 Angiosperm Checklist of Espírito Santo and date. We performed intensive and extensive exploration analyses covering all vouchers accessible at the HVFF database. Vouchers with provenance from ES and identified by specialists were prioritized, replacing the vouchers listed by the contributors to the Brazilian List, when these were not from ES. When a particular ES collected record could not be tracked for a given taxon name, the voucher cited in the Brazilian List was maintained. For vouchers listed in the Brazilian List that were not included in the HVFF, literature sources were added. In order to expand the available information, specific bibliography comprising species records from ES was searched. Taxonomic revisions and scientific articles that report new species from ES and published in the last 10 years were included in the search using the scientific literature databases ISI (< SCOPUS (< scopus.com/home.url>), Google Scholar (< scholar.google.com.br/>), Portal de Periódicos CAPES (< and JSTOR (< Species were listed alphabetically by family and genus (circumscription following APG (2009)), indicating endemic species ( ) and new records (*) to BFG (2015). Cultivated and species naturalized were highlighted by (!) and (+), respectively. Herbaria acronyms follow Index Herbariorum (Thiers, continuously updated). Results The revised Angiosperm list from ES comprises 6,364 species belonging to 1,390 genera and 180 families (see supplementary material < - DOI: / ) which 6,204 are native, 74 are cultivated, and 110 are naturalized. The richest families (Fig. 2) are Orchidaceae (744 spp.), Bromeliaceae (386 spp.), Fabaceae (384 spp.), Asteraceae (291 spp.), Myrtaceae (291 spp.), Rubiaceae (239 spp.), Melastomataceae (197 spp.), Apocynaceae (175 spp.), Cyperaceae (175 spp.) and Poaceae (174 spp.). The five richest genera are Solanum L. (Solanaceae - 96 spp.), Eugenia L. (Myrtaceae - 95 spp.), Vriesea Lindl. (Bromeliaceae 88 spp.), Myrcia DC. (Myrtaceae - 81 spp.), and Piper L. (Piperaceae - 74 spp.). According to the results obtained, there are 516 species endemic to Espírito Santo. Furthermore, ES houses 8.3% of the all endemic species 1147 Figure 2 Richest angiosperm families and their endemic species numbers from the state of Espírito Santo. registered for Brazil and 7.4% of all Atlantic Forest species. Bromeliaceae and Orchidaceae include the highest number of endemic species for the state (142 spp. and 80 spp., respectively), followed by Myrtaceae (33 spp.), Melastomataceae (30 spp.), and Araceae (23 spp.) (Fig. 2). Cryptanthus Otto & A.Dietr. and Alcantarea (E.Morren ex Mez) Harms (Bromeliaceae) showed the highest percentage of endemic species with 81% and 75%, respectively. Several common widespread taxa were not listed in the BFG (2015) as occurring in Espírito Santo, e.g. Gallesia integrifolia (Phytolaccaceae), Microsthachys bidentata (Mart. & Zucc.) Esser (Euphorbiaceae), and Cucumis anguria L. (Cucurbitaceae), despite their wide distribution throughout Brazil. Other examples are Chomelia oligantha Müll.Arg. (Rubiaceae) and Agarista eucalyptoides (Cham. & Schltdl.) G.Don (Ericaceae) that occur in adjacent areas of the bordering states of Bahia, Minas Gerais and Rio de Janeiro and were found to occur in ES. In addition, many examples of species first recorded for the Atlantic Forest represent new cases of disjunction between it and the Amazon Forest, such as Crepidospermum rhoifolium (Benth.) Triana & Planch (Burseraceae), and Fridericia patellifera (Schlecht.) L.G. Lohmann, the latter previously recorded for the Amazon Forest, Cerrado and Pantanal biomes. Furthermore, the known geographically distribution of a larger number of species was expanded by being recorded in ES (Appendix 1). Discussion and conclusion With an increase of 900 species over the number recorded for ES by the BFG (2015), the Rodriguésia 66(4):

217 1148 Dutra, V.F., Alves-Araújo, A. & Carrijo, T.T. present results suggest that ES is no longer the tenth state in species richness in the country (BFG 2015) but in reality it occupies the seventh position, overtaking the states of Goiás, Mato Grosso and Paraná. A simple area comparison makes it obvious that the ES is amongst the richest states in terms of plant diversity in Brazil, with a relatively small area (23rd amongst Brazilian states with an area of ,925 km 2 ) harbouring extremely high number of species. The state s plant richness has already been highlighted by Forzza et al. (2012) who stated that it holds 32% of the angiosperm richness from the Atlantic Forest. The number of angiosperm species registered by Fraga et al. (2007) and by the BFG (2015) for the ES were increased here by around 50% and 16%, respectively. The ranking of the Orchidaceae, Bromeliaceae, Fabaceae and Asteraceae for ES remains very similar to previous results reported for the Atlantic Forest by Stehmann et al. (2009), who reported Fabaceae in second and Bromeliaceae in third place. Despite the fact that Stehmann et al. (2009) highlight Orchidaceae, the richest family, as having the highest number of endemic species for the Atlantic Forest, partial results shown here for the state show that Bromeliaceae has more records of endemic species than Orchidaceae in ES. In more specific terms, it is known that the Atlantic Forest is the diversity centre of speciose genera such as Alcantarea and Cryptanthus (Martinelli et al. 2008) that are particularly expressive in ES, partially explaining the large number of endemic species for Bromeliaceae in this state. The complete documentation of the ES flora by the year 2020, in accordance to the goals of the Convention on Biological Diversity and the Global Strategy for Plant Conservation (CBD 2010), represents a big challenge. Geographical gaps in plant collections are notorious despite a reasonably well-scored species per area relationship. Exploration in such poorly-sampled areas is of strategic importance if we are to improve species documentation. In this sense, performing regional inventories is highly recommended to improve the quality of the knowledge of the state and consequently of the Flora of Brazil. The improvement of floristic knowledge is currently one of the greatest research challenges in Brazil. The present case study in the ES state establishes online plant databases as fundamental tools to record and retrieve species richness information. Considering that there are still many plant collections from Brazil that have not been databased and included in the available virtual herbaria, and that there are parts of the state that are less accessible and presently undercollected, it is most likely that the checklist of angiosperms in ES will need readjustments, with ongoing studies still revealing many further records. Acknowledgements We are grateful to Dra. Rafaela Forzza for supporting and encouraging us from the early stages to the completion of this study; to the coordination and team of the Brazilian List by providing relevant data about ES; to FAPES and CNPq for financial support of floristic surveys that allowed the collection of material in poorly sampled areas; to Dr. Mário L. Garbin, Dra. Daniela Zappi and two anonymous reviewers for their helpful comments and suggestions. References Aguiar, A.P.; Chiarello, A.G.; Mendes, S.L. & Matos, E.N Os Corredores Central e da Serra do Mar na Mata Atlântica brasileira. In: Galindo-Leal, C. & Câmara, I.G. (eds.). Mata Atlântica: biodiversidade, ameaças e perspectivas. Fundação SOS Mata Atlântica, Conservação Internacional e Centro de Ciências Aplicadas à Biodiversidade, Belo Horizonte. Pp Almeida, R.F. & Mamede, M.C.H Checklist, conservation status, and sampling effort analysis of Malpighiaceae in Espírito Santo State, Brazil. Brazilian Journal of Botany: 37: Alves, M. & Luz, C.L Juncaceae. In: Lista de espécies da flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < gov.br/jabot/floradobrasil/fb8084>. Access on 19 June Amaral, M.C.E.; Pellegrini, M.O.O. & Lima, C.T Nymphaeaceae. In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < FB10937>. Access on 12 October APG An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. Botanical Journal of the Linnean Society 161: Araujo, A.O.; Chautems, A. & Ferreira, G.E Gesneriaceae. In: Lista de espécies da flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < FB119>. Access on 19 June Barthlott, W. & Taylor, N.P Notes towards a monograph of Rhipsalideae (Cactaceae). Bradleya 13: Rodriguésia 66(4):

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221 1152 Dutra, V.F., Alves-Araújo, A. & Carrijo, T.T. Available at < floradobrasil/fb10909>. Access on 28 June Saiter, F.Z. & Thomaz, L.D Revisão da lista de espécies arbóreas do inventário de Thomaz & Monteiro (1997) na Estação Biológica de Santa Lúcia: o mais importante estudo fitossociológico em florestas montanas do Espírito Santo. Boletim do Museu de Biologia Mello Leitão 34: Sarnaglia Junior, V.B.; Thomaz, L.D. & Guimarães, E.F O gênero Peperomia Ruiz & Pav. na Área de Proteção Ambiental do Mestre Álvaro, Espírito Santo, Brasil. Boletim do Museu de Biologia Mello Leitão 35: Sano, P.T.; Giulietti, A.M.; Costa, F.N.; Trovo, M.; Echternacht, L.; Tissot-Squalli, M.L.; Watanabe, M.T.C.; Hensold, N.; Andrino, C.O. & Parra, L.R Eriocaulaceae. In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < floradobrasil/fb7548>. Access on 9 October Secco, R.; Cordeiro, I. & Martins, E.R Phyllanthaceae. In: Lista de espécies da flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < floradobrasil/fb24166>. Access on 8 June Simpson, B Krameriaceae. In: Lista de espécies da flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < floradobrasil/fb140>. Access on 19 June Sobral, M Musaceae. In: Lista de espécies da flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < floradobrasil/fb85800>. Access on 28 June Sobral, M.; Proença, C.; Souza, M.; Mazine, F. & Lucas, E Myrtaceae. In: Lista de espécies da flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < floradobrasil/fb10275>. Access on 28 June Somner, G.V.; Ferrucci, M.S.; Acevedo-Rodríguez, P.; Perdiz, R.O.; Coelho, R.L.G. & Medeiros, H Sapindaceae. In: Lista de espécies da flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < FB2016>. Access on 10 October Souza, V.C. & Hassemer, G Plantaginaceae. In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < jbrj.gov.br/jabot/floradobrasil/fb191>. Access on 11 October Standley, P.C Rubiaceae. In: Dahlgren, B.E. (ed.). Studies of american plants VI. Field Museum of Natural History, Botanical series 8: Stefano, M.V.; Calazans, L.S.B. & Sakuragui, C.M Meliaceae. In: Lista de espécies da flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < floradobrasil/fb19740>. Access on 28 June Stehmann, J.R.; Forzza, R.C.; Salino, A.; Sobral, M.; Costa, D.P. & Kamino, L.H.Y Plantas da Floresta Atlântica. Jardim Botânico do Rio de Janeiro, Rio de Janeiro. 516p. Stehmann, J.R.; Mentz, L.A.; Agra, M.F.; Vignoli- Silva, M.; Giacomin, L. & Rodrigues, I.M.C Solanaceae. In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < floradobrasil/fb225>. Access on 10 October Tabarelli, M.; Silva, J. & Gascon, C Tropical forest fragmentation, synergisms and the impoverishment of neotropical forests. Biodiversity and Conservation 13: Taylor, N.; Santos, M.R.; Larocca, J. & Zappi, D Cactaceae. In: Lista de espécies da flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < floradobrasil/fb1733>. Access on 8 June Thiers, B. [continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden s Virtual Herbarium. Available at < org/ih/>. Access on 5 May Walter, B.M.T Hydroleaceae. In: Lista de espécies da flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < jbrj.gov.br/jabot/floradobrasil/fb132>. Access on 19 June Wanderley, M.G.L.; Shepherd, G.J.; Martins, S.E.; Estrada, T.E.M.D.; Romanini, R.P.; Koch, I.; Pirani, J.R.; Melhem, T.S.; Harley, A.M.G.; Kinoshita, L.S.; Magenta, M.A.G.; Longhi- Wagner, H.M.; Barros, F.; Lohmann, L.G.; Amaral, M.C.E.; Cordeiro, I.; Aragaki, S.; Bianchini, R.S. & Esteves, G.L Checklist das Spermatophyta do Estado de São Paulo, Brasil. Biota Neotropica 11: Zappi, D Crassulaceae. In: Lista de Espécies da Flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < floradobrasil/fb95>. Access on 10 October Zappi, D.; Guimarães, E.F.; Manoel, E.A. & Siqueira, C.E Loganiaceae. In: Lista de espécies da flora do Brasil. Jardim Botânico do Rio de Janeiro. Available at < floradobrasil/fb8663>. Access on 19 June Artigo recebido em 09/08/2015. Aceito para publicação em 23/10/2015. Rodriguésia 66(4):

222 Rodriguésia 66(4): Epílogo DOI: / Epilogue The huge impact of the publication of the Brazilian Catalogue of Plants and Fungi in 2010 spread further from the plant taxonomy arena, influencing the stability of nomenclature from biodiversity surveys to ecology and conservation research and also becoming the reference for plant names within other areas. The Lista de Espécies da Flora do Brasil (< or Brazilian List, answers to the demand for plant species name certification coming from customs, policy making, legislation and above all, the scientists and general public. The species list became the reference for floristic and plant survey studies in Brazil and is currently used by ecologists, foresters, agronomists and geographers. Since 2010, the website has received over 1.5 million visits from half a million users hitting over 13 million pages, remaining in the site in average for 12 minutes. Herbarium curators and the users of plant collections who need to check plant names and their occurrence within Brazil can count on this list as their main place for consultation. Formerly dominant sites, such as Missouri Botanical Gardens ( and International Plant Names Index ( are nowadays secondary to the Brazilian List for researchers concerned with Brazilian plants. The stabilization of nomenclature and provision of a taxonomic backbone has impacted positively towards the country s contribution for species conservation status assessments (GSPC Target 2) carried out by the Centro Nacional de Conservação da Flora/CNCFlora. The improved data quality has also positively impacted individual researchers and herbaria, since it provides a secure source for correct and accepted names. Other consequence of the project was to bring evidence to poorly known regions where botanical collections are still needed in order to provide a more complete picture of the regional plant diversity. Increasing the knowledge about the species distribution area is important to develop a framework for Brazilian state floras and checklists. The project has led to a resurgence of plant taxonomy as a worthwhile subject to be studied and consequent increase of the number of contributors from 416 in 2010 to 575 in The Brazilian government recognition of taxonomic activities was re-kindled by the whole process and the Reflora Programme was launched. This two pronged programme includes scholarships for Brazilian botanists to visit collections abroad (initially Royal Botanic Gardens, Kew/RBG Kew and the Museum Nationelle d Histoire Naturelle, Paris/NHM Paris) and also funds, through the State Research Councils (FAPs) and the private sector, the repatriation of herbarium data and images to compose a virtual herbarium, Herbário Virtual Reflora (< An unprecedented number of individual and collective bursaries have generated a high and varied output of scientific work based on historical botanical collections. Together with the images provided by the INCT - Herbario Virtual da Flora e dos Fungos, another government funded programme, the Brazilian List was populated by images of vouchers in due course. While the Catalogue published in 2010 included fundamental features concerning accepted species, bibliographic reference, biome, distribution and endemism, and endeavoured to include a voucher specimen for each species and infraspecific taxon cited, it lacked a facility to include the images of the specimens listed. From March 2013 the Brazilian List on-line was migrated into the server of the Rio de Janeiro Botanical Gardens under the umbrella of the Reflora Project, alongside with the Virtual Herbarium Reflora, launched in September Amongst the new functionalities featured, it became possible to include the life-form, substrate and vegetation type. The rapid growth and linking of the virtual herbaria to the Brazilian List meant that images of vouchers (including type-specimens) could be added to the system, and researchers were also welcome to link images of plants in the field, making the list more user-friendly for non-botanists and for researchers based far away from the collections. Georeferenced herbarium records from the Reflora Virtual Herbarium and specieslink Network (< also helped to suggest possible new state records and refine the endemism category, but these are to be used cautiously as the data are not always accompanied by images and their determination might in some cases be inaccurate.

223 The final edition of the Brazilian List Project aims to mark this particular step in the process of recording the known Brazilian plant diversity prior to its transformation and upgrade into a new project, the Brazilian Flora Online 2020, aiming to describe and differentiate among the known species. Much work has been required to get to this point, but the fundamental shift in the involvement and focus demanded by large-scale floristic descriptive work imposes an intensification and concentration around several new issues, for instance: to design new functionalities to include descriptive and identification key data taking advantage of the widely successful basic Brazilian List framework; to create a system that makes it possible to compare between descriptions to indicate where information is lacking and when all the species of a given genus have been described; to gauge how many species a single researcher can describe in 4 years, considering that there are groups where monographs already exist, while the largest genera, such as Eugenia, Mimosa, Croton and Paepalanthus still lack in-depth studies to permit their differentiation beyond large species groups. Such issues meant that a fundamental change in the authorship system had to be effected in order to provide a democratic, effective and meritorious system of attribution of responsibility and authorship, and to create a less top-heavy, more dynamic way for the contributors (including the provision of opportunities to new players) to add descriptions to the system. In order to achieve this objective, researchers bid individually or as research-groups to complete taxonomic plant groups comprising up to 300 species within a system that aims to adhere to strict deadlines. The final numbers for the Brazilian List Project 2015 are shown below. It is an excellent moment to take stock and consider the difference made by the huge effort involved in holding an on-line updatable system to register plant biodiversity coordinating the work of 575 contributors and making it instantly available for the botanical community and public worldwide Species total Endemics % Fungi 5, Algae 4, Bryophytes 1, Ferns and Lycophytes 1, Gymnosperms 30 (23*) Angiosperms 32,831 (32,026*) 18, Total 46,097 19,503 43,1 *native species (endemism percentage calculated based on native species) Daniela C. Zappi, Rafaela Campostrini Forzza, Vinicius Castro Souza, Vidal de Freitas Mansano e Marli Pires Morim Acknowledgements: