January 2022 Wetland Science & Practice by Society of Wetland Scientists

Wetland Science Practice

INCUDES A FOCUS ON LATIN AMERICAN WETLANDS

published by the Society of Wetland Scientists

Vol. 40, No. 1 January 2022 ISSN: 1943-6254

FROM XXXXXXTHE EDITOR’S DESK Greetings to all and Happy New Year! I’m looking forward to this year with the hope that we can get back to normal and do a bit more travelling including to the SWS Conference in Grand Rapids in May, although the threat of more Covid impacts remains. My wife and I did get to travel some in 2021 and look forward to getting together with more friends in 2022. Besides visits with family, we took two memorable trips this past year – one was the trip to Maine that was the source of my Notes from the Field for the October issue of WSP. The Ralph Tiner other was a cycling trip to Greek WSP Editor islands which was an experience as we rode with some effort up mountains but flew down around the S-curves and more carefully around hairpins. At the end of the day, we did have a sense of accomplishment and could eat whatever we wanted without guilt. This issue of WSP is a special one dedicated to publishing articles about research and conservation activities involving Latin American wetlands (see our introduction to that section). One exception is an article that summarizes presentations from the Ramsar Section’s symposium at our annual (virtual) conference on challenges facing international wetlands. The issue also contains some SWS news, especially an announcement about our 2022 annual meeting that will be held as part of the Joint Aquatic Sciences Meeting (JASM) in Grand Rapids, Michigan, May 16-20 – a call for papers. An earlier announcement was published on the SWS website. We offer a special thanks to all our contributors for WSP issues in 2021 as well as to those for the current issue. We hope that wetland scientists, managers, and other folks with an interest in wetlands will continue to submit articles and information for publication in 2022. Perhaps our chapters or sections would like to have their members prepare articles on wetlands in their geographic area or area of interest. Yes…this will require some effort, but I’m sure that this information would be of great interest and worth the effort. Please contact me if you have any questions; general guidelines appear in the back of each issue. Meanwhile, Happy Swamping! n

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CONTENTS Vol. 40, No. 1 January 2022 ISSN: 1943-6254 4 / President's Address 5 / Webinars 6 / SWS News ARTICLES 7 / After Fifty Years of Ups and Downs, What is Needed for International Wetland Conservation to Become a Relevant Force for the Challenges of the Future? Nick C. Davidson, Robert J. McInnes, and others. 15 / Introduction to This Special Issue Ralph Tiner and Tatiana Lobato-de Magalhães 16 / Juncus in Northern Patagonian Wetlands Adriel I. Jocou and Ricardo Gandullo 26 / From Marsh to Swamps: Vegetation Gradient Linked to Estuarine Hydrology Hugo López Rosas and Patricia Moreno-Casasola 35 / Aquatic Groups in Freshwater Systems in the Semidesier to Queretano, Mexico: Algae, Bryophytes, Vascular Plants, and Odonata Patricia Herrera-Paniagua, Mahinda Martínez, and others 48 / Veredas: Upper Basin Wetlands Feed the Pantanal Year-round Arnildo Pott, Vali Joana Pott, and Suzana Neves Moreira 51 / Tolerance and Recovery Capacity of Tropical Native Tree Species to Water and Salt Stress: An Experimental Approach for Wetland Rehabilitation Wilmer O. Rivera-De Jesús and Elsie Rivera-Ocasio 59 / Experiences in the Restoration of Tropical Freshwater Swamps Patricia Moreno-Casasola, Hugo López Rosas, and others 70 / Assessment and Perceptions of the Environmental Quality of the Urban and Suburban Wetlands of Leticia (Amazonas, Colombia) María J. Arias-B, Jhon Ch. Donato- Rondón, and others

Wetland Science Practice 82 / Effective Conservation and Good Governance at the Ramsar Site Bahía Lomas, Tierra del Fuego, Chile Carmen Espoz, Ricardo Matus, and others 86 / Water and Wetlands Outreach Project in the Sierra Gorda Biosphere Reserve, Mexico Tatiana Lobato-de Magalhães, Marinus L. Otte, and others 92 / The Pan-American Treatment Wetland Network, HUPANAM: A Brief Sketch of the Treatment Wetland Research Groups in the Latin-American and Caribbean Region M.A. Rodriguez-Dominguez, P.H. Sezerino, and C.A. Arias 98 / Cuatrociénegas Wetlands Coahuila de Zaragoza, Mexico 100 / Notes from the Field 113 / Wetlands in the News 114 / Wetlands Bookshelf 115 / WETLANDS 116 / About WSP/Submission Guidelines

COVER PHOTO: Brackish forested wetland dominated by Pterocarpus officinalis in Rio Guajataca, Quebradillas Municipality, Puerto Rico. (Photo by Elsie Rivera-Ocasio)

SOCIETY OF WETLAND SCIENTISTS 1818 Parmenter St., Ste 300, Middleton, WI 53562 (608) 310-7855 www.sws.org Social icon

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Note to Readers: All State-of-the-Science reports are peer reviewed, with anonymity to reviewers.

PRESIDENT / Gregory Noe, Ph.D. PRESIDENT-ELECT / William Kleindl, Ph.D. IMMEDIATE PAST PRESIDENT / Loretta Battaglia, Ph.D. SECRETARY GENERAL / Leandra Cleveland, PWS TREASURER / Lori Sutter, Ph.D. EXECUTIVE DIRECTOR / Erin Berggren, CAE DIGITAL MARKETING SPECIALIST / Moriah Meeks WETLAND SCIENCE & PRACTICE EDITOR / Ralph Tiner, PWS Emeritus CHAPTERS ASIA / Wei-Ta Fang, Ph.D. CANADA / Susan Glasauer, Ph.D. CENTRAL / Tim Fobes, PWS CHINA / Xianguo Lyu EUROPE / Matthew Simpson, PWS INTERNATIONAL / Ian Bredlin, Msc; Pr.Sci.Nat and Tatiana Lobato de Magalhães, Ph.D., PWS MID-ATLANTIC / Jason Traband, PWS NEW ENGLAND / Dwight Dunk, PWS NORTH CENTRAL / Casey Judge, WPIT OCEANIA / Phil Papas PACIFIC NORTHWEST / Josh Wozniak, PWS ROCKY MOUNTAIN / Rebecca Pierce SOUTH ATLANTIC / Brian Benscoter, Ph.D. SOUTH CENTRAL / Jodie Murray Burns, PWS, MEd, MS WESTERN / Richard Beck, PWS, CPESC, CEP SECTIONS BIOGEOCHEMISTRY / Beth Lawrence, Ph.D. EDUCATION / Darold Batzer, Ph.D. GLOBAL CHANGE ECOLOGY / Wei Wu, Ph.D. PEATLANDS / Bin Xu, Ph.D. PUBLIC POLICY AND REGULATION / John Lowenthal, PWS RAMSAR / Nicholas Davidson, Ph.D. STUDENT / Steffanie Munguia WETLAND RESTORATION / Andy Herb WILDLIFE / Andy Nyman, Ph.D. WOMEN IN WETLANDS / Jennifer Karberg, Ph.D. COMMITTEES AWARDS / Siobhan Fennessy, Ph.D. EDUCATION AND OUTREACH / Jeffrey Matthews, Ph.D. HUMAN DIVERSITY / Kwanza Johnson and Jacoby Carter, Ph.D. MEETINGS / Yvonne Vallette, PWS MEMBERSHIP / Leandra Cleveland, PWS PUBLICATIONS / Keith Edwards WAYS & MEANS / Lori Sutter, Ph.D. WETLANDS OF DISTINCTION / Roy Messaros, Ph.D. Bill Morgante, Steffanie Munguia and Jason Smith, PWS REPRESENTATIVES PCP / Scott Jecker, PWS WETLANDS / Marinus Otte, Ph.D. WETLAND SCIENCE & PRACTICE / Ralph Tiner, PWS Emeritus ASWM / Jill Aspinwall AIBS / Dennis Whigham, Ph.D.

PRESIDENT'S ADDRESS Fellow Wetlanders,

Gregory B. Noe, Ph.D. Florence Bascom Geoscience Center, U.S. Geological Survey SWS President

One perk of being President of SWS is exposure to all of the good people who volunteer to improve the society; its activities, its knowledge base, and its benefits to members. It’s humbling and amazing to be a part of our collective community. As COVID-19 still continues to harm people and change our daily lives, I’d like to take the opportunity to highlight just a few of the good things that we are doing.

Our Chapters of SWS have been tremendously busy hosting virtual conferences to advance wetland science and management. I’d like to mention three: the Asia Chapter held the “International Wetland Convention”, the Oceania Chapter held the “Fire in Wetlands Forum”, and the International Chapter held their firstever African Region meeting. These are notable not only for their information content and opportunity for virtual networking, but also because of the continuing globalization of SWS’ activities. Many of our other Chapters also have been busy hosting virtual meetings too. Looking forward, SWS is one of nine scientific societies that is hosting the 2022 Joint Aquatic Sciences Meeting (JASM). We won’t be holding our own standalone annual meeting. Instead, we’ll hold our traditional and beloved SWS-specific activities during JASM. So while JASM is certain to have stellar scientific content, many SWS volunteers are laboring to ensure that the meeting meets our own needs. We’ll hold our usual SWS Chapter and Section annual meetings, Awards banquet, Business Meeting, SWaMMP mentoring, and much more. While the meeting is being built as a hybrid meeting style, with videos of all scientific presentations available online, the emphasis

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(and hope!) is that most participants will be there in person in Grand Rapids, Michigan U.S. We know this will be a ‘can’t miss’ meeting and we look forward to seeing you there or virtually! A strategic emphasis that SWS is currently working to bolster is our benefits to early career professionals. The Education and Outreach Committee is currently developing several new exciting initiatives. The Committee is currently finalizing an Early Career Networking program that will include a fun event at JASM. It also is developing two mentoring initiatives: Joint Mentoring with the Association of State Wetland Managers; and the International Mentoring Program. While we develop these programs, please reach out to us with your suggestions on what you want them to do or to volunteer to help. What mentoring do you need? The Education Section also has been very busy developing a new effort to provide online educational materials, termed “Foundations of Wetland Science.” This program will provide the general public, students, instructors, and professionals with online educational modules that can be freely accessed to gain general knowledge about wetland science, or be used as resources in the classroom and for various outreach activities. If you would like to contribute, please join the Section and participate in their annual meeting at JASM where final planning will occur. For SWS to grow and sustain its role as the professional home for wetland professionals, we need to have worthy benefits available to our members during the arc of their careers from students, to early career professionals, established professionals, and retired experts. If you’d like to become more involved in these rewarding efforts, please reach out and we’ll engage you with a good opportunity. And let your colleagues know about all that SWS offers! With appreciation, Greg Noe SWS President (2021-2022); gnoe@usgs.gov

WEBINARS

SOCIETY WETLAND SCIENTISTS

ENGLISH: January 21 | 1:00 PM ET

November 17 | 1:00 PM ET

Student Section Juried Student Presentation

Wetland identification in ecologically challenging and high-stakes interface environments Julia Alards-Tomalin

December 15 | 1:00 PM ET

February 17 | 1:00 PM ET

SPANISH:

Cat Branch Stream and Wetland Restoration Restoration of a degraded, urban stream system into a complex of floodplain and wetlands using baseflow channel and valley restoration techniques. Nasrin Dahlgren March 17 | 1:00 PM ET

USACE AJD Requests and Waters of the U.S. definition Jonathan G. Barmore April 21 | 1:00 PM ET

Constructed wetlands for (waste)water treatment: types and use for various wastewaters Jan Vymazal

Retrospective of Research Lifetime Achievement Recipient

January 12 | 1:00 PM ET

The Central American Waterbird Count: the first ten years / El Censo Centroamericano de Aves Acuáticas: los primeros diez años Dr. John van Dort and Arne Lesterhuis March 23 | 1:00 PM ET

Hydrologic model of sediment transport: a tool applied to recover and manage Santa Marta wetlands, Colombia/ Modelo hidrosedimentológico de la Ciénaga Grande de Santa Marta, Colombia, herramienta para su recuperación y manejo Dr. Constanza Ricaurte-Villota June 22 | 1:00 PM ET

More info coming soon

Wetland Connectivity & Podcast series HumMentor Mentees (Flor, Cynthia, Adad)

July 21 | 1:00 PM ET

September 21 | 1:00 PM ET

June 16 | 1:00 PM ET

August 18 | 1:00 PM ET

Biodiversity and ecosystem functioning of coral reefs: a holistic study for their conservation from microorganisms to ecosystem Dr. Fabián Alejandro Rodríguez Zaragoza

More info coming soon

December 12 | 1:00 PM ET

Karst wetlands in the Yucatan Peninsula Eduardo Cejudo

September 15 | 1:00 PM ET

More info coming soon October 20 | 1:00 PM ET

The Central American Waterbird Count: the first ten years / El Censo Centroamericano de Aves Acuáticas: los primeros diez años Dr. John van Dort and Arne Lesterhuis

More info coming soon

Wetland Science & Practice January 2022 5

SWS NEWS SWS Member Dr. Muhammad Afzal Receives Nature Research Award Muhammad Afzal, Principal Scientist at the National Institute for Biotechnology and Genetic Engineering (NIBGE), Faisalabad, Pakistan, was chosen as the runner-up for Nature Research Awards for Driving Global Impact . He is the first Asian scientist to win this prestigious award which includes a cash prize of SWS Member Dr. $10,000 in recognition for his Muhammad Afzal work addressing polluted soil and wastewater through self-sustaining and environmentfriendly technologies. Muhammad is a member of Society of Wetland Scientists and was just certified as SWSPCP’s very first WPIT (Wetland Professional in Training) from Pakistan. Previously he was awarded a Gold Medal in Earth and Environmental Sciences by the Pakistan Academy of Sciences. Muhammad has pioneered the “Floating Treatment Wetland” (FTW) technology in Pakistan at the field-scale for the treatment of sewage and industrial wastewater. In Pakistan, more than 95% of wastewater is discharged untreated into the environment, contaminating groundwater, surface water, soil, and the food chain. This contamination contributes to 80% of morbidity and 40% of mortality in Pakistan. One of the main reasons for the discharge of

untreated wastewater is the high capital and operational costs of conventional wastewater treatment technologies. To counter this problem, NIBGE has developed a cheaper, ecologically safe, and indigenous FTW technology, also known as floating gardens and floating parks. Indigenous wetland plants were screened to develop FTWs over a locally designed and fabricated buoyant mat. Bacteria were isolated and characterized and applied in FTWs to enhance their wastewater treatment efficiency. This method proved to be 500 times cheaper than currently available floating treatment systems in the international market, saving millions on imports of material required for the development of FTWs. Moreover, the design and fabrication of floating mat done locally made it possible to apply FTWs more widely in Pakistan. Since 2014, floating treatment wetlands technology and constructed wetlands have been applied at more than 20 sites in Pakistan to improve the quality of ~700 million m3 of wastewater. As an additional benefit, these wetlands are now used by many species of birds for nesting. We offer our congratulations to Dr. Afzal on receiving this award. His entry was one of 350 applications from 62 countries considered for the award.

Upcoming International Chapter Meeting for Latin America and the Caribbean

SWS Student Mentoring Program for Latin America and the Caribbean - the HumMentor

Join us for the first-ever International Chapter Meeting for Latin America and the Caribbean (1-day event, virtual option) – February 9 at 10AM (CST). Register your interest here. We will contact you soon with more information about the confirmed schedule and speakers. Thank you so much for being so interested! Are you not a member? Please join the SWS, see more information here. Following us @SWSLAC 6 Wetland Science & Practice January 2022

Nature Research Awards are given out by the publisher Springer Nature to support researchers for their work addressing challenges across healthcare protection, environmental intervention and food/water security.

HumMentor is a mentoring program sponsored by the Society of Wetland Scientists (SWS) for senior undergraduate and early graduate students from Latin America and the Caribbean (LAC) countries who are conducting research or scientific outreach in wetland science. Its goals are: 1) to stimulate and promote wetland science education in LAC countries, 2) to encourage the publication of collaborative wetland research from LAC countries, and 3) to grow a network of wetland scientists and practitioners across Latin American and the Caribbean. The next call for applications in 2022 will open on World Wetlands Day on February 2, 2022. Visit our web page and learn more about the program.

INTERNATIONAL WETLANDS CONSERVATION

After Fifty Years of Ups and Downs, What is Needed for International Wetland Conservation to Become a Relevant Force for the Challenges of the Future? Nick C. Davidson1,2, Robert J. McInnes2,3, Gillian T. Davies4,5, Matthew Simpson2,6 and C. Max Finlayson2,7

INTRODUCTION This article has been developed from the presentations made in the Society of Wetland Scientists (SWS) Ramsar Section’s symposium during the SWS virtual conference on December 3, 2020. THE CRISIS FACING WETLANDS AND WETLAND GOVERNANCE Despite 50 years of global governments' commitments to delivering wetland conservation and wise use, and stemming the loss and degradation of wetlands, we face an accelerating biodiversity crisis and a climate crisis and none of the world's twenty 2020 Aichi Biodiversity Targets have been delivered (Convention on Biological Diversity 2020). This is very much the case for wetland biodiversity, as shown through the Global Wetland Outlook (Ramsar Convention 2018), global citizen science assessments of the world’s wetlands (McInnes et al. 2020; McInnes et al. 2021), and an analysis of information contained in national reports submitted by Contracting Parties to the Convention (Davidson et al. 2020). Similarly, given the importance of wetlands for storing carbon, their alarming loss and mismanagement is contributing to increased carbon emissions and contributing to the climate crisis, while also being highly vulnerable to changes in the climate (Moomaw et al. 2018a). The data that are now available confirm that existing policy and governance are not sufficient to stem wetland loss and degradation. Finlayson and Gardner (2020) contend that “A switch from documenting the change in wetland biodiversity towards more emphasis on taking decisions is needed to implement effective responses and reverse the negative trends for wetlands.” and “…that Nick Davidson Environmental, Wigmore, UK Institute for Land, Water and Society, Charles Sturt University, Albury, NSW, Australia 3 RM Wetlands & Environment Ltd, Littleworth, Oxfordshire, UK 4 BSC Group Inc., Worcester, Massachusetts, USA 5 Global Development and Environment Institute, Tufts University, Medford, Massachusetts, USA 6 35percent, Stroud, UK 7 IHE Delft, Institute for Water Education, Delft, Netherlands 1 2

failure to place greater emphasis on effective responses could lead to the [Ramsar] Convention becoming an irrelevant force for the wise use of wetlands.” This is a harsh critique from authors who have over many years ardently supported the Ramsar Convention. Given these multiple lines of evidence something needs to change - continuing "business as usual" is not an option. The data speak for themselves – the condition of wetlands globally is dire, and getting worse. Since 1971 the Ramsar Convention has established itself as the main international voice for the conservation and wise of wetlands. As at 25 November 2021, it has 171 national governments as Contracting Parties. These governments have met on 13 occasions to consider the wise use of wetlands and agree formal decisions covering administrative and technical issues. There are now 2,434 Wetlands of International Importance (Ramsar Sites) covering 254,618,717 ha (as at 25 November 2021). However, as shown by the data from multiple sources, this delivery of wetland wise use is not enough – wetland loss and degradation has not been stopped, let alone reversed. Approaching the 50th anniversary of the signing of the text of the Convention in the Iranian city of Ramsar in February 1971 we took the opportunity of the Society of Wetland Scientists (SWS) Virtual Conference (1-3 December 2020) to examine what has gone wrong, and why. We then outlined a number of new approaches and paradigms for better addressing efforts for achieving the wise use of wetlands, including the new SWS initiatives on the Rights of Wetlands (Simpson et al. 2020; Fennessy et al. 2021) and on Climate Change and Wetlands (Finlayson et al. 2020a), and how SWS can respond to the "Statement of World Aquatic Scientific Societies on the Need to Take Urgent Action against Human-Caused Climate Change". INTERNATIONAL AND NATIONAL WETLAND GOVERNANCE – WEAKNESSES AND DEVELOPING A ROADMAP FOR CHANGE We face a recognised global biodiversity crisis. Wetlands are not exempt. In 1971, 50 years ago, the Ramsar Convention on Wetlands was established by governments because of then increasing concerns over wetland loss and degradation – and its impacts on wetland-dependent species. But since 1970 the area of wetlands has progressively continued to decline, through deliberate drainage and conversion, in all parts of the world (Darrah et al. 2020). Deterioration in the state of our remaining wetlands is becoming progressively more widespread, including for designated Wetlands of International Importance (Ramsar Sites) (Davidson et al. 2020). Populations of freshwater (inland wetland-dependent) species have declined since 1970 by 84% - far more than species depending on other biomes (WWF 2020). Wetland Science & Practice January 2022 7

The world’s governments failed to meet any of their twenty 2020 Aichi Targets for biodiversity, including for wetlands (Convention on Biological Diversity 2020). Nor are we on track to deliver the 2030 UN Sustainable Development Goals (SDGs) for wetlands. Yet governments, including through their intergovernmental processes, seem to be just continuing with “business as usual”. In 2022 the same governments are now preparing to agree yet another suite of biodiversity goals and targets, this time for 2030. Their draft 2030 targets are similar to previous targets. Despite some successes for wetlands, and increasing recognition of the benefits and value of wetlands to people and their livelihoods (Davidson et al. 2019), overall sectoral nature conservation actions and protected area approaches have not delivered sufficiently to stem the loss and degradation of wetlands worldwide. They will continue to fail if the drive for economic growth, rather than truly sustainable development (Bradshaw et al. 2021), continues to over-ride efforts to achieve wetland wise use. This begs the question as to whether the intergovernmental agreements such as the Ramsar Convention on Wetlands are any longer “fit-for-purpose”? There is now an urgent need to change mindsets and approaches and develop new paradigms finally to begin to achieve wetland wise use. FROM NATURE CONSERVATION TO THE UTOPIA OF GREEN CAPITALISM - HOW TO SHIFT FROM THE MOLLIFYING AND ANODYNE REBRANDING OF THE ‘WISE USE OF WETLANDS’ TO GENUINE ACTION Wise use is the central tenet of the Ramsar Convention. Defined as the “maintenance of ecological character, achieved through the implementation of ecosystem approaches, within the context of sustainable development”, ‘Wise Use’ can be considered as the brand slogan for the Ramsar Convention, in the way that “Think different” is to technology or “Just do it” is to sportswear. Unfortunately, given the plight of wetlands reported in this paper, the Ramsar Convention has been considerably less effective than Apple or Nike in promoting and, more concerningly, protecting its product. In the marketing world, ‘Ramsar’ can be considered as the brand, ‘wetlands’ the product and ‘wise use’ the slogan or strapline. The brand name should give the product an identity whilst the slogan should play an important role in protecting the brand identity (Abdi and Irandoust 2013). Every good marketing executive knows that a slogan should aim to be timeless. The slogan (wise use), in contrast to the brand name (Ramsar), is capable of telling where the brand is going (Kohli et al. 2007). This could not be more apposite for wetlands as non-wise use has left them in the parlous state 8 Wetland Science & Practice January 2022

they find themselves in today – without wise use there will be very few wetlands remaining in the future. Rather than emphasising and promoting the slogan (wise use) and forming strong links between the slogan and the product (wetlands), the slogan has taken a backseat whilst the name and role of the product has continually changed in an attempt to appeal to differing and emerging audiences. Since the Ramsar Convention came into being in 1971, wetlands, and the language around wise use, has undergone continuous and evolving rebranding attempts. Terms such as natural or green infrastructure (Kumar et al. 2017; Stefanakis 2019), blue carbon (Sheehan et al. 2019), nature-based solutions (Thorslund et al. 2017), natural capital (Cahoon and Guntenspergen 2010) and natural climate solutions (Griscom et al. 2017), have been thoughtup and rolled out in successive rebranding attempts, each time stressing a new version of the value proposition that wetlands offer. The messaging between the type of product (wetlands) and what it delivers (the value proposition) has become confused. Wise use has been supplanted by a variety of slogans each seeking to reposition wetlands to find favour with a certain audience. Meanwhile wetlands have continued to be degraded and lost. One of the main human connections with wetlands is their aesthetical qualities (Buell 2005; Do and Kim 2020). From John Constable’s landscape paintings of rural England to the sun-kissed beaches and coral lagoons of holiday companies’ promotional materials, we can all picture wetlands in our minds. However, one of the challenges faced by wetlands as a brand is their diversity and the vast contrasts in their visual elements. Urban ponds in London, tropical coral reefs fringing the Maldives and swamp forests of Amazonia are visually and eudaemonically very different products. This presents significant challenges if the marketing goal is to achieve the delivering of visual unity within diversity (Phillips et al. 2014). Nevertheless, all humans, both as individuals and collectively, have relationships with wetlands. However, confusion has arisen through the rebranding of wetlands, often because the focus has been primarily on the hedonic and moral values (see Costanza et al. 1989; Richardson 1994; Mitsch and Gosselink 2000; Ghermandi et al. 2010; Davidson et al. 2019) and an attempt to commodify wetlands within capitalist nature (O’Connor 1994; Robertson 2000) and the lack of attention to the relational or eudaemonic values. Positive relational values offer important opportunities for the evolution of values that may be necessary for transformative change towards wetland wise use (Chan et al. 2018). However, the eudaemonic or relational values which genuinely connect people to wetlands remain for the main part overlooked (van den Born et al. 2018). The

Ramsar Convention (the brand), and its various agencies and acolytes, needs to rebuild and stress the link between the aesthetic or visual qualities of wetlands (the product) through the eudemonic or relational values (wise use). The focus should shift from the hedonic or moral values to a promotion of the meaning of wetlands to people rather than jumping on the latest bandwagon or buzzword. Under this construct wise use becomes the delivery vehicle for the maintenance of the meaning that wetlands give to peoples’ lives, a meaning that can extend beyond the economic or moral. A SHIFT IN PARADIGMS – NEW WAYS OF RECOGNISING OUR RELATIONSHIP WITH WETLANDS The world is at a tipping point, as the World Scientists’ Warning to Humanity: A Second Notice (Ripple et al. 2017) starkly warns us. David Attenborough (2020) uses his own close to century-long lifetime (94 years) as illustration. When he was 11 years old in 1937, the world’s human population numbered 2.3 billion, atmospheric carbon dioxide stood at 280 parts per million (ppm), and 66% of the earth was wilderness. By 1989, humanity had more than doubled, growing to 5.1 billion, carbon dioxide in the atmosphere had reached 353 ppm, and only 49% of the earth remained wilderness. Just 31 years later in 2020, wilderness has dwindled to a mere 35% of the earth, carbon dioxide in the atmosphere has shot up to 415 ppm, something not seen in more than 800,000 years (Lindsey 2009), and the world’s human population has ballooned to 7.8 billion (Attenborough 2020). The Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES 2019) reports that 1 million species are in imminent danger of extinction. The climate change and biodiversity crises and associated loss and decline of wetlands are accelerating at paces unimaginable to most of us even just a few years ago, not to mention the concomitant social upheavals and crises. As we continue with “business as usual”, no end is in sight for these accelerating downward trajectories in the condition of our planet and its atmosphere. Thus, new approaches are required if we wish to restore a stable climate, healthy biodiversity, and a viable, sustainable future for humans and other species, all of which depend on functioning wetlands and other ecosystems. Along with non-governmental organizations (such as the Global Alliance for the Rights of Nature, the Earth Law Center, and the Community Environmental Legal Defense Fund) and others, Indigenous peoples and local communities have been leading a growing Rights of Nature movement that provides an alternative paradigm, one that fundamentally realigns the human-Nature relationship and recognizes the legal personhood of Nature based on the

living beingness of Nature, and Nature’s inherent rights, including the right to exist. By fundamentally shifting our relationship with Nature and with wetlands – to one that is based on respect, reciprocity and gratitude, similar to the relationship that many Indigenous peoples and local communities have had with Nature for millennia, we may find a truly sustainable and restorative way to manage wetlands and live on this planet. Recent successes in recognizing the rights of Nature include the passage of the Te Awa Tupua Act (Whanganui River Claims Settlement Act) in 2017 (New Zealand Parliament 2017) by the New Zealand Parliament in agreement with the Maori iwi (iwi = tribes), in which the legal rights and living personhood of the Whanganui River, as well as the special relationship between the Maori iwi and the river, were recognized in law. Other examples include the 2008 recognition of the rights of Nature in the Ecuadorian Constitution (Morales 2013), the 2018 issuance of the Kawsak Sacha (Living Forest) Declaration (Kichwa Native People of Sarayaku 2018) by the Ecuadorian Sarayaku, and the 2010 issuance of the Universal Declaration of the Rights of Mother Earth (World People’s Conference on Climate Change and the Rights of Mother Earth 2010) at the World People’s Conference on Climate Change and the Rights of Mother Earth in Cochabamba, Bolivia, where the inherent rights of Mother Earth and status as a living being were declared. In 2020, the Earth Law Center and International Rivers issued a Universal Declaration of the Rights of Rivers (Earth Law Center and International Rivers 2020), which includes riparian wetlands. Although many may find the recognition of the legal personhood and rights of Nature, and possibly the living beingness of Nature, to be unfamiliar, this perspective has deep cultural, legal, religious, and philosophical roots around the world, including in western thinking (Nash 1989; Stone 2010; Davies et al. 2020). As a wetlands-specific response to the global crises and the rights of Nature movement, and through the SWS Ramsar Section, the SWS Climate Change and Wetlands Initiative and the SWS Rights of Wetlands Initiative, a group of scientists and attorneys have proposed a Universal Declaration of the Rights of Wetlands (Declaration) (Davies et al. 2020). Davies et al. (2020) include a timeline and world map as supplementary material to the Declaration article, as illustration of the global and historical depth and breadth of rights of Nature. However, this timeline and world map are far from comprehensive. The authors welcome and encourage the addition of further examples from others.

Wetland Science & Practice January 2022 9

The preamble and proposed Declaration are as follows: UNIVERSAL DECLARATION OF THE RIGHTS OF WETLANDS Acknowledging that wetlands are essential to the healthy functioning of Earth processes and provision of essential ecosystem services, including climate regulation at all scales, water supply and water purification, flood storage, drought mitigation and storm damage prevention; Acknowledging that wetlands have significance for the spiritual or sacred inspirations and belief systems of many people worldwide, but particularly for Indigenous peoples and local communities living in close relationship to wetlands, and that wetlands provide opportunities to learn from and about Nature, which supports scientific understanding and innovation, cultural expression and artistic creativity; Further acknowledging that humans and the natural world with all of its biodiversity depend upon the healthy functioning of wetlands and the benefits that they provide, and that wetlands play a significant role in global climate regulation; Alarmed that existing wetland conservation and management approaches have failed to stem the loss and degradation of wetlands of all types around the globe; Further alarmed that global climate destabilization and biodiversity losses are accelerating and that efforts to reverse these trends are failing; Acknowledging that peoples around the world of many cultures and faiths have recognized for millennia that Nature, or elements of Nature, are sentient living beings with inherent value and rights independent of their value to humans, and that Indigenous peoples, local communities and non-governmental organizations have been contributing to a global movement to recognize the rights of Nature; Aware that continued degradation and loss of wetlands threatens the very fabric of the planetary Web of Life upon which depend the livelihoods, wellbeing, community life and spirituality of many people, particularly Indigenous peoples and local communities who live in close relationship with wetlands; Guided by recent legal recognition of the inherent rights of Nature, including recognition of the entire Colombian Amazon as an “entity subject to rights” by the Colombian Supreme Court; recognition of the rights and legal and living personhood of the Whanganui River through the Te Awa Tupua Act (Whanganui River Claims Settlement Act) agreed upon by the Māori iwi and the New Zealand Parliament; and Ecuador’s first-in-the-world recognition of the rights of Nature in their Constitution; Convinced that recognizing the enduring rights and the legal and living personhood of all wetlands around the world will enable a paradigm shift in the human – Nature relationship towards greater understanding, reciprocity and respect leading to a more sustainable, harmonious and healthy global environment that supports the well-being of both human and non-human Nature; Further convinced that recognizing the rights and legal and living personhood of all wetlands and the paradigm shift that this represents will lead to increased capacity to manage wetlands in a manner that contributes to reversing the destabilization of the global climate and biodiversity loss; WE DECLARE that all wetlands are entities entitled to inherent and enduring rights, which derive from their existence as members of the Earth community and should possess legal standing in courts of law. These inherent rights include the following: 1. The right to exist. 2. The right to their ecologically determined location in the landscape. 3. The right to natural, connected, and sustainable hydrological regimes. 4. The right to ecologically sustainable climatic conditions. 5. The right to have naturally occurring biodiversity, free of introduced or invasive species that disrupt their ecological integrity. 6. The right to integrity of structure, function, evolutionary processes and the ability to fulfil natural ecological roles in the Earth’s processes. 7. The right to be free from pollution and degradation. 8. The right to regeneration and restoration.

10 Wetland Science & Practice January 2022

WETLAND CONSERVATION – WITHOUT COMMUNITY ENGAGEMENT AND PARTICIPATION IT DOES NOT EXIST As outlined above, are facing global biodiversity and climate change crises that include the widespread loss and degradation of wetlands but how best do we stem this loss and conserve and protect wetlands? Historically, conservation efforts have concentrated on protecting specific areas and endangered species. This has led to positive outcomes for nature within many protected areas but with only a fragment of the globe protected, 15% of land and 3% of the oceans (UNEP-WCMC 2020), this has not addressed the steep decline in biodiversity outside of these areas (WWF 2020). The original goal for protected areas was to keep nature safe from people resulting in people being kept out of an area. Although, more recently, this approach has changed and the requirements of people are more generally considered within protected area plans, protected areas can lead to considerable negative social and economic impacts on local people although evidence is limited (Dasgupta et al. 2017). At the same time, a focus on endangered species, whilst leading to success for some critically endangered species, has seen a catastrophic decline in overall species populations (WWF 2020). The problem with a protected area and endangered species conservation approach is that it can exclude the needs of people and draw attention away from wider landscape. This approach to conservation can result in a disconnect of conservation efforts from local and regional populations. This disconnect means a focus on key drivers of poverty and consumption, which directly impact on nature, may not occur and support for conservation from local communities does not happen as a result. To address this situation focus should be on integrating biodiversity conservation into every aspect of land and water management, whether agriculture, urban or industrial planning. Wetland conservation cannot exclude people as local communities must be integral to conservation actions ensuring their needs are met and their voices heard (Shrestha 2013). Without the active participation of local communities ultimately wetland conservation will fail. Wetland conservation cannot remain behind a fence protecting an area but integrated across the landscape with local communities involved in planning and management. To achieve community and stakeholder involvement you need to invest resources because talking to people and hearing their views requires both effort and time (Tschirhart et al. 2016). It is also important to use participatory techniques to ensure all voices are heard and respected, creating a space where scientific and non-scientific knowledge can be exchanged (Project COBRA Consortium 2014). Participatory techniques such as interviews, focus groups

or participatory video can be used to resolve conflicts and build a consensus of the most appropriate actions to benefit the community and conservation. It is useful to use a learning cycle approach to all stages of wetland conservation – planning, implementing, monitoring and reviewing because reflecting on what is working and what is not working helps improve community and conservation outcomes and build a consensus for the most robust solutions (Project COBRA Consortium 2014). A focus should always be on ensuring conservation actions are embedded into community actions, so communities are supported to protect their own natural resources. An approach, termed ‘community owned solutions’, developed working with Indigenous communities in South America (Mistry 2016), attempted to do this by testing, with the community, whether an intervention really was something a community needed, whether they would undertake it, control it, benefit from it, whether it was fair and whether the action was good for the environment. If a conservation intervention achieves these measures, then it will become embedded within community practices and it is more likely to be successful and continue beyond the lifetime of an initiative or funding. Increasing evidence demonstrates that when local communities actively manage their own natural resources there are positive outcomes for biodiversity (Corrigan et al. 2018). To achieve wetland conservation local communities must actively be engaged, participate in all aspects of conservation and ideally manage their own resources. RESPONDING TO THE STATEMENT OF WORLD AQUATIC SCIENTIFIC SOCIETIES ON THE NEED TO TAKE URGENT ACTION AGAINST HUMAN-CAUSED CLIMATE CHANGE On 14 September 2020, the Society of Wetland Scientists (SWS) joined 110 other aquatic science societies, representing more than 80,000 scientists across the world, to release a statement on climate change entitled “Statement of World Aquatic Scientific Societies on the Need to Take Urgent Action against Human-Caused Climate Change” (https:// climate.fisheries.org/world-climate-statement/ accessed 14 December 2020). The release of the Statement was coordinated by the American Fisheries Society. The Statement highlights the problems posed by climate change for aquatic ecosystems, and suggested actions to help turn things around. Impacts from climate change on wetlands are already occurring and many of these will be irreversible, and will continue if current socio-economic trajectories are not changed. Given that wetlands are in decline globally, the impact of climate change is expected to make a bad situation worse (Finlayson et al. 2018; Ramsar Convention 2018). Wetlands, particularly disturbed wetlands, are also an important contributor of emissions that Wetland Science & Practice January 2022 11

are driving climate change (Moomaw et al. 2018a). Despite an increase in the level of attention being directed towards adaptation measures to respond to the impact of climate change on wetlands, far more effort is needed to develop appropriate policy and management responses, including at an international level (Finlayson et al. 2017a). The Statement of World Aquatic Science Societies included recommendations for responses with four of those of relevance to wetlands described below, with the addition of specific suggestions for members of SWS. 1. Rapid action is necessary to curb the release of greenhouse gases and remove carbon dioxide from the atmosphere to help decrease the adverse consequences of climate change on wetlands, as outlined in Moomaw et al. (2018a). The biogeochemical processes that enable carbon emissions and the storage of carbon on wetlands are well known scientifically (Kang and Jang 2018), but policy outcomes for wetlands are lagging internationally and nationally and warrant further attention (Finlayson et al. 2017a). Communication and raising awareness about the importance of wetlands for carbon emissions and potential sequestration is needed from scientists to help inform wetland decision and policy makers about the important role of wetlands in mitigating climate change and to build public support for such policies. Individuals can help by converting their science into readily understandable messages and guidance for policy makers and the general public, as done by Moomaw et al. (2018b), including for managing ‘carbon in wetlands’ is needed to ensure the benefits of carbon storage in wetlands are realised and carbon emissions minimised. Such activities may need to be supported by incentives to reward good practice (possibly seen as a system of payment for carbon services) – we can help by converting our science into practical guidance for management and to support incentives to reward good practice. 2. Global/national targets are necessary to help protect and restore carbon dense ecosystems, such as peat, salt marshes, sea grasses, and other wetlands to sequester carbon, prevent greenhouse gas emissions, and reduce the impacts of climate change. Efforts to set targets and encourage better outcomes for wetland biodiversity have not been successful (Davidson et al. 2020). Biodiversity targets for wetlands are needed, but underlying data needs attention. Wetland scientists can assist by suppling accurate data for targets for individual wetland sites, and more generically for wet12 Wetland Science & Practice January 2022

land types and species, and encourage policy-makers to confirm the efficacy of the underlying data and data collection mechanisms to avoid setting unrealistic targets. This is well illustrated by the inadequate state of wetland inventory globally, as reported to the Ramsar Convention in 1999 (Finlayson et al. 1999) and again by Davidson et al. (2020). 3. Robust adaptation measures: identification and easing of other environmental stressors that act synergistically with climate change; and additional resources for data collection, mapping, and research to better understand potential impacts and to arm natural resources agencies with the tools to mitigate these impacts are essential. Adaptation to climate change impacts on wetlands is needed and dependent on effective vulnerability assessment – supply information on wetland dynamics, trends and trajectories, and likely responses to climate scenarios. Wetland managers need practical tools to respond and manage for the future – wetland scientists need to convert their science into guidance for managing complex systems, and provide training. 4. It is time to acknowledge the urgent need to act to address climate change. Delaying action to control greenhouse gas emissions is not an option if humankind wishes to conserve wetlands and the environmental safety of the world. Science information needs to be actively used to influence policy and decision-making – scientists and practitioners need to actively engage in policy dialogue and ensure that policy mechanisms for climate change are aware of the role and importance of wetlands. Scientists and practitioners need to more actively engage in policy dialogue and advocate and question the efficacy of existing policy and management mechanisms that have hitherto not achieved the sustainable future of wetlands globally. Members of the Society of Wetland Scientists are invited to consider these needs and to work with the recently established SWS Climate Change and Wetlands Initiative (Finlayson et al. 2020a). The Initiative builds on climate change statements signed by individuals at SWS meetings in 2017, 2018 and 2019 (Finlayson et al. 2017b, 2019, 2020b). The purpose of the Initiative is to develop sciencebased responses, share knowledge and do more as a group of wetland experts concerned about the future of wetlands, and about climate change in particular. In addition to the climate change statements, individuals now involved with the Initiative have contributed to a review of wetlands and

climate change (Moomaw et al 2018a) and a response to the Second Warning to Humanity from World Scientists (Finlayson et al 2018). Further activities are being considered, including a review of information on carbon stocks in different wetland types, and a collation of key issues from SWS special interest Sections about wetlands and climate change. CONCLUSION Based on the concepts presented in the combined presentations in the Ramsar Section’s symposium in the December 2020 SWS virtual conference, and from associated recent activities (as outlined in Finlayson et al. 2020a; Fennessy et al. 2021; Simpson et al. 2020) our vision for the SWS is as follows: “The Society of Wetland Scientists engages with wetland professionals and wider society (including reinvigorated Multilateral Environmental Agreements) to promote wetland research, education, and management in support of a sustainable future: develop new approaches; link with our communities; and get past the rhetoric to support effective actions, including those that reduce the impact of climate change on wetlands.” With this in mind, speakers in the symposium are keen to engage in further dialogue with others interested in wetlands to develop and refine new approaches and paradigms for better addressing efforts for achieving the wise use of wetlands. REFERENCES

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Ghermandi, A., J.C. Van Den Bergh, L.M. Brander, H.L. De Groot, and P.A. Nunes. 2010. Values of natural and human‐made wetlands: A meta‐analysis. Water Resources Research 46(12). https://doi. org/10.1029/2010WR009071 Griscom, B.W., J. Adams, P.W. Ellis, R.A. Houghton, G. Lomax, D.A. Miteva, ... and J. Fargione. 2017. Natural climate solutions. Proceedings of the National Academy of Sciences 114: 11645-11650. IPBES. 2019. Summary for policymakers of the global assessment report on biodiversity and ecosystem services of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services. IPBES Secretariat, Bonn, Germany. Kang, H. and I. Jang. 2018. Carbon flux from wetlands. In Finlayson, C.M., M. Everard, K. Irvine, R.J. McInnes, B.A. Middleton, A.A. van Dam and N.C. Davidson (eds) 2018. The Wetland Book I: Structure and Function, Management and Methods. Springer Publishers, Dordrecht. Pp. 277-284.

O’Connor, M. 1994. On the misadventures of capitalist nature. In M. O’Connor (Ed.) Is Capitalism Sustainable?: Political Economy and the Politics of Ecology. Guilford Press, New York, USA. pp. 125–151. Phillips, B.J., E.F. McQuarrie and W.G. Griffin. 2014. The face of the brand: How art directors understand visual brand identity. Journal of advertising, 43(4), 318-332. Project COBRA Consortium. 2014. How to find and share community owned solutions - A Handbook. https://cobracollective.org/practitionershandbook/ Accessed 25 January 2021. Ramsar Convention. 2018. Global Wetland Outlook; State of the World’s Wetlands and Their Ecosystem Services. Ramsar Convention Secretariat, Gland, Switzerland. Richardson, C.J. 1994. Ecological functions and human values in wetlands: a framework for assessing forestry impacts. Wetlands 14(1): 1-9.

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Kumar, R., S. Tol, R.L. McInnes, M. Everard, and A.A. Kulindwa. 2017. Wetlands for disaster risk reduction: Effective choices for resilient communities. Ramsar Policy Brief No. 1. Ramsar Convention Secretariat, Gland, Switzerland.

Sheehan, L., E.T. Sherwood, R.P. Moyer, K.R. Radabaugh and S. Simpson. 2019. Blue carbon: an additional driver for restoring and preserving ecological services of coastal wetlands in Tampa Bay (Florida, USA). Wetlands 39(6): 1317-1328.

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LATIN AMERICAN WETLANDS

Introduction to this Special-focus Issue

n October 2020, we published a special issue on Latin American wetlands – our first and only issue emphasizing wetland-related activities in a particular region of the world. That issue contained 19 articles and notes covering various research, conservation initiatives, and outreach activities being conducted in Central and South America. It was greeted with such success that we decided to initiate a second special issue to give researchers, managers, and others interested in Latin American wetlands a chance to bring attention to their work and improve the connections among individuals from this region. So in April 2021, we made an announcement about another issue and a call for papers. Draft manuscripts were due by October 2021. The contributions were reviewed and edited by the WSP editor

and returned to the corresponding author for final preparation. To start the new year off, we are pleased to publish 11 articles about ongoing activities in and concerns for wetlands from the Caribbean, Central America, and South America. A special thanks go out to the authors for taking time to prepare these articles. We also solicited images of wetlands for the accompanying Notes from the Field. For this we received numerous candidates and selected a good number of wetland scenes and wetland animal pictures to illustrate some of the diversity of the region’s wetlands and wildlife. Thanks to all who sent images for our consideration. We hope that readers enjoy this special contribution as well.

Ralph Tiner, WSP Editor and Tatiana Lobato de Magalhães, Special Issue Coordinator

Capybara, the world's largest rodents, are native to South America. By: (reisegraf – stock.adobe.com) Wetland Science & Practice January 2022 15

PATAGONIAN RUSHES

Juncus in Northern Patagonian Wetlands Adriel I. Jocou and Ricardo Gandullo1,2 INTRODUCTION The genus Juncus L. (Juncaceae Juss.) contains about 315 species, distributed abundantly in both the northern and southern hemispheres, but they are rarer in the tropics (Balslev 1996; Kirschner 2002; Romero Zarco 2010). Juncus is currently divided into two subgenera and 10 sections (Kirschner 2002): subgenus Juncus (six sections) and

Subgenus

subgenus Agathryon Raf. (four sections). Although various studies on Juncus have been carried out in Argentina and neighbouring countries (e.g., Barros 1945, 1953, 1969; Novara 2009, 2012; Zuloaga et al. 2019; Jocou and Brignone 2020), a thorough and current review of this genus is necessary. In Argentina, the genus Juncus is represented by ca. 27 species and 10 infraspecific taxa (Table 1).

Section

Number of species in the world

Distribution

Number and species in the Argentinean Flora

Forskalina Kuntze

Distributed in the whole Mediterranean region.

not represented

Juncotypus Dumort.

World-wide distribution. Mainly in Australia and Pacific North America.

Steirochloa Griseb.

Widespread, with species native to all temperate regions except for South Africa. Mainly in W and E North America, Central Asia and temperate South America.

7: Juncus capillaceus Lam.; J. cordobensis Barros; J. dichotomus Elliott; J. imbricatus Laharpe; J. occidentalis (Coville) Wiegand (incl. J. tenuis var. congestus); J. tenuis Willd.; J. venturianus Castillón

Tenageia Dumort.

W of Mediterranean region; some species very widespread.

2: Juncus bufonius L. [var. bufonius]; J. sorrentinii Parl. (= J. bufonius var. condensatus Cout.)

Caespitosi Cout.

Southern Africa, W North America.

not represented

Graminifolii Engelm.

W North America, southern Africa, Australasian region.

2: Juncus cyperoides Laharpe; J. marginatus Rostk.

Iridifolii Snogerup & Kirschner

W North America, Eastern Asia.

Not geographically circumscribed. It includes species from Europe, America, Africa and East Asia.

Agathryon Raf.

Juncus Juncus

Ozophyllum Dumort.

Stygiopsis [Grand. ex] Kuntze

E North America, South America, SW Europe, the Far East, North Africa, S Africa, Oceania.

In high mountains of North Hemisphere and subarctic regions, centred in the SinoHimalaya.

Departamento de Biología Aplicada. Facultad de Ciencias Agrarias. Universidad Nacional del Comahue. Cinco Saltos, Río Negro, Argentina. Correspondence author contact: adrieljocou@gmail.com 2 Herbario ARC (Agronomía Región Comahue). Facultad de Ciencias Agrarias. Universidad Nacional del Comahue. Cinco Saltos, Río Negro, Argentina. 1

16 Wetland Science & Practice January 2022

4: Juncus balticus Willd. [subsp. mexicanus (Willd. ex Roem. & Schult.) Kirschner; subsp. andicola (Hook.) Snogerup]; J. effusus L. [subsp. effusus]; J. procerus E.Mey.; J. uruguensis Griseb.

not represented 1: Juncus acutus L. [subsp. leopoldii (Parl.) Snogerup]

11: Juncus articulatus L. [subsp. articulatus]; J. burkartii Barros; J. densiflorus Kunth; J. diemii Barros; J. ernesti-barrosii Barros; J. llanquihuensis Barros; J. micranthus Schrad. ex E.Mey.; J. microcephalus Kunth; J. pallescens Lam. [var. pallescens; var. achalensis (Barros) Novara]; J. scheuchzerioides Gaudich.; J. stipulatus Nees & Meyen [var. stipulatus, var. chilensis(Gay) Kirschner] not represented

Table 1. Summary of subgenera and sections within Juncus, number of species in each section and their distributions worldwide, and the number and species per section in the Argentinean flora. Infraspecific rank taxa in brackets, synonyms in parenthesis.

The northern Patagonian wetlands flora contains 48 families, 137 species and 172 infraspecific taxa of vascular plants (Jocou and Gandullo 2020). In this flora, Juncaceae has the fourth most species and Juncus is the largest genus (Jocou and Gandullo 2020). This paper contributes to the correct identification of Juncus taxa which are crucial in floristic studies of wetlands in this region. Since Juncus is a taxonomically complicated genus and it has not been revised in Argentina, the aim of this work is to review the taxa that occur in northern Patagonian wetlands, while providing a dichotomous key, morphological and ecological information, and photographs of each taxon. STUDY AREA The study wetlands are located in the northern Argentinian Patagonia, between 38° 30’S and 39° 30’S and 66° 55’W and 69° 20’W at altitudes between 190 and 380 m a.s.l. (Figure 1); they are the same wetlands studied by Jocou and Gandullo (2020). The dominant climate is cold temperate, arid to semi-arid, with average annual temperatures of 14ºC, rainfall from 150–300 mm annually with a Mediterranean regime, and average annual evapotranspiration of 800 mm (Peri 2004; Gandullo and Gastiazoro 2009; Lozeco 2014; Jocou et al. 2018).

Figure 1. Study area, location of the surveyed wetlands. Diamond indicates the natural riparian lagoon Balsa Las Perlas, triangle indicates Maturana lagoon, circle indicates Ezequiel Ramos Mexía dam. Dashed line indicates study area of rivers, and irrigation and drainage canals. (Source: Satellite layer, from Google Earth ®)

The taxa of Juncus occur in six environments with marked and fluctuating water conditions along the year (for a detailed description see Jocou and Gandullo 2020): 1. Maturana lagoon (Figure 2A, B) – an artificial lagoon located in the city of Senillosa (province of Neuquén) a few meters from National Route 22 (39°00’41’’S and 68°26’20’’W). 2. Ezequiel Ramos Mexía dam (Figure 2C, D) - an artificial lake located on the boundaries of Río Negro and Neuquén provinces (39º30’S and 69º00’W). 3. Balsa Las Perlas lagoon (Figure 2E, F) – a small natural riparian lagoon, located at 60 m from the shore of the Limay River (38°58’47’’S and 68°7’23’’W). 4. Drainage canals (Figure 3A–C) – an extensive network of artificial canals (more than 500 km) that collect excess water for irrigation, infiltration and rain, as well as urban and industrial waste; they are located throughout the valleys. 5. Irrigation canals (Figure 3D–F) – an extensive network of artificial canals (more than 2000 km) that divert the watercourse of the rivers; they are used for the irrigation of fruit and vegetable production in the valleys of the Limay, Neuquén and Negro Rivers.

Figure 2. Environments inhabited by taxa of Juncus. A, B. Maturana lagoon. C, D. Ezequiel Ramos Mexía dam. E, F. Balsa Las Perlas lagoon. (Photographs: A.I. Jocou)

6. Rivers - Limay, Negro and Neuquén- (Figure 3G, H). The Negro River Basin is among the most important hydrographic systems in Argentina; it is the main source of water for fruit production in the lower valleys of the Neuquén and Limay Rivers, and the upper valley of the Negro River. The water level varies during the year depending on the use of hydroelectric dams, irrigation, and seasonal climatic conditions in the upper basin. MATERIALS AND METHODS Previous works on the genus Juncus were consulted (Barros 1945; 1953; 1969; Balslev 1996; Kirschner 2002; Novara 2009; Romero Zarco 2010; Novara 2012; Balslev and Duno de Stefano 2015; Balslev 2018; Zuloaga et al. 2019; Wetland Science & Practice January 2022 17

Jocou and Brignone 2020). We created a list of taxa of Juncus from our ongoing ecological and floristic studies in the wetlands of Patagonia in Argentina (e.g., Fernández et al. 2018; Jocou et al. 2018; Gandullo et al. 2019; Jocou and Gandullo 2020; Jocou and Brignone 2020). We reviewed the Argentinean specimens of Juncus kept at the ARC (herbarium acronyms following Thiers 2021) and confirmed their identity using classical methods in taxonomy. The morphological study of the specimens was performed under a stereo-microscope Leica EZ4 HD, and the photographs were taken using LAS EZ 3.4.0. RESULTS AND DISCUSSION For each name, the earliest published name or basonym (indicated as ≡) and its type specimens are provided. The heterotypic synonyms are indicated as = symbol. The exclamation mark after citing a type specimen indicates that the specimen was seen on a scanned image. The examined specimens cited are kept at ARC; those cited as Specimens of reference are from northern Patagonian wetlands, whereas Additional examined specimens include those from other areas from Patagonia in Argentina. The water characteristics of wetlands where the studied taxa of Juncus were found are shown in Table 2. The taxa from the northern Patagonian wetlands can be distinguished using the following dichotomous key. For precise identification, the specimens must have both vegetative

Figure 3. Environments inhabited by taxa of Juncus. A-C. Drainage canals, distinct wet conditions and dimensions. D-F. Irrigation canals, distinct dimensions: primary (F), secondary (E), and tertiary (D) canals. G, H. Shores of Limay River. (Photographs: A.I. Jocou)

(rhizomes, stem, leaves) and reproductive structures (flowers, fruits). We recommend taking detailed field photographs and preparing dry specimens to be kept in herbaria for further studies.

Taxon J. acutus subsp. leopoldii J. articulatus subsp. articulatus J. balticus subsp. andicola J. balticus subsp. mexicanus J. bufonius var. bufonius J. effusus subsp. effusus J. microcephalus J. pallescens var. pallescens J. tenuis

pH 7.7–8.4 7.2–9 5.8–8.6 5.8–8.6 7.2–8 6–7.7 5.8–9 6–9 7.2–8

Water characteristics EC (µS/cm) DO (mg/L) 230–250 5.9–7.9 70–280 8–10 187–1620 0.4–12 187–1620 0.4–12 70–180 9–10 79–254 3.8–9.2 70–1620 0.4–12 70–280 3.8–10 70–80 9–10

Table 2. Water characteristics of wetlands inhabited by taxa of Juncus. EC: electrical conductivity, DO: dissolved oxygen.

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KEY OF THE TAXA OF JUNCUS PRESENT IN NORTHERN PATAGONIAN WETLANDS 1. Inflorescence racemose, flowers without a pair of bracteoles below; flowers usually in 2 heads or clusters, rarely borne singly or in loose groups 1’. Inflorescence cymose, flowers with a pair of bracteoles below; flowers usually borne 5 singly or in loose groups 2. Leaves stem-like, pungent, not septate; lower bract apparently forming a prolongation J. acutus subsp. leopoldii of stem 2’. Leaves not stem-like, perfectly septate; lower bract not appearing as a stem 3 prolongation J. articulatus subsp. articulatus 3’. Capsule shorter than the perianth 4 4. Tepals rigid and castaneous to dark castaneous; flower head peduncles usually 0.45 J. pallescens subsp. pallescens mm thick or thicker. 3. Capsule exceeding the perianth

4’. Tepals soft and stramineous-castaneous to castaneous; flower head peduncles usually J. microcephalus 0.4 mm thick or thinner. 5. Annuals J. bufonius var. bufonius 5’. Perennials 6 6. Inflorescence obviously terminal, lower bract flat, not appearing as a prolongation of J. tenuis the stem; basal leaves with a blade, and flat (grass-like) 6’. Inflorescence seemingly lateral (pseudo-lateral); lower inflorescence bract terete, 7 erect, apparently forming a prolongation of the stem; basal leaves usually bladeless, if bladed then terete or ± compressed and twisted, but never completely flat 7. Densely caespitose plants; rhizome very short-noded; tepals 2–3 mm long, acute to J. effusus subsp. effusus acuminate, green to light brown 7’. Not caespitose plants; rhizome sparsely branching with mostly 5–10 mm long nodes. 8 Tepals 3–5 mm long, obtuse to apiculate, light to dark brown 8. Stem usually 3–10 mm diam., not compressed or twisted; cataphylls with an awn-like J. balticus subsp. andicola blade, but never with well-developed blade 8’. Stem 1–3 mm diam., usually compressed and twisted; 1–2 uppermost cataphylls with J. balticus subsp. mexicanus a well-developed blade

Wetland Science & Practice January 2022 19

PROFILES OF THE TAXA 1. Juncus acutus L. subsp. leopoldii (Parl.) Snogerup, Bot. Not. 131: 187. 1978. ≡Juncus leopoldii Parl., Giorn. Bot. Ital. 2: 324-325. 1846. TYPE (lectotype designated by Snogerup, Willdenowia 23: 37. 1993): South Africa, Sommerset, Stellenbosch, Ecklon & Zeyer 4308 (FI! barcode 000909; isolectotypes S! barcode S-G-008001, W! barcode 0015534). Brief description. It is a densely caespitose plant with basal, very pungent leaves. Inner tepals mucronate, with two auricles. Stamens 6, anthers 2.5–4 times as long as filaments. Mature capsules brown to dark brown, rounded and exceeding the perianth. (Figure 4A–D). Ecology and occurrence. Worldwide this subspecies inhabits coastal sandy, salt marshes and inland moist saline or marshy habitats, and alkaline seeps, sometimes growing in places with a carbonate crust and sulphurous salts and in areas with groundwater; from sea level to 1500 m (Snogerup 1993; Balslev 1996; Brooks and Clemants 2000; Kirschner 2002; Romero Zarco 2010; Balslev and Duno de Stefano 2015). Novara (2012) considered this taxon as a halophytic indicator of very saline soils. In the northern Patagonian wetlands, this taxon grows on the shores of the Maturana lagoon at ca. 290 m a.s.l. The water is classified as alkaline, slightly saline and moderately to well-oxygenated (Table 2). The soils where it grows are sandy to sandy-clay. Specimen of reference. Prov. Neuquén: Senillosa, Laguna Maturana, 20-XII-2018, Jocou, Minué & Gandullo 2242 (ARC). 2. Juncus articulatus L., Sp. Pl. 1: 327. 1753. subsp. articulatus. TYPE (lectotype designated by Ferrer-Gallego, Taxon 68: 145. 2019): Herb. A. van Royen, (L! barcode 0221767). Brief description. Densely to loosely densely caespitose plants with perfectly septate leaves. Tepals maroon or straw-coloured, completely brown or with a green central band, ovate to lanceolate. Stamens 6, anthers usually equalling filaments. Mature capsules castaneous to dark brown, trigonous to ovoid, and exceeding the perianth. (Figure 4E–I). Ecology and occurrence. Worldwide, this subspecies inhabits various moist and wet natural or man-made places, margins of watercourses, usually on acid soils, often behaving like a calcicole, from sea level to 3700 m (Grime et al. 1988; Brooks and Clemants 2000; Guofang and Clemants 2000; Kirschner 2002; Romero Zarco 2010; Ferrer-Gallego 20 Wetland Science & Practice January 2022

Figure 4. Juncus acutus subsp. leopoldii: A. Habit, Maturana lagoon. B. Close-up of inflorescences, in vivo. C. Fruits, dry specimen. D. Closeup of perianth with capsule, dry specimen. Juncus articulatus subsp. articulatus: E. Habit, irrigation canal. F–G. Variation of inflorescence, in vivo. H. Close-up of inflorescence, dry specimen. I. Close-up of a head, dry specimen. (Photographs: A.I. Jocou)

2019; Jocou and Brignone 2020). In the northern Patagonian wetlands, this taxon grows on the shores of Neuquén and Limay Rivers, and in irrigation canals, from 250–290 m a.l.s. The water is near-neutral to alkaline, sparingly to moderate saline, and oxygen-saturated (Table 2). The soils where it grows are mainly sandy. Specimens of reference. Prov. Neuquén: Centenario, Balneario municipal, 29-XII-2016, Jocou & Minué 2313 (ARC); Neuquén, Balsa Las Perlas, 17-XI-2018, Jocou, Minué & Gandullo 2117 (ARC). Prov. Río Negro: Allen, 12-XII-2018, Jocou et al. 2314 (ARC); Cinco Saltos, 3-II2020, Jocou 2318 (ARC); Cipolletti, 12-XII-2018, Jocou et al. 2315 (ARC); General Fernández Oro, 9-I-2020, Jocou

2316 (ARC); General Fernández Oro, 9-I-2020, Jocou 2317 (ARC). 3. Juncus balticus Willd., Mag. Neuesten Entdeck. Gesammten Naturk. Ges. Naturf. Freunde Berlin 3: 298. 1809. Worldwide this species has seven geographically and morphologically recognized subspecies (Kirschner 2002). However, the taxonomy of J. balticus in South America remains unexplored (Kirschner 2002). In the northern Patagonian wetlands, two subspecies are recognized up to date. 3a. subsp. andicola (Hook.) Snogerup, Preslia 74: 258. 2002. ≡Juncus andicola Hook., Hooker’s Icon. Pl., ser. 2, 8: t. 714. 1848. TYPE (holotype): Ecuador, Andes of Quito, W. Jameson 51 (K! barcode 000574359, isotypes BM, not seen; G! barcode 00098687, 00098686). Brief description. Plants up to 110 cm tall, usually robust, not caespitose, forming large and often dense stands. Stems 3–10 mm thick, terete, not compressed (rarely twisted). Cataphylls bladeless. Inflorescence much-branched, bracteoles light to reddish-brown. Tepals castaneous to dark castaneous, with central band green to light brown. Stamens 6, anthers 1.5–2 times as long as filaments. Capsule ellipsoidovoid, trigonous, light brown, shorter than or equalling perianth, acuminate. (Figure 5A–D). Ecology and occurrence. This subspecies inhabits wet pastures in the mountains, margins of lakes, rivers, lagoons, ponds, and forest, gravelly banks of streams, often in cattle fields and other places under human influence; up to 4000 m a.s.l. (Balslev 1996; Kirschner 2002; Balslev and Duno de Stefano 2015). In the northern Patagonian wetlands, this taxon grows on drainage canals and shores of Maturana and Balsa Las Perlas lagoons, and rivers. The water is nearneutral to alkaline, slightly to highly saline and hypoxic to oxygen-saturated (Table 2). The soils where this taxon grows are mainly sandy, occasionally sandy-clay to clay. Specimens of reference. Neuquén, Senillosa, Laguna Maturana, 20-XII-2018, Jocou, Gandullo & Minué 2243 (ARC); Neuquén, Laguna Río Limay, Balsa Las Perlas, 17-XI-2018, Jocou, Minué & Gandullo 2115 (ARC); Río Limay, 17-XI2018, Jocou, Minué & Gandullo 2118 (ARC). Additional examined specimens. Prov. Neuquén: Camino El Huecú – El Cholar, s.d., Pérez s.n. (ARC); ANP Domuyo, Vega Los Menucos camino C° Las Papas, 01-III-2005, Gandullo GR563 (ARC). Prov. Río Negro: s.d., “Mallín 2”, 19I-1983, Ojeda s.n. (ARC).

Figure 5. Juncus balticus subsp. andicola: A. Habit, Maturana lagoon. B. Close-up of inflorescence, Limay River. C. Close-up of flowers. D. Close-up of perianth with capsule. Juncus balticus subsp. mexicanus: E. Habit, drainage canal. F. Herbarium sheet, arrows indicate the uppermost cataphyll with well-developed blade. G. Close-up of uppermost cataphyll. (Photographs: A.I. Jocou)

3b. subsp. mexicanus (Willd. ex Roem. & Schult.) Snogerup, Preslia 74: 257. 2002. ≡Juncus mexicanus Willd. ex Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 7(1): 178. 1829. TYPE (holotype): Mexico, Humboldt & Bonpland s.n. (B! barcode B -W 06843 -01 0, isotypes P barcode 00738780, not seen). Brief description. Plants up to 70 cm tall, not caespitose, forming loose to dense stands. Uppermost 1–2 cataphylls with well-developed blades. Stems usually up to 3 mm thick. Inflorescence diffuse or with several loose clusters on short, usually flexuose peduncles, bracteoles pale stramineous, sometimes pale castaneous-brown at the base. Tepals are not much different from those found in subsp. andicola. Stamens 6, anthers much longer than filaments. Capsule Wetland Science & Practice January 2022 21

more or less equalling perianth, narrowly oblong-ovoid, subtrigonous, abruptly contracted, castaneous-brow to brown. (Figure 5E–G). Ecology and occurrence. This taxon inhabits open soils and sunny places, wet meadows, swamps, streams and ponds, often on sandy and more or less saline soils; it can be weedy and found on road banks, grazing areas, cropped areas, and even in crevices; from sea level to 4000 m a.s.l. (Kirschner 2002; Balslev and Duno de Stefano 2015). In the northern Patagonian wetlands, this taxon grows on the shores of the natural riparian lagoon Balsa Las Perlas, and along drainage canals. The water is near-neutral to alkaline, slightly to highly saline and hypoxic to oxygen-saturated (Table 2). The soils where this taxon grows are mainly sandy, occasionally sandy-clay to clay. After J. tenuis, this subspecies is the second most common taxon on drier soils in the studied wetlands. Specimen of reference. Prov. Neuquén: Neuquén, Laguna Río Limay, Balsa Las Perlas, 17-XI-2018, Jocou, Minué & Gandullo 2119 (ARC).

rary wet habitats with suppressed competition, behaviour as weedy species; from 0–2900 m a.s.l. (Balslev 1996; Brooks and Clemants 2000; Guofang and Clemants 2000; Kirschner 2002; Balslev and Duno de Stefano 2015). In the northern Patagonian wetlands, this taxon inhabits the shores of Limay and Negro Rivers. The water is near-neutral to alkaline, sparingly saline and saturated with oxygen (Table 2). The soils where this taxon grows are mainly sandy. Specimens of reference. Prov. Neuquén: Neuquén, Balsa Las Perlas, 17-XI-2018, Jocou, Minué & Gandullo 2116 (ARC). Prov. Río Negro: General Fernández Oro, 12-XII2018, Jocou & Gandullo 2203 (ARC). Additional examined specimens. Prov. Neuquén, Caviahue, 10-III-1980, Conticello 156 (ARC). Prov. Río Negro: s.d., 19-I-1983, Ojeda 73 (ARC). 5. Juncus effusus L., Sp. Pl. 1: 326. 1753. subsp. effusus. TYPE (lectotype designated by Lye, in Edwards, Sebsebe and Hedberg, Fl. Ethiop. Eritr. 6: 387. 1997): Herb. Linn. (LINN! barcode 449.6).

4. Juncus bufonius L., Sp. Pl. 1: 328. 1753. var. bufonius. TYPE (lectotype designated by Cope & Stace, Watsonia 12: 121. 1978): Herb. A. van Royen, (L! barcode 0052756). Brief description. Small annual caespitose plants, usually up to 25 cm tall. Inflorescence constitutes most of the plant height, lax, open, bracteoles hyaline, scarious. Tepals broadly lanceolate with greenish central band and broad membranous margins. Stamen usually 6, anthers usually as long as filaments. Capsule narrowly ovoid to subellipsoid, shorter than perianth, acute to subacute. (Figure 6A–D). Taxonomic remarks. Juncus bufonius is part of a complex of very similar and closely related taxa (Balslev 1996; Brooks and Clemants 2000; Kirschner 2002; Romero Zarco 2010). It is an extremely variable species in general habit as a result of a variety of habitats (Kirschner 2002) which led to the publication of numerous names of infraspecific level (see Barros 1953; Cope and Stace 1978; Balslev 1996; Brooks and Clemants 2000; Kirschner 2002). The specimens collected in northern Patagonian wetlands are treated here as var. bufonius, according to regional literature (see Zuloaga et al. 2019). Ecology and occurrence. This cosmopolitan taxon most frequently inhabits open muddy flats, denuded bottoms of ponds and periodic ponds, moist soils in meadows, lakeshores or stream banks, ditches, roadsides, drawdown areas, wet tracks, wet fields, swamps, and other tempo22 Wetland Science & Practice January 2022

Figure 6. Juncus bufonius var. bufonius: A. Habit, shores of Limay River. B. Herbarium sheet, size variation of specimens. C. Close-up of inflorescence, dry specimen. D. Close-up of perianth with capsule, dry specimen. Juncus effusus subsp. effusus: E. Inflorescence, Balsa Las Perlas lagoon. F. Habit, E.R. Mexía dam. G. Herbarium sheet. H. Closeup of inflorescence, dry specimen. I. Close-up of perianth with capsule, dry specimen. (Photographs: A.I. Jocou)

Brief description. Densely caespitose perennial plants, up to 100 cm tall. Inflorescence pseudolateral, many-flowered dense to moderately lax, bracteoles often unequal, ovate, acute. Flowers small, 2–3 mm long, tepals green to light brown, with scarious margin, lanceolate, acute to acuminate, thin and soft, central band greenish to stramineousbrown. Stamens usually 3, anthers equalling the filaments. Capsule trigonous-ovoid, obtuse, light brown, subequal to or slightly exceeding perianth. (Figure 6E–I). Ecology and occurrence. This taxon inhabits a wide range of wet sites from meadows and pastures, ditches, ponds, puddles in full sun, shores of bodies of water near roads, shores to mountain tracks and other places under the human influence (Balslev 1996; Kirschner 2002; Balslev and Duno de Stefano 2015). In the northern Patagonian wetlands, this taxon inhabits the shores of Ezequiel Ramos Mexía dam and the riparian lagoon Balsa Las Perlas. The water is nearneutral to alkaline, slightly to moderately saline and poorly oxygenated to saturated with oxygen water (Table 2).

treat it here as a heterotypic synonym of J. microcephalus, according to Balslev (1996) and Kirschner (2002). Some specimens collected in the northern Patagonian wetlands have agglomerated heads, similar to the type specimens of J. involucratus. Ecology and occurrence. This species inhabits wet highland sites and swampy places, along streams, ditches, canal banks, ponds, streams, creek bottoms (Kirschner 2002; Balslev and Duno de Stefano 2015). In the northern Patagonian wetlands, this taxon inhabits the shores of rivers and grows in drainage and irrigation canals. The water is nearneutral to alkaline, sparsely to highly saline and hypoxic to saturated with oxygen (Table 2). The soils where this taxon grows range from sandy to clay. Specimens of reference. Prov. Neuquén: Plottier, 19-I-2018, Jocou & Gandullo 2114 (ARC). Prov. Río Negro: General Fernández Oro, 09-I-2020, Jocou 2384 (ARC); Cipolletti, 14-IX-2016, Jocou, Fernández & Gandullo 2385 (ARC). 7. Juncus pallescens Lam., Encycl. 3: 268. 1789. var. pallescens.

Specimens of reference. Prov. Neuquén: Embalse E.R. Mexía, 10 km de Villa Chocón, 06-I-2019, Jocou & Minué 2240 (ARC); Neuquén, Laguna Río Limay, Balsa Las perlas, 17-XI-2018, Jocou, Minué & Gandullo 2120 (ARC).

TYPE (holotype): Argentina, buenos Aires, 1767, P. Commerson s.n. (P! barcode 00744272, isotype P?, not seen).

6. Juncus microcephalus Kunth., Nov. Gen. Sp. [H.B.K.] 1: 237. 1816

=Juncus dombeyanus J.Gay ex Laharpe, Mém. Soc. Hist. Nat. Paris 3: 132. 1827.

TYPE (holotype): Colombia, Quindio, Humboldt & Bonpland s.n. (P! barcode 00135241; isotypes A! (ex GH) barcode 00029703, B! barcode B-W 06860-01 0).

TYPE (holotype, sensu Balslev 1996): Peru, Dombey s.n. (P! barcode 00135244; isotypes P! barcodes 00135245, 00744275, 00744277, 00744278).

=Juncus involucratus Steud ex. Buchenau, Abh. Naturwiss. Vereins Bremen 4: 121. 1874.

Brief description. Caespitose, up to 60 cm tall. Inflorescence anthelate, of 3–25 semiglobose to globose, 8–20-flowered heads, each head 5–10 mm wide. Tepals rigid, castaneous to dark castaneous. Stamen usually 6, anthers shorter to equalling perianth. Capsule ellipsoid to obovoid, truncate to acute, trigonous, brown, glossy. (Figure 7E–H).

TYPE (lectotype designated by Balslev, Fl. Neotrop. Monogr. 68: 106. 1996): Peru, Tabina, Jul. 1854, Lechler 2078 (GOET! barcode 004188; isolectotypes K! barcode 000574353, MO! barcode 104757, S! barcode S-R-3087, O! barcode O-V-2014452, AAU, not seen). Brief description. Caespitose plants, up to 100 cm tall. Inflorescence anthelate, of 7–50 conical to globose, 10–35-flowered heads, each head 5–10 mm wide; heads often agglomerated. Tepals soft, stramineous to castaneous. Stamens 6, anthers shorter than filaments. Capsule ellipsoid to obovoid, obtuse, subequalling perianth, stramineousbrown to blackish; valves inflexed at the apex. (Figure 7A–D). Taxonomic remarks. This species is extremely variable and closely related to Juncus pallescens. Although Zuloaga et al. (2019) treated J. involucratus as separated species, we

Taxonomic remarks. This is an extremely variable species, with a number of forms sometimes recognized as separate species, varieties or subspecies (see Barros 1953; Balslev 1996; Kirschner 2002). Since the regional literature recognizes the var. achalensis (Barros 1953; Novara 2009; Novara 2012; Zuloaga et al. 2019) we treated the northern Patagonian specimens as var. pallescens, however other authors do not recognize the varieties (Balslev 1996; Kirschner 2002). The thickness of the inflorescence smallest branches was proposed as a primary feature for distinguishing J. pallescens (>0.45 mm) from J. microcephalus (<0.4 mm) (Balslev 1996; Kirschner 2002). However, we Wetland Science & Practice January 2022 23

8. Juncus tenuis Willd., Sp. Pl., ed. 4 [Willdenow] 2(1): 214. 1799. TYPE (lectotype designated by Balslev, Fl. Neotrop. Monogr. 68: 79. 1996): “America Boreali”, s.d. (B! barcode B -W 06888 -01 0, isolectotypes B! barcodes B -W 06888 -02 0; -03 0; -04 0; NY! fragment, barcode 00247725; HBG and AAU, not seen). Brief description. Perennial caespitose plants, usually 15–50 cm tall. Leaves basal, flat. Inflorescence usually lax, less often in denser clusters, of 1–5 primary branches, 5–40-flowered, flowers arranged in irregular groups, bracteoles broadly ovate, membranous. Tepals lanceolate, acuminate, central band green, paler and membranous towards the margins. Stamens 6, anthers shorter than filaments. Capsule sub-trigonous, ellipsoid, subacute to obtuse, tan or light brown, shorter than perianth. (Figure 7I–L).

Figure 7. Juncus microcephalus. A. Habit, irrigation canal. B. Herbarium sheet. C. Close-up of inflorescence, dry specimen. D. Close-up of heads. Juncus pallescens var. pallescens: E. Habit, irrigation canal. F. Herbarium sheet. G. Close-up of inflorescence, dry specimen. H. Close-up of heads. Juncus tenuis: I. Habit, the arrow indicates the plant, growing together with Mentha pulegium, Plantago lanceolata, and Phyla nodiflora var. minor in the shore of Limay River. J. Closeup of inflorescence, in vivo. K. Close-up of perianth with capsule, dry specimen. L. Close-up of inflorescence, dry specimen. (Photographs: A.I. Jocou)

noticed that the character is variable within each specimen, and seems to be not constant between species in the northern Patagonian material. Ecology and occurrence. This taxon has a wide geographical range and inhabits edges of lakes, bogs, disturbed wet sites at a variety of altitudes (Kirschner 2002). In the northern Patagonian wetlands, this taxon inhabits shores of rivers, the natural riparian lagoon Balsa Las Perlas and it grows in irrigation canals. The water is near-neutral to alkaline, sparsely to slightly saline and poor to saturated with oxygen (Table 2). The soils where this taxon grows vary from sandy to clay. Specimens of reference. Prov. Neuquén: Neuquén, Laguna Río Limay, Balsa Las Perlas, 17-XI-2018, Jocou, Minué & Gandullo 2122 (ARC). Prov. Río Negro: Mainqué, 28-XI2018, Jocou 2201 (ARC); General Fernández Oro, 01-XII2018, Jocou 2202 (ARC). 24 Wetland Science & Practice January 2022

Taxonomic remarks. Although Jocou and Gandullo (2020) included var. congestus Engelm. in their floristic study, the examined specimens actually correspond to J. tenuis. In addition, the var. congestus is recognized today as a synonym of J. occidentalis (Coville) Wiegand (Kirschner 2002). Ecology and occurrence. Juncus tenuis inhabits exposed or shaded areas, often along roads and disturbed sites in sandy to clay soils, under moist or drier conditions; wet grasslands, forests, on nitrified, and more or less compacted substrates; it is considered a weed whose distribution is influenced by human activity; from 1000–3000 m a.s.l. (Kirschner 2002; Romero Zarco 2010. Balslev and Duno de Stefano 2015). Balslev and Duno de Stefano (2015) stated that the dispersal of this species is facilitated by the ability of the seminal coating to swell and become sticky, jelly-like and sticking easily to grazing animals. In the northern Patagonian wetlands, this taxon inhabits the shores of Limay River, at an altitude much lower than reported by Balslev and Duno de Stefano (2015) for Mexico. The water is nearneutral to alkaline, sparingly saline and saturated with oxygen (Table 2). The soils where this taxon grows are mainly sandy. In the studied wetlands, it is the taxon that is most adapted to drier conditions. Specimen of reference. Prov. Neuquén: Senillosa, 20-XII2018, Jocou, Minué & Gandullo 2241 (ARC). EXCLUDED TAXA Juncus stipulatus Nees & Meyen, Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 19 (Suppl. 1): 126. 1843. TYPE (holotype): Chile, Valparaiso, Feb. 1831, Meyen s.n. (PR, not seen).

Although Jocou and Gandullo (2020) included this taxon in their floristic analysis, the examined specimens identified as Juncus stipulatus (Jocou, Minué & Gandullo 2117 ARC) correspond to depauperate and young plants of J. articulatus subsp. articulatus. Due to this, we exclude Juncus stipulatus from the flora of the northern Patagonian wetlands.

Gandullo, R., C. Fernández, and A.I. Jocou. 2019. Sintaxonomía de las comunidades de plantas vasculares del sistema de drenaje del Alto Valle de Río Negro, Patagonia, Argentina. Boletín de la Sociedad Argentina de Botánica 54: 567–587. Doi: 10.31055/1851.2372.v54.n4.24826

CONCLUSION Eight species and nine infraspecific taxa of Juncus occur in the northern Patagonian wetlands, inhabiting different water and soil conditions. Juncus stipulatus was previously recorded based on a misidentification and is excluded here for the northern Patagonian wetlands. A comprehensive revision of the species of Juncus in Argentina is needed for the clarification of both taxonomic and nomenclatural aspects within the genus.

Guofang, W. and S.E. Clemants. 2000. Juncaceae. In: Z.Y. Wu, and P.H. Raven (eds.). Flora of China 24. Science Press, Beijing, and Missouri Botanical Garden Press, St. Louis. pp. 44–69.

ACKNOWLEDGMENTS We are grateful to M. Appelhans (GOET herbarium) for providing the scan of specimen in his care and to H. Balslev (Aarhus University) for comments about Juncus and detailed review of the first version of this manuscript. REFERENCES

Balslev, H. 1996. Juncaceae. Flora Neotropica 68: 1–167. http://www. jstor.org/stable/4393863 Balslev, H. 2018. Two new species of Juncus (Juncaceae) from South America. Phytotaxa 375: 97–102. Doi: 10.11646/phytotaxa.376.2.3 Balslev, H. and R. Duno de Stefano. 2015. La familia Juncaceae en México. Acta Botanica Mexicana 111: 61–164. Doi: 10.21829/ abm111.2015.182 Barros, M. 1945. Juncáceas argentinas. Holmbergia 4: 101–112. Barros, M. 1953. Las Juncáceas de la Argentina, Chile y Uruguay. Darwiniana 10: 279–460. Barros, M. 1969. Juncaceae. In: N.M. Correa (dir.). Flora Patagónica, Parte II: Typhaceae a Orchidaceae (excepto Gramineae). INTA, Buenos Aires. pp. 109–137. Brooks, R.E. and S.E. Clemants. 2000. Juncus. In: Flora of North America Editorial Committee (eds.). Flora of North America 22. Oxford University Press, New York. pp. 211–255. Cope, T.A. and C.A. Stace. 1978. The Juncus bufonius L. aggregate in western Europe. Watsonia 12: 113–128. Fernández, C.J., A.I. Jocou, and R. Gandullo. 2018. Vegetación acuática bioindicadora de eutrofización del Alto Valle de Río Negro (Argentina). Ernstia 28: 45–93.

Grime J.P., J.G. Hodgson, and R. Hunt. 1988. The Autecological Accounts. In: J.P. Grime, J.G. Hodgson, and R. Hunt (eds.). Comparative Plant Ecology. Springer, Dordrecht. pp. 53–615. Doi:10.1007/978-94017-1094-7_5

Jocou, A.I. and N.F. Brignone. 2020. First record of Juncus articulatus subsp. articulatus (Juncaceae) for the Southern Cone flora. Boletín de la Sociedad Argentina de Botánica 55: 631–640. Doi: 10.31055/1851.2372. v55.n4.29938 Jocou, A.I. and R. Gandullo. 2020. Diversidad de plantas vasculares de los humedales de la Norpatagonia (Argentina). Revista del Museo Argentino de Ciencias Naturales 22: 131–154. Doi: 10.22179/REVMACN.22.688 Jocou, A.I., C. Fernández, and R. Gandullo. 2018. Macrófitas acuáticas vasculares del Sistema de drenaje del Alto Valle de Río Negro, Patagonia (Argentina). Revista del Museo de La Plata 3: 296–308. Doi: 10.24215/25456377e060 Kirschner, J. 2002. Juncaceae 2: Juncus subg. Juncus, Species Plantarum: Flora of the World 7. Australian Biological Resources Study (ABRS), Canberra, Australia. Lozeco, C.V. 2014. Desarrollo de un esquema de gestión integrada para los colectores de drenaje de la ciudad de Cipolletti (Río Negro, Argentina). Tesis de Mestría. Universidad Nacional del Litoral. Doi: 10.29104/ phi-aqualac/2015-v7-1-04 Novara, L.J. 2009. Juncaceae. In: R. Kiesling (dir.). Flora de San Juan, Vol. IV: Monocotiledóneas. Zeta Editores, Mendoza. pp. 377–391. Novara, L.J. 2012. Flora del Valle de Lerma: Juncaceae. Aportes Botánicos de Salta – Ser. Flora, Vol. 1 (edición digital). Herbario MCNS, Salta. Peri, G. 2004. La agricultura irrigada en Río Negro y su contribución al desarrollo regional. World Bank, Buenos Aires. Romero Zarco, C. 2010. Juncus. In: S. Talavera, M.J. Gallego, C. Romero Zarco, and A. Herrero (eds.). Flora Iberica: Plantas vasculares de la Península Ibérica e Islas Baleares 17. CSIC, Madrid. pp. 123–187. Snogerup, S. 1993. A revision of Juncus subgen. Juncus (Juncaceae). Willdenowia 23: 23–73. Thiers, B. 2021 [continuously updated]. Index Herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. http://sweetgum.nybg.org/science/ih Zuloaga, F.O., J.M. Belgrano, and C.A. Zanotti. 2019. Actualización del Catálogo de las Plantas Vasculares del Cono Sur. Darwiniana, nueva serie 7: 208–278. Doi: 10.14522/darwiniana.2019.72.861

Ferrer-Gallego, P.P. 2019. Typification of four Linnaean names in the genus Juncus (Juncaceae). Taxon 68: 142–151. Doi: 10.1002/tax.12013 Gandullo, R. and J. Gastiazoro. 2009. Suaedetum neuquenensis nueva asociación de ambientes salinos. Multequina 18: 31–36.

Wetland Science & Practice January 2022 25

TIDAL WETLANDS

From Marsh to Swamps: Vegetation Gradient Linked to Estuarine Hydrology Hugo López Rosas1 and Patricia Moreno-Casasola2 INTRODUCTION The distribution of ecosystems in coastal landscapes responds to environmental gradients generated by the continent-ocean interaction (Silva et al. 2017). The wetlands of the coastal zone are often influenced by the tide and receive input of saltwater from the ocean. However, since they are located in the lowest part of the basin, they continuously receive freshwater from the upper parts of the basin, either superficially or subsurface through the water table as well as from rainwater (Day et al. 2019). This interaction generates a mosaic with different types of coastal ecosystems, connected to each other by hydrology, and sharing species (Sheaves 2009). The excess of water and salinity are stressors for plant establishment (Perata et al. 2011; Rasool et al. 2013). In wetlands, hydrophytes have developed adaptations that allow them to tolerate or avoid these stressful conditions (Blom 1999; Howard and Mendelssohn 1999). This is especially true for plants growing in tidal saline wetlands (Tiner 2013). It is expected that, in a salinity and flood gradient, the greatest diversity or productivity will be present in the areas with less saline influence and less flooding. In this study we sampled the vegetation, interstitial water salinity and water level in a coastal wetland system where there is a clear zonation between herbaceous and arboreal vegetation and, within the latter, different types of plant formations. Our objective was to determine the presence of a combined water level and salinity gradient in relation to the different observed plant formations. STUDY SITE The wetland system where the present study is located is in the south and southeast of Laguna La Mancha, in the central coastal region of the State of Veracruz, in Mexico. (Figure 1). The climate in this region is warm sub-humid with a summer rainy season and a Precipitation/Temperature quotient greater than 55.3 (type AW2; García 1970). Mean annual precipitation varies between 1200 and 1650 mm. The mean annual temperature is 25.5°C with a minimum of 17°C in January and a maximum of 27.3°C in June Academia de Desarrollo Regional Sustentable, El Colegio de Veracruz, Carrillo Puerto 26, Zona Centro, Xalapa, 91000, Veracruz, México. Correspondence author contact: hugo.loper@gmail.com 2 Red de Ecología Funcional, Instituto de Ecología, A.C., Carretera Antigua a Coatepec No. 351, El Haya C. P. 91073, Xalapa, Veracruz, México, patricia. moreno@inecol.mx 1

26 Wetland Science & Practice January 2022

(Campos et al. 2011). There are three climatic seasons in the region: (a) the dry season (March to June) characterized by a mean monthly rainfall of 6 mm; (b) the rainy season (July to October) with a mean monthly rainfall from 200 to 400 mm; and (c) the nortes season (November to February) characterized by lower temperatures, strong northern winds (7 m/s on average at 10 m elevation), and a mean monthly rainfall of 24 mm (Castillo and Carabias 1982). The “La Mancha” lagoon has permanent and intermittent inputs of fresh water through streams, and also has input of salt water from the sea through intermittent communication with the Gulf of Mexico. The cold fronts of winter (nortes season) generate strong gusts of wind that lift the sand from the beach and the dunes, and deposit it at the mouth of the lagoon, forming a sandy bar that prevents communication with the Gulf of Mexico. During this season, the lagoon continues to receive freshwater through streams or by subsurface (Yetter 2004), which causes the water level of the entire system to rise (Moreno-Casasola et al. 2010). When the water level exceeds the level of the sand bar, it breaks through, so the lagoon once again communicates with the Gulf of Mexico, and the water level of the wetland system is offset by the sea level. The study site is within Ramsar Site 1336 “La Mancha y El Llano” and is partially within a private conservation area belonging to “Residencial Ecológico Diada La Mancha.” As shown in the map in Figure 1, The largest area of wetlands in the study site is covered by mangroves, but being located at the southern end of the lagoon, the distance to the mouth of the lagoon is two or more kilometers, and it is close to the mouth of the river called Caño Gallegos, which permanently carries water to the lagoon. Due to this location, the study site has a marked hydrological gradient, which is reflected in the presence of other types of vegetation, such as the flooded palm grove, the flooded freshwater forest and the freshwater herbaceous wetland. METHODS Sampling Design and Data Collection We distributed 19 permanent monitoring units (MUs) in four types of wetlands in the system. We placed three MUs in the freshwater swamp (FWS, Figure 2), three in the palm forest swamp (PFS, Figure 3), and nine in the flooded grassland (FG, Figure 4). For this study we did not place MUs in the mangrove, but we used previous information gathered by our work team (Moreno-Casasola et al. 2009; Utrera López and Moreno-Casasola 2008) to select another four MUs from this wetland. The mangrove physiognomy at the site is shown in Figure 5. The geographical location and the characteristics of the vegetation type and main species of each MU are presented in Table 1. Figure 6 shows

Figure 1. Map showing the different types of vegetation in the study site.

Wetland Science & Practice January 2022 27

Monitoring Unit (MU)

Vegetation type

Geographic location Latitude

Dominant species

Accompanying species

Longitude

FWS_01 FWS_02

Freshwater swamp Freshwater swamp

19.568587° 19.566294°

-96.379556° Annona glabra -96.379593° Sabal mexicana, Conocarpus erectus

FWS_03 PFS_01

Freshwater swamp Palm forest swamp

19.563870° 19.566827°

-96.379356° Annona glabra -96.380280° Sabal mexicana

PFS_02

Palm forest swamp

19.567565°

-96.380106° Sabal mexicana, Conocarpus erectus

PFS_03

Palm forest swamp

19.569334°

-96.380058° Sabal mexicana

FG_01

Flooded grassland

19.555920°

FG_02

Flooded grassland

19.555925°

FG_03

Flooded grassland

19.555929°

FG_04

Flooded grassland

19.554973°

FG_05

Flooded grassland

19.554977°

FG_06

Flooded grassland

19.554990°

FG_07

Flooded grassland

19.554025°

FG_08

Flooded grassland

19.554038°

FG_09

Flooded grassland

19.554042°

MS_01

Mangrove swamp

19.558693°

MS_02

Mangrove swamp

19.557859°

MS_03 MS_04

Mangrove swamp Mangrove swamp

19.568718° 19.564902°

-96.378484° Mimosa pigra, Echinochloa pyramidalis -96.378770° Echinochloa pyramidalis -96.379065° Echinochloa pyramidalis -96.378509° Echinochloa pyramidalis -96.378786° Mimosa pigra, Eleocharis mutata -96.379071° Mimosa pigra, Eleocharis mutata -96.378515° Echinochloa pyramidalis, Eleocharis interstincta -96.378810° Echinochloa pyramidalis -96.379105° Echinochloa pyramidalis -96.379735° Rhizophora mangle, Laguncularia racemosa -96.379530° Rhizophora mangle, Laguncularia racemosa -96.389261° Rhizophora mangle -96.388875° Avicennia germinans

None Laguncularia racemosa, Ficus cotinifolia Sabal mexicana Ginoria nudiflora, Pithecellobium dulce Avicennia germinans, Laguncularia racemosa Ginoria nudiflora, Avicennia germinans Ipomoea tiliacea, Pontederia sagittata Mimosa pigra Eleocharis mutata, Annona glabra Mimosa pigra Fuirena simplex, Echinochloa pyramidalis Eclipta prostrata, Leersia hexandra Leersia hexandra, Hymenocallis littoralis Mimosa pigra Annona glabra, Dalbergia brownei Avicennia germinans Avicennia germinans Laguncularia racemosa Laguncularia racemosa, Rhizophora mangle

Table 1. Geographical location, characteristics of the vegetation type, and main species of the monitoring units in this study.

28 Wetland Science & Practice January 2022

Figure 2. Freshwater swamp with a mix of species that include Annona glabra, Sabal mexicana, Ficus cotinifolia, and the terrestrial Pithecellobium dulce. (Photo by Hugo López Rosas)

Figure 3. Palm forest swamp dominated by Sabal mexicana. (Photo by Hugo López Rosas)

Figure 4. Flooded grassland dominated by the African invader grass Echinochloa pyramidalis. (Photo by Hugo López Rosas)

Figure 5. Mangrove swamp with Rhizophora mangle. (Photo by Hugo López Rosas)

the location of these MUs in the system. The minimum distance between contiguous MUs was 30 m in herbaceous vegetation and 80 m in arboreal vegetation The MUs were 10 x 10 m (100 m2) squares, with two 4 x 4 m (16 m2) sub-squares and four 1 x 1 m (1 m2) sub-sub-squares, according to the methodology proposed by Valdez Hernández (2002). In the 10 x 10 m squares, we measured the diameter at breast height (DBH) and the height of trees with DBH greater than 2.5 cm, using the clinometer methodology (Brower et al. 1998). We use the 4 x 4 m squares to measure the percent cover (Kent 2011) and average height of shrubs and juvenile trees (i.e., those with height greater than 30 cm and DBH less than 2.5 cm). In the 1 x 1 m squares we estimated the percent cover and determined the average height for woody seedlings (less

than 30 cm tall) and herbaceous species. In the center of each MU we installed a water table well (Peralta Peláez et al. 2009) to a depth of 50 cm to measure the water level and obtain interstitial water samples, from which we measured the salinity with Ultrameter II (Mod. 6PFC, Myron L). We did the vegetation and water sampling in October 2018. In addition to water table wells, to monitor hydrology, in June 2018 we installed level loggers (HOBO® model U20L) in three MUs: FWS_01, FWS_03, and FG_01. We programmed these probes to measure the water level every 30 min for slightly more than two years (until August 2020). Relative Importance Value (RIV) We obtained de RIV of each species in each MU. For tree species, we obtained the RIV using the formula:

Wetland Science & Practice January 2022 29

RIVtrees =

Relative basal area + Relative height 2

For shrub and herb species:

RIVshrub and herbs =

Percent cover + Relative height 2

We obtained the relative basal area of each tree species by adding the basal areas (in m2) of the individuals of the same species in the MU and dividing by 100. To obtain the relative height, we divide the average height of the species in the MU by the maximum height obtained in the sampling, which was 14.4 m (López Rosas et al. 2021; Moreno-Casasola et al. 2016). DATA ANALYSIS With the values of each species, we constructed a data matrix with 19 MUs and 30 species. We transform these data with square root, to later create Bray-Curtis similarity matrices that we use to place each MU in groups (CLUSTER analysis; Legendre and Legendre 1998) or in an ordination space (MDS; Faith et al. 1987; Minchin 1987). To detect differences between groups of MUs, we applied an analysis of the percentage of similarity using permutation and randomization methods of the similarity matrix (SIMPROF test; Clarke et al. 2008). To identify the determining species of each group formed, we described the groups with statistical significance obtained from SIMPROF with the support of the technique of analysis of similarity between percentages and species contribution (SIMPER test; Clarke 1993). We constructed a second matrix (abiotic) with the salinity and water level data. We transformed these data through standardization before running an indirect gradient analysis (BEST/BIOENV test; Clarke et al. 2008) to interpret the CLUSTER or MDS in relation to environmental variables. We used PRIMER 6.0 to run the CLUSTER, SIMPROF,

Figure 6. Distribution of the monitoring units in the study site. 30 Wetland Science & Practice January 2022

SIMPER, and BEST/BIOENV tests, and PCORD 6 to run the MDS ordination. RESULTS We recorded a total of 30 species, 11 trees or tree-like species, 13 herbs, and 6 shrubs or sub-shrubs. Table 2 shows the list of these species and their habit. The seven species with the highest average RIV (±1 SE, n = 19 MUs), in order from highest to lowest, were: Antelope Grass (Echinochloa pyramidalis; 15.0 ± 4.88), White Mangrove (Laguncularia racemosa; 11.8 ± 4.42), Black Mangrove (Avicennia germinans; 11.2 ± 5.21), Red Mangrove (Rhizophora mangle; 10.8 ± 4.78), Mexican Palm (Sabal mexicana; 9.6 ± 4.02), Giant Sensitive Plant (Mimosa pigra; 6.4 ± 2.69), and Pond Apple (Annona glabra; 6.1 ± 3.42). According to the composition of species, the MUs cluster into five significant (SIMPROF test; Pi = 5.73, p = 0.001) groups (Figure 7). From left to right, Group I had a similarity of 35.2%, and includes the nine MUs from flooded grassland. In this group the contribution of E. pyramidalis was 64.76 % (SIMPER test), while Mimosa pigra contributed with 21.41%. This was the only group dominated by herbaceous vegetation. Group II had a similarity of 67.4%, and included two MUs from the freshwater swamp: 100% of the contribution of this group was Annona glabra, although there was also a presence of the palm S. mexicana. This group included the MU “FWS_01” that was monospecific Annona glabra (Figure 8). Group III had a similarity of 54.53%; it included two MUs from palm forest swamp. The species that contributed the most to the formation of this group were S. mexicana (32.75%) and Guayabillo (Ginoria nudiflora; 30.12%), but the terrestrial Bullhorn Acacia (Acacia cornigera) was also present and contributed with 20.91%. Group IV had a similarity of 70.4%, and includes the four MUs from mangrove swamp that had co-dominance of R. mangle (40.70%) and L. racemosa (40.57%). A. germinans also was present in the mangrove, but with low contribution to the group (18.73%). Group V included one MU from the freshwater swamp and one from the palm swamp. This group had a similarity of 65.66% and the vegetation was a mixture of S. mexicana with mangrove species (L. racemosa and Buttonwood - Conocarpus erectus) and other trees from freshwater swamp (e.g., Strangler Fig - Ficus cotinifolia and Annona glabra). Sabal mexicana contributed 32.75%, while L. racemosa and C. erectus accounted for 27.68% and 24.57%, respectively. The 2-dimensional MDS ordination of 20 MUs and plant species (stress = 10.34 for two dimensions; final instability = 1.0 x 10-8 with 75 iterations) shows a gradient from high salinity and low water levels to low salinity and high water levels (Figure 9). Along Axis 1, the herbaceous

Table 2. List of species found in the different wetlands of this study.

Family

Species

Habit

Acanthaceae

Avicennia germinans (L.) L.

Tree

Vegetation type* FWS PFS FG x

Amaryllidaceae

Hymenocallis littoralis (Jacq.) Salisb.

Herb

Annonaceae

Annona glabra L.

Tree

Arecaceae

Sabal mexicana Mart.

Rosette tree (palm)

Asteraceae

Eclipta prostrata (L.) L

Herb

Combretaceae

Conocarpus erectus L.

Tree

Combretaceae

Laguncularia racemosa (L.) C.F. Gaertn.

Tree

Commelinaceae

Commelina sp.

Herb

Convolvulaceae

Ipomoea tiliacea (Willd.) Choisy

Herb (vine)

Cucurbitaceae

Melothria pendula L.

Herb (vine)

Cyperaceae

Eleocharis interstincta (Vahl) Roem. & Schult. Herb (sedge)

Cyperaceae

Eleocharis mutata (L.) Roem. & Schult

Herb (sedge)

Cyperaceae

Fuirena simplex Vahl

Herb (sedge)

Fabaceae

Acacia cornigera (L.) Willd

Shrub

Fabaceae

Dalbergia brownei (Jacq.) Schinz

Climbing shrub

Fabaceae

Mimosa pigra L.

Shrub

Fabaceae

Pithecellobium dulce (Roxb.) Benth

Tree

Lythraceae

Ginoria nudiflora (Hemsl.) Koehne

Tree

Moraceae

Ficus cotinifolia Kunth

Tree

Onagraceae

Ludwigia octovalvis (Jacq.) P.H. Raven

Sub-shrub

Poaceae

Echinochloa pyramidalis (Lam.) Hitchc. & Chase

Herb (grass)

Poaceae

Leersia hexandra Sw.

Herb (grass

Polygonaceae

Coccoloba barbadensis Jacq.

Tree

Pontederiaceae

Pontederia sagittata C. Presl

Herb

Pteridaceae

Acrostichum aureum L.

Herb (fern)

Rhizophoraceae

Rhizophora mangle L

Tree

Verbenaceae

Lippia nodiflora (L.) Michx.

Herb

Unknown

Unknown (Bush_sp1)

Shrub

Unknown

Unknown (Bush_sp2)

Shrub

Unknown

Unknown (Tree_sp1)

Tree

MS x

x x x x

x x

x x x x x x

x x

* FWS = freshwater swamp, PFS = palm forest swamp, FG = flooded grassland, and MS = mangrove swamp.

Wetland Science & Practice January 2022 31

Figure 7. Dendrogram of the numeric classification of the 19 monitoring units. The red color indicates groups, significantly different (SIMPROF test, p < 0.01).

Figure 8. Monospecific formation of Annona glabra, as a type of freshwater swamp (MU FWS_01). (Photo by Hugo López Rosas)

vegetation (MUs from flooded grassland) is ordered to right side, where the water levels are higher and there are low salinity levels. The MUs of the palm forest swamp are ordered on the far right of the plot, where the water level values are the lowest and salinity values are high. On this side of the graph the MUs of the mangrove swamp are also located, but slightly more to the right of the MUs from the palm swamp grove - lower salinity and higher water level than the palm swamp. The MUs from the freshwater swamp and the palm forest swamp, with exception of FWS_01 and PFS_03, had a heterogeneous species composition, sometimes mixing with palm or mangrove species. These MUs are located in an intermediate space of the gradient, although the MU FWS_02 has more affinity with the MUs of the palm grove. With the BEST we obtained that the combination of salinity and water level had a high and significant correlation (0.574; BEST/BIOENV; Rho = 0.598, p = 0.01), indicating that both variables form the principal environmental gradient in the study site. Hydroperiods show a similar pattern in herbaceous or arboreal wetlands (Figure 10), indicating hydrological connectivity. The wetlands have two flood peaks, one during 32 Wetland Science & Practice January 2022

the rainiest months (September-October), and one during the winter months when the sandy bar of the lagoon is closed and the water level rises gradually with the permanent entry of water from the river. The year 2019 presented a prolonged drought, and the water level of the wetlands did not recover until the end of the rainy season (Figure 10). The salinity and water level values of the different types of wetlands are presented in Table 3. In general, salinity values are low, even for the mangroves. According to the classification proposed by Montagna et al. (2017), the flooded grassland has oligohaline conditions and the freshwater swamp has oligohaline or slightly mesohaline conditions, while the mangrove swamp had mesohaline conditions. The palm forest swamp had a high variation, with a minimum value of 4.1 ppt (oligohaline) and a maximum of 13.6 ppt (mesohaline). The study area is a coastal plain in a tropical rainy region, receiving permanently subterranean flows from the mountain range close by and from the extensive dune system (Yetter 2004), as well as from the Caño Gallegos River. These sources supply freshwater to all the wetlands, thereby reducing salt stress. The water levels correspond to the beginning of the rainy season, with the variation in these levels due to the different topographic levels of the system (Flores-Verdugo et al. 2007) and the geomorphology of the landscape (Brinson 1993). The mangrove swamp and the flooded grassland are located in the floodplain, so they have the highest water levels, whereas the palm forest swamp and the freshwater swamp are in a topographic depression (dune slack) at the edge of the dune system, so they had low water level values. DISCUSSION The wetlands in this study are hydrologically connected and they respond to a salinity and water level gradient. The herbaceous vegetation is dominated by the African grass Echinochloa pyramidalis, and occurs in the areas with the highest flooding and lowest salinity levels. In the areas of higher salinity, the mangrove and the palm forest swamp are located, but the latter is found at higher elevations, with less flooding. In intermediate zones of the gradient is the freshwater swamp, which may support monospecific Annona glabra formations, but may also have a diverse mix of mangrove and palm elements. Figure 11 presents a hypothetical model of the zonation of the different tropical coastal ecosystems in response to the combination of flood and salinity gradients (MorenoCasasola 2016; Silva et al. 2017). Small salinity changes and flood levels result in a variety of wetlands with different species. The results of the present study confirm that these two variables are the main determinants of the distribution of vegetation in tropical coastal wetlands. Nonethe-

less, the actual delimitation between types of vegetation is complex because the species possess wide tolerances. In this study, it was possible to find mangrove species, such as L. racemosa and R. mangle, mixed in areas dominated by the palm Sabal mexicana. Also, although we observed areas dominated by S. mexicana, this species also mixes with species of flooded forest, such as Ficus cotinifolia or Conocarpus erectus. The higher species richness in the flooded grassland (16) is explained by the low salinity levels; while in the palm forest swamp its high richness (14) is due to its transitional location between oligohaline and mesohaline environments, and the low stress by inundation, sharing species with the freshwater swamp and the mangrove swamp, and with the presence of terrestrial plants. The MUs from the freshwater swamp only had six species due to the high dominance of A. glabra. This type of plant formation is rare in the coastal plain of Veracruz, and we have very little information about its natural history. The type of vegetation poorest in species (3) was the mangrove forest, because of saline stress. Rincón-Perez et al. (2020) found a similar pattern in relation to lower species richness associated with higher salinity in a gradient of coastal wetlands on the Mexican Pacific coast. In this study we did not obtain productivity data, but Infante-Mata et al. (2012) made the comparison of this variable in five freshwater coastal swamps of the Gulf of Mexico, including one in the study region, and they concluded that the litter production of this wetland type is similar to that of mangroves. The area with the highest flooding and least saline influence was dominated by E. pyramidalis. This species had the highest RIV in this study. Despite being an herbaceous species with a maximum height of 2.0 m, this species had a higher RIV than the arboreal species due to its high coverage. This species has been reported as an invasive species of freshwater wetlands in this region of the country (López-Rosas et al. 2006). Due to the presence of isolated individuals of Annona glabra immersed in the grassland, we consider that this area was originally a swamp, but the historical management in the area caused the loss of Vegetation type Freshwater swamp (n=3)

Salinity (ppt) Water level (cm) 5.9 ± 0.87 11.6 ± 1.18

Palm forest swamp (n=3)

8.0 ± 2.87

-36.3 ± 16.23

Flooded grassland (n=9)

1.2 ± 0.08

16.3 ± 1.42

Mangrove swamp (n=4)

11.6 ± 1.18

11.3 ± 1.18

Table 3. Salinity and water level by vegetation type.

Figure 9. MDS ordination of 19 monitoring units (MUs) and wetland species. Ordination based on relative importance values of species data. Abiotic data are represented by vectors. The angle and length of vectors indicates the direction and strength of the relationships between abiotic factors and ordination scores.

Figure 10. Hydroperiods for three type of wetlands in the study area from June 2018 to August 2020. The dashed line is ground level and the solid lines indicate the daily maximum (black) and minimum (gray) levels of the hydroperiods: A - freshwater swamp dominated by Annona glabra (MU FWS_01), B - freshwater swamp mixed with palm and mangrove species (MU FWS_03), and C - flooded grassland dominated by Echinochloa pyramidalis (MU FG_01).

Figure 11. Schematic representation of the zonation of coastal ecosystems in a combined salinity and flood gradient. (Modified from MorenoCasasola 2010 and Moreno-Casasola 2016) Wetland Science & Practice January 2022 33

the vegetation cover and its transformation into a flooded grassland. In this region of Mexico, freshwater wetlands are replaced by grasslands for extensive livestock farming. In mangroves this practice is not successful due to salt stress, which does not allow the survival of pasture grasses. Ecological studies of tropical coastal wetlands show a strong bias towards only one type of wetland: mangroves. However, the results of the present study, as well as those of other authors (López-Rosas et al. 2021; Rincón Pérez et al. 2020), indicate that it is necessary to recognize and understand the patterns and processes of other types of coastal wetlands, that may be connected with mangroves. The bias in the research has prevented finding those relationships that are imperative for the generation of management and conservation proposals and obscure differences in ecosystem functioning such as productivity. ACKNOWLEDGMENTS We thank Boni Azua, Luis Miguel Rodríguez, and Máximo Azua for their support with the field work and Roberto Monroy for his support preparing the map and editing the pictures. We also thank DIADA SA de CV for the funding that helped us carry out this work. REFERENCES

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Howard, R.J, and I.A. Mendelssohn. 1999. Salinity as a constraint on growth of oligohaline marsh macrophytes. I. Species variation in stress tolerance. Am. J. Bot. 86: 785–794. Infante Mata, D. P. Moreno-Casasola, and C. Madero-Vega. 2012. Litterfall of tropical forested wetlands of Veracruz in the coastal floodplains of the Gulf of Mexico. Aquatic Bot. 98: 1–11. Kent, M. 2011. Vegetation Description and Data Analysis: A Practical Approach. 2nd Edition. Wiley-Blackwell, London. Legendre, P., and L. Legendre. 1998. Numerical Ecology. 2nd Edition. Elsevier, New York. López Rosas, H., V.E. Espejel González, and P. Moreno-Casasola. 2021. Variaciones espacio-temporales del nivel y salinidad del agua afectan la composición de especies del manglar-tular. Ecosistemas y Recursos Agropecuarios 8(I): e2674. López Rosas, H., P. Moreno-Casasola, and I.A. Mendelssohn. 2006. Effects of experimental disturbances on a tropical freshwater marsh invaded by the African grass Echinochloa pyramidalis. Wetlands 26(2): 593–604. Minchin, P.R. 1987. An evaluation of the relative robustness of techniques for ecological ordination. Vegetatio 69: 89–108. Montagna, P.A., T.A. Palmer, and J.B. Pollack. 2013. Hydrological Changes and Estuarine Dynamics. SpringerBriefs in Environmental Sciences, New York. Moreno-Casasola, P., 2010. Mar de Arena. Gobierno del Estado de Veracruz para la Conmemoración de la Independencia Nacional y de la Revolución Mexicana, y Universidad Veracruzana, Xalapa, Veracruz, México. (Available in www.cienagasyhumedales.org). Moreno-Casasola, P. 2016. La zona costera y sus ecosistemas. In: Moreno-Casasola, P. (ed.). Servicios Ecosistémicos de Selvas y Bosques Costeros de Veracruz. INECOL-ITTO-CONAFOR-INECC, Xalapa. pp. 18–36. (Available in www.cienagasyhumedales.org). Moreno-Casasola, P., H. López Rosas, D. Infante Mata, L.A. Peralta, A.C. Travieso-Bello, and B.G. Warner. 2009. Environmental and anthropogenic factors associated with coastal wetland differentiation in La Mancha, Veracruz, Mexico. Plant Ecol. 200: 37–52. Moreno-Casasola, P., E. Cejudo-Espinosa, A. Capistrán Barradas, D. Infante-Mata, H. López-Rosas, G. Castillo-Campos, J. Pale-Pale, and A. Campos-Cascaredo. 2010. Floristic composition, diversity and ecology of freshwater marshes in the central coastal plain of Veracruz, Mexico. Bol. Soc. Bot. Mex. 87: 29–50. Moreno-Casasola, P., H. López Rosas, J. Vázquez Benavides, F. LópezBarrera, V.E. Espejel González, and L. Sánchez Higueredo. 2016. Restauración de un popal: estado de la vegetación y nivel de inundación después de siete años del manejo de una gramínea invasora en Veracruz. In: E. Ceccon, C. Martínez-Garza (eds.), Experiencias Mexicanas en la Restauración de los Ecosistemas. UNAM/UAEM/CONABIO, México. pp. 433–455. Perata, P., W. Armstrong, L.A., and C.J. Voesenek. 2011. Plants and flooding stress. New Phytologists 190: 269–273.

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MEXICAN WETLANDS Silva R., M.L. Martínez, P. Moreno-Casasola, E. Mendoza, J.L. LópezPortillo, D. Lithgow, G. Vázquez, R.E. Martínez-Martínez, R. MonroyIbarra, J.I. Cáceres-Puig, A. Ramírez-Hernández, and M. Boy-Tamborell. 2017. Aspectos Generales de la Zona Costera. UNAM, INECOL. Mexico. Tiner, R.W. 2013. Tidal Wetlands Primer. An Introduction to their Ecology, Natural History, Status, and Conservation. University of Massachusetts Press, Amherst.

Aquatic Groups in Freshwater Systems in the Semidesierto Queretano, Mexico: Algae, Bryophytes, Vascular Plants, and Odonata

Utrera-López, M.E., and P. Moreno-Casasola. 2008. Mangrove litter dynamics in la Mancha Lagoon, Veracruz, Mexico. Wetlands Ecol. Manage. 16(1): 11–22.

Patricia Herrera-Paniagua1, Mahinda Martínez1,2, and Olga Gómez Nucamendi1

Valdez Hernández, J.I. 2002. Aprovechamiento forestal de manglares en el estado de Nayarit, costa Pacífica de México. Maderas y Bosques 8 (Special number 1): 129–145.

INTRODUCTION Freshwater ecosystems occupy an estimated 5.4-6.8% of the global land surface and host almost 9.5% of the Earth´s described species, including one-third of vertebrates. A large proportion of aquatic biodiversity is threatened by factors such as climate change, pollution, biological invasions, infectious diseases, and salinization (Reid et al. 2019). Freshwater habitats in arid zones are especially vulnerable due to isolation, and because they have relicts and endemic species (Davis et al. 2013). Mexico has a large proportion of mountains and deserts, and as a consequence, wetlands are scarce, accounting for barely 0.6% of the world’s wetlands. Olmsted (1993) estimates that only 3.3 million hectares are wetlands, 44% of which are either estuarine or coastal. Continental wetlands span over 6,500 km2 (Berlanga et al. 2008) including dams or small reservoirs, but estimates of lotic areas are lacking (Mora et al. 2013). Querétaro is a small state in the central portion of Mexico (Figure 1) which mirrors the situation of the country. It has mountains and deserts and large wetlands are absent. Only freshwater systems are present since the state lacks coastal areas. The state has two ecoregions, the Pánuco that drains to the Atlantic, and the Lerma-Chapala which runs to the Pacific (Abell et al. 2008). A narrow strip in central Querétaro is locally known as “Semidesierto Queretano” (SQ). Floristically, it is the southernmost portion of the largest and richest Mexican arid zone, the Chihuahuan Desert (Figure 2a, b; GranadosSánchez et al. 2011; Bayona 2016). The area has been under a long human occupation by Otomis and Mestizos that survive on extensive grazing of cows and goats, and rain-fed agriculture (CONAGUA 2020). Therefore, the few water resources available in the area are under high human pressure (Bezaury-Creel et al. 2017; CONAGUA 2020). Lotic environments are represented with first, second, and third-order rivers such as the Moctezuma, Estorax, and Tolimán (Pineda et al. 2009). Natural lentic

Yetter, J. 2004. Hydrology and Geochemistry of Freshwater Wetlands on the Gulf Coast of Veracruz, Mexico. (Master’s Thesis). University Waterloo. Ontario, Canada

Laboratorio de Botánica, Facultad de Ciencias Naturales, Universidad Autónoma de Querétaro, Avenida de las Ciencias s/n, 76230, Juriquilla, Querétaro, Mexico 2 Corresponding author: mahinda@uaq.mx 1

Wetland Science & Practice January 2022 35

environments include springs, phytotelmata, and temporary ponds. Cattle watering holes and dams are also present (Díaz-Pardo 2016). A few studies have addressed the SQ aquatic biota, mostly as part of a general description of Querétaro, the Pánuco drainage, or Central Mexico. Studied groups include fish (Gutiérrez-Yurrita and Morales-Ortiz 2004), aquatic macroinvertebrates (Pineda et al. 2009; Torres-Olvera et al. 2018), Odonata (González-Soriano and NoveloGutiérrez 1996; Alonso-Eguía et al. 2002; GonzálezSoriano and Novelo-Gutiérrez 2014), and finally, algae, vascular plants, and fish parasites (Pineda et al. 2009). Monitoring biodiversity in freshwater ecosystems provides information on population status and their extinction risk (Reid et al. 2019). Therefore, systematic sampling is important to estimate the biodiversity of aquatic species per site and their conservation (Heino et al. 2009; Johnson and Hering 2009). Literature of the Pánuco watershed identified 47 algae genera, 40 species of vascular plants (Pineda et al. 2009), and 78 species of Odonata (Alonso-Eguía et al. 2002). However, only three

Figure 1. Study sites: 1 - Guamuchil, 2 - El Salado, 3 - Adjuntas de los Guillén, 4 - San Miguel Palmas, 5 - El Manantial, 6 - Tzibanzá, 7 - Pathé, 8 - Las Moras, 9 - Maconí, 10 - La Cañada, 11 - El Chilar, 12 Crucitas, 13 - La Vereda, 14 - Bomintzá, 15 - El Zapote, 16 - Gudiños, 17 - Panales, 18 - El Púltpito, 19 - El Oasis, and 20 - Rancho Quemado. 36 Wetland Science & Practice January 2022

of our sampled localities are in common with previous works for Odonata (Río Victoria-Palmas, El Oasis, and Rancho Quemado) and two for algae and vascular plants (El Oasis and Río Extórax). Bryophytes (sensu lato) had not been previously collected in our study sites The aims of our study were to 1) establish the biodiversity of macroalgae, bryophytes, vascular plants, damselflies and dragonflies in the different natural aquatic environments in the area, and 2) establish which localities are conserved or deteriorated using biota richness as an indicator.

Figure 2. View of two study sites: a) Salix riparian forest at El Chilar, Tolimán and b) spring at Rancho Quemado, Cadereyta. (Photos by Mahinda Martínez)

Table 1. Sampling sites – characteristics and summary results for hydrophilic bryophytes, vascular plant water associations, and aquatic invasives. Site

Locality

River/ Stream

Predominant cover

Hydrophilic bryophyte

Unvegetated, isolated Salix trees

Water association Aquatic 1

Aquatic invasive Sub-aquatic Riparian taxa

Guamuchil

Extoraz

El Salado

Extoraz

Salix riparian forest

Adjuntas de los

Victoria

Guillén San Miguel Palmas Victoria

Unvegetated riverbank Salix/Taxodium riparian forest

El Manantial

Spring

Tzibanzá

Moctezuma

Platanus/Salix riparian forest Unvegetated area

Pathé

Spring

Salix riparian forest

Las Moras

Moctezuma

Taxodium riparian forest

Maconí

Spring

Platanus/ Salix riparian forest

La Cañada

Tolimán

Salix riparian forest

El Chilar

Tolimán

Salix riparian forest

Crucitas

Tolimán

La Vereda

Tolimán

Unvegetated riverbank Unvegetated, isolated Salix trees

Bomintzá

First order

El Zapote

river Spring

Gudiños

Tolimán

Panales

Tolimán

El Púlpito

19 20

1 1

3 1

Platanus/Salix riparian forest Oak forest

1 1

Salix/Platanus riparian forest

El Púlpito

Taxodium riparian forest Desert shrubs

El Oasis

Spring

Salix forest

Rancho Quemado

Spring

Unvegetated area

1 4 1

Wetland Science & Practice January 2022 37

MATERIALS AND METHODS Study Area and Site Selection We sampled 20 sites in the municipalities of Peñamiller, Tolimán, and Cadereyta de Montes in the SQ (Figure 1). Site selection depended on accessibility and water seasonality. Aquatic vegetation in any of its forms had to be present, so ephemeral localities were not sampled. From the 20 sites, two sites were along the Extoraz, Victoria, and Moctezuma rivers and six for the Tolimán (Table 1, Figure 2a). Finally, we found five springs (Figure 2b) that feed the rivers. Overall, our 20 sites belong to 12 independent systems. Data Collection Fieldwork was initiated in June 2020 and ended in July 2021. Vegetation cover near water-logged areas includes riparian forest (e.g., Salix humboldtiana, Taxodium mucronatum, and Platanus mexicana), oak forest, and grasslands, with some highly degraded areas devoid of vegetation (Table 1). Table 1 indicates the sampling site, the river to which it belongs, the vegetation cover, and the water association of bryophytes and vascular plants.

Algae and plants. In each site algae and plants were collected using linear transects perpendicular to the water flow. Because of the different forms of the watersheds, we sampled two or three 50x50 cm quadrants from the river shore to the watercourse. We sampled a total of 127 quadrants. All algae observed by naked eye as well as all bryophytes and vascular plants were recorded and collected. Bryophytes and vascular aquatic plants were classified according to their association to water. Bryophytes were considered hydrophilic (sensu Glime and Chavoutier 2017) when found in water-logged microhabitats, or splashed-out places. For vascular plants, we defined their water affinities as either strictly aquatic or subaquatic using the criteria of Lot et al. (2015). We also included trees and shrubs defined as riparian by Lot (2015). Algae were analyzed under an Olympus BX43 microscope and identified to genus using Wehr and Sheath (2003) and Bellinger and Sigee (2015). Mosses were determined with Sharp et al. (1994) and Allen (2002). Aquatic vascular plants were determined using Lot (2015) and Flora del Bajío (Rzedowski et al. 2021) and Lot et al. (2015) for aquatic vegetation. Nomenclature follows AlgaeBase (Guiry and Guiry 2021), the electronic version of LATMOSS (Delgadillo http://www.ibiologia.unam.mx/briologia/ 38 Wetland Science & Practice January 2022

www/index/latmoss.html), the Classification of the Bryophyta (Goffinet and Buck, http://bryology.uconn. edu/classification/), and Tropicos (Tropicos.org). Mosses and vascular plants are deposited at QMEX (Index Herbariorum http://sweetgum.nybg.org/science/ ih/). Odonata. We collected adults of odonates flying near water bodies or on the surrounding vegetation using conventional sampling techniques with entomological nets. The sampling effort at each site was two hours. Captured specimens were injected with acetone and submersed at 100% acetone for 24 hours to preserve color (Morse 1998). Each individual was then placed in a glassine bag with the collection data. Samples were determined using Abbott (2005) Odonata Central (https://www.odonatacentral.org) and the help of specialists. Specimens are deposited at the University entomological collection (UAQ-E). Data Analyses Algae, bryophytes, vascular plants, damselflies and dragonflies were recorded in a presence/absence data matrix. Taxonomic biodiversity (expressed as species number) for each river or stream was estimated. Occurrence frequencies for each species (F) were estimated using the MouraJúnior et al. (2013) equation: In which: ni = number of sites where the i species was found N = number of sampled sites RESULTS1 Biodiversity We aggregated the results of our 20 sampling sites into 12 independent systems as shown in Table 1. Therefore, results and discussion centers around the 12 systems. We found 17 taxa of five algae groups (Cyanobacteria, Charophyta, Chlorophyta, Bacillariophyceae, and Xantophyceae); for bryophytes we found 18 taxa, 15 genera, and 11 families; for vascular plants 67 taxa, 64 genera, and 39 families, and for Odonata 28 taxa, 20 genera, and six families (Appendix). The Tolimán river, with 49 taxa was the most diverse, while El Púlpito was the least with eight (Figure 3). Not all groups were present at each site, algae were absent from Maconí, and Odonata were not found at Bominzá and El Zapote (Figure 3). Cyanobacteria (e.g., Anabaena sp., Oscillatoria sp., and Phormidium sp.) and Chlorophytes (e.g., Cladophora sp. Hydrodictyon sp. and Stigeoclonium sp.) with six taxa each were the most diverse algae. Although diatoms (Bacillariophyceae) are not filamentous, we found Synedra 1

Datasets are available from the corresponding author on reasonable request

sp. and Terpsinoe sp. (Appendix) with a large biomass that made them visible in San Miguel Palmas (Río Victoria) and Gudiños (Río Tolimán). Rio Victoria had the highest algae richness (Figure 4a-c). Six bryophyte species were classified as hydrophilic because they grow on water-logged places, either on

Figure 3. Species richness at the study sites. A - algae, P - plants (bryophytes and vascular plants), and O -odonates. Site codes: Ex Extorax river, Vi - Victoria river, Em - El Manantial, Mo - Las Moras, Pt Pathé, Ma - Maconí, To - Tolimán, Bo - Bomintzá, Ez - El Zapote, Ep - El Púlpito, Eo - El Oasis, and Rq - Rancho Quemado.

Figure 4. Some algal and bryophyte species found at the Semidesierto Queretano: filamentous algae - a) Chladophora (in the field), b) Oscillatoria, c) Vaucheria; bryophytes - d) Barbulla bolleana, e) Philonitis elongata, and f) Splachnobryum obtusum. (Photos by Patricia Herrera-Paniagua)

rocks or soil or in waterfalls or streams. They included

Amblystegium varium, Barbula bolleana (Figure 4d) and Splachnobryum obtusum (Figure 4f). The place with the highest number of aquatic bryophytes was El Oasis. Twelve strictly aquatic vascular plants were found,

notably Marsilea mollis and Zannichellia palustris. San Miguel Palmas (Río Victoria) was the richest site with five taxa (Table 1; Appendix). Subaquatic vascular plants included 15 taxa, especially Bacopa monnieri and Commelina coelestis. Tolimán was the river with most subaquatics. Five tree taxa constitute the riparian vegetation along with shrubs, such as Heimia salicifolia. Riparian forest was best developed at Maconí and El Chilar (Table 1). Five exotic invasive vascular plants were present in the area: Arundo donax, Egeria densa, Eichhornia crassipes, Plantago major, and Rorippa nasturtium-aquaticum (Appendix). Forty percent of our sampling sites contained exotic species, while Pathé, Gudiños and Rancho Quemado had two each (Table 1). The Anisoptera suborder (Odonata), had the highest diversity with 17 species belonging to 13 genera of three families (Aeshnidae, Gomphidae, and Libellulidae). Of the Zygoptera suborder we found 11 species of six genera from three families (Calopterygidae, Coenagrionidae, and Lestiidae; Figure 5 a-f). Fifty-five percent of the sampled sites had five or more Odonata. Pathé with 10 species was the

Figure 5. Frequent Odonata species at the Semidesierto Queretano: a) Acanthagrion quadratum, b) Pseudoleon superbus, c) Orthemis ferrugínea, d) Hetaerina americana, e) Argia anceps, and f) Libellula saturata. (Photos by Olga Gómez-Nucamendi) Wetland Science & Practice January 2022 39

richest place, followed by La Cañada with 9, and El Chilar with 8. The poorest sites were La Vereda and El Manantial with one species each. Odonata were absent from Bominzá and El Zapote (Figure 3). Thirteen species of all the taxonomic groups were widely distributed in the area (Figure 6). Cladophora (algae) was present at nine sites, Bacopa monnieri (vascular plant) at 8, and Libellula saturata (Odonata; Figure 5f) at 12 (see Appendix). None of the bryophytes was present at more than five localities, but Splacnobryum obtusum (Figure 4f) was recorded at three (15% frequency). Site Conservation All our taxa were widely distributed, and we did not find relicts or endemic species that could be used as a strong biological indicator of conserved sites. La Vereda (site 13, a portion of the Tolimán River) is a locality where the riparian forest has been devastated by human impact. We found one bryophyte and one Odonata, plus Chladophora (associated to human eutrophication) was present in large quantities (Figure 4a). Other degraded sites that lack aquatic vegetation cover were Tzibanzá (site 6) and Las Moras

(site 8 both part of the Moctezuma). The Extoraz (sites 1 and 2), Victoria (sites 3 and 4) and the upper portion of the Tolimán (sites 10, 11, 16, 17) had well preserved areas with aquatic and subaquatic plants (Table 1). The lower drainage of the Tolimán and the Moctezuma had several sites with low aquatic diversity that represented degraded areas because of water scarcity and probable eutrophication. We found other places with low species diversity: El Manantial (site 5), Bominzá (site 14), and El Púlpito (site 18, Table 1). There are several explanations for such diversities, or for the absence of some groups. El Manantial is a spring surrounded by riparian forest that prevents light from reaching the water. Bominzá is a first-order river fed by intermittent springs, whereas El Púlpito is a small pond surrounded by xerophytic shrubs. Therefore, in spite of their low diversity, these places do not seem to represent degraded areas. DISCUSSION The Tolimán River had the highest species diversity of vascular plants and Odonata. Since it is the river where we

Figure 6. Species frequency (in percentage) per site. A – algae, P – plants (bryophytes and vascular plants), and O – odonates. Aqu – Acanthagrion quadratum; Psu – Pseudoleon superbus; Ofe – Orthemis ferruginea; Ham – Hetaerina americana; Aan – Argia anceps; Lsa – Libellula saturata. Ver – Verbesina tumacensis; Bsa – Baccharis salicifolia; Com – Commelina coelestis; Bac – Bacopa monnieri. Vau – Vaucheria; Osc – Oscillatoria; Cla – Cladophora.

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had the largest number of sampling sites, the result might be biased. Nevertheless, adjacent areas have endemic plants (Machuca-Machuca 2017) and unique bird records for Querétaro have been reported there (González-García et al. 2004). Consequently, the river was found to support a large biodiversity, and more species are likely to be found with additional effort. The river is the most important surface water flow of the Tolimán valley and is a tributary of the Extorax and Moctezuma (CONAGUA 2020). The frequent algae genera we found are common in Mexican aquatic ecosystems, either oligotrophic or eutrophic (e.g., Ramírez et al. 2002; Carmona et al. 2016). Cladophora is found in a variety of marine and freshwater systems worldwide. While its presence might be due to cultural eutrophication, it provides habitat and food for numerous organisms (Dodds and Gudder 1992). Cyanobacteria is the largest and most diverse group of photosynthetic prokaryotes in freshwater (Bellinger and Sigee 2015) and Oscillatoria was the most ubiquitous filamentous blue-green algae in the study area. The taxon is considered pollution-tolerant and is present at high densities during the warmer months in Mexico (Ramírez et al. 2002; Cuttah et al. 2008). We found Vaucheria (Xantophytes) which is associated with oligotrophic habitats (Carmona et al. 2016), but the specimen did not have reproductive structures so determination to species level was not possible (BonillaRodríguez et al. 2013). To our surprise, the genus had not been previously collected in Querétaro, as we found it in rivers, streams, and artificial ponds. Bryophytes are considered key elements of some aquatic ecosystems, but their presence and ecology are poorly documented in many parts of the world (Stream Bryophyte Group 1999; Schevock et al. 2017). We found B. arcuata and S. obtusum, which also occur in other Mexican states; they are pantropical and widely distributed (e.g., Delgadillo et al. 2014). However, their association with water is still poorly understood (Herrera-Paniagua et al. 2018). We collected 18 species of bryophytes associated with the aquatic systems, six of which are considered hydrophilic. Several site characteristics, such as elevation, water seasonality (perennial, intermittent, ephemeral), water velocity, light incidence, and water pollution influence their frequency and composition (e.g., Fritz et al. 2007; Vieira et al. 2012; Gecheva et al. 2013; Schevock et al. 2017). Therefore, places devoid of vegetation cover, modified river banks, and general degradation probably explain why hydrophilic bryophytes are significantly less diverse in the SQ than vascular plants. Bryophytes were frequent in Salix, Taxodium or Platanus riparian forest. The few that were found in disturbed habitats were mostly associated with the few relict trees. Bryophytes are likely to have been extirpated

from La Vereda, since it is a strongly modified watershed. According to Martínez and García (2001), there are 117 species of aquatic vascular plants (ferns, gymnosperms, and angiosperms) in Querétaro. Lot (2015) estimates that 982 aquatic and subaquatic species grow in Mexico, excluding weeds that can grow for short periods of time under inundation. Therefore, Querétaro has only about 12% of the aquatic diversity of Mexico. Based on our study, the SQ had 36 strictly aquatic and subaquatic plants (Appendix) that represent 3.6% of the Mexican aquatic and subaquatic vascular plant diversity, and 30% of Querétaro. We only observed eight aquatic species in our study. This low diversity was no surprise since most of our environments were lotic - part of rivers in dry area mountains. The highest diversity of aquatic plants in Mexico is found in lentic tropical areas (Mora et al. 2013). Odonates have been used as pollution and climate change bioindicators. They are also important for determining ecological conditions and provide important environmental services such as mosquito controllers (Bried and Samways 2015). González-Soriano and Novelo-Gutiérrez (2014) reported 355 species for Mexico, but 10 states (Chiapas, Jalisco, Michoacán, Morelos, Nayarit, Oaxaca, San Luis Potosí, Sonora, Tamaulipas, and Veracruz) have over 100 species. These states are not only highly diverse because of their tropical conditions but they have also been subjected to stronger collection efforts. Alonso-Eguía et al. (2002) found 16 species in the three localities they had in common with our study sites. We did not find Progomphus belisshevi, Macrothemis pseudimitans or Orthemis discolor, but observed Ischnura denticollis (Zygoptera) and Sympetrum corruptum (Anisoptera) that had not been previously recorded for the SQ. Two of the six frequent species (Acanthagrion quadratum and O. ferruginea; Figure 6 a, c) were only rarely encountered by Alonso-Eguía et al. (2002). All the species we found have a wide geographic distribution from the southern U.S. to South America, and were therefore expected to occur in our study area (GBIF, www.gbif.org). Invasive vascular plants, a major threat to freshwater biodiversity (Reid et al. 2019), were present in several sites. Most commonly distributed were A. donax (3 localities) and R. nasturtium-aquatium (4 sites). We found two other highly invasive species (E. crassipes and E. densa) at only one site, whereas they have already invaded other aquatic systems elsewhere in Querétaro (Martinez and García 2001). The presence of these species can affect odonates as invertebrate assemblages in México are known to be sensitive to floristic composition changes (e.g., Rocha-Ramírez et al. 2007; Mora-Olivo et al. 2013; Chediack et al. 2018). Monitoring the presence of these Wetland Science & Practice January 2022 41

invasive plants in neighboring places could prevent further invasions in the study area.

Bayona, A. 2016. El Estado de Querétaro. In: R. Jones and V. Serrano (eds.) Historia Natural de Querétaro. Universidad Autónoma de Querétaro, Querétaro, pp 15-24.

CONCLUSION Our study shows that aquatic biodiversity in the Mexican semi-deserts wetlands is high, since we found 130 taxa directly associated to water. Biodiversity is still poorly known. We established new records for the algae Vaucheria, the bryophyte Philonotis elongata, and two Odonates Ischnura denticollis and Sympetrum corruptum. Species richness alone cannot be used as a conservation indicator, because not all groups were present in all sites. We found differences in community composition between the different streams. Absence and/or presence of a given species can indicate direct or indirect anthropogenic stresses, as demonstrated by bryophytes and invasive vascular plants at some sites. Our results emphasize the need of more research to improve our understanding of species assemblages. The monitoring of streams diversity and the description of environmental parameters are of crucial importance. Therefore, although difficult, different taxonomic groups can be used to monitor aquatic biodiversity. Freshwater conservation lags when compared to terrestrial ecosystems (Abell et al. 2008). The SQ aquatic systems are part of the Pánuco river basin, which has been considered threatened (Bezaury-Creel et al. 2017) because of high human water demand and inefficient water use. Our sites are further threatened by river basin alteration, invasive species, and severe drought.

Bellinger, E.G. and D.C. Sigee. 2015. Freshwater algae: identification, enumeration, and use as bioindicators.1ra Edition. John Wiley and Sons, Singapore.

ACKNOWLEDGMENTS Ana Elia Chávez Rojas helped with the algae microscopic observations, and Enrique Cantoral Urquiza confirmed the determinations. Enrique Soriano confirmed the Odonata species. Yolanda Pantoja Hernández provided Figure 1. Research funding was provided by the Universidad Autónoma de Querétaro through the grant FONDEC-UAQ02FCN2020. REFERENCES

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APPENDIX. List of algae, bryophytes, vascular plants and odonates recorded at the Semidesierto Queretano. Occurrence = X. River/Stream: Ex - Extoraz; Vi Victoria; Em - El Manantial; Mo - Moctezuma; Pt - Pathé; Ma - Maconí; To - Tolimán; Bo - Bomintzá; Ez - El Zapote; Ep - El Púlpito; Eo - El Oasis; Rq - Rancho Quemado. Water association = *hydrophilic bryophyte, **aquatic vascular plant, ***sub-aquatic vascular plant, ****riparian.

Group/Taxon

Ex Vi

Em Mo

X X

Algae Anabaena sp. Chara sp. Cladophora sp. Hydrodictyon sp. Leptolyngbya sp. Lyngbya sp. Microcoleus sp. Oedogonium sp. Oscillatoria sp. Phormidium sp. Prasiola sp. Pithophora sp. Spirogyra sp. Stigeoclonium sp. Synedra sp. Terpsinoë sp. Vaucheria sp. Bryophytes and vascular plants Acalypha sp. Amblystegium varium (Hedw.) Lindb.* Artemisia ludoviciana Nutt. Arundo donax L.*** Aster subulatus Michx.*** Azolla filiculoides Lam.** Baccharis conferta Kunth Baccharis pteronioides DC. Baccharis salicifolia (Ruiz & Pavón) Pers.*** Bacopa monnieri (L.) Wettst.*** Barbula arcuata Griff.* Barbula bolleana (Müll. Hal.) Broth.* Brachythecium occidentale (Hampe) A. Jaeger Bryoerythrophyllum campylocarpum (Müll. Hal.) H.A. Crum Bryum miniatum Lesq.* Bryum sp.

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X X

X X X

X X X X X X

X X

X X X

X X

X X X

X X

X X X

X X X X X X

Group/Taxon Callitriche deflexa A. Braun ex Hegelm.** Calyptocarpus vialis Less. Chenopodium ambrosioides L. Commelina coelestis Willd.*** Croton ciiato-glandulifer Ort. Cynodon sp. Cyperus canus Presl.*** Cyperus sp.*** Didymodon sp. Dodonaea viscosa (L.) Jacq. Dryopteris cinnamonea (Cav.) C. Christens Eclipta prostrata (L.) L.*** Egeria densa Planch.** Eichhornia crassipes (C. Mart.) Solms** Eleocharis acicularis (L.) Roem. & Schult.*** Entodon beyrichii (Schwaegr.) Müll. Hal. Erythranthe glabrata (Kunth) G.L. Nesom** Euphorbia sp. Fraxinus uhdei (Wenz.) Lingelsh.**** Gnaphalium stramineum Kunth Grimmia trichophylla Grev. Haplocladium angustifolium (Hampe & Müll. Hal.) Broth. Heimia salicifolia (Kunth) Link**** Heteranthera sp. Hydrocotyle sp.** Ipomoea sp. Iva sp. Juncus acuminatus Michx.*** Lemna gibba L.** Lilaeopsis schaffneriana (Schltdl.) Cham.** Ludwigia peploides (Kuth) P. H. Raven** Marchantia sp. Marsilea mollis Rob. & Fern.** Melinis repens (Willd.) Zizka Monstera sp.***

Ex Vi

Em Mo

X X

Ez X

X X

X X X X X

X X X

X X

X X X X X X X

X X X

X X X X X X

X X

X X X Wetland Science & Practice January 2022 45

Group/Taxon Nissolia pringlei Rose Oxalis sp. Peperomia campylotropa A. W. Hill Philonotis elongata (Dism.) H.A. Crum & Steere Philonotis uncinata (Schwägr) Brid.* Piper auritum Kunth Plantago major L.*** Platanus mexicana Moric.**** Pluchea carolinensis (Jacq.) G. Don Polygonum hydropiperoides Michx.*** Polygonum punctatum Ell.*** Prosopis laevigata (Hum. & Bonpl. ex Willd.) M.C. Johnst. Racopilum tomentosum (Hedw.) Brid. Ricinus communis L. Rorippa nasturtium-aquaticum (L.) Hayek** Rozea andrieuxii (Müll. Hal.) Besch. Salix humboldtiana Willd.**** Salvia sp. Schinus molle L. Schistidium apocarpum (Hedw.) Bruch & Schimp. Sedum sp. Selaginella sp. Sida sp. Solanum americanum L. Splachnobryum obtusum (Brid.) Müll. Hal.* Taxodium mucronatum Tenn.**** Tropaeolum majus L. Verbena litoralis Kunth Verbena longifolia M. Martens & Galeotti Verbesina turbacensis Kunth Viguiera dentata (Cav.) Spreng. Vichellia farnesiana (L.) Wight & Arn. Xanthosoma robustum Schott Zannichellia palustris L.**

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Ex Vi

Em Mo

Bo X

X X X X

X X

X X X X

X X

X X X X X X X X

X X X

X X

X X X

X X

Group/Taxon Odonata Acanthagrion quadratum Selys, 1876 Aeshna persephone Donnelly, 1961 Archilestes grandis (Rambur, 1842) Argia anceps Garrison, 1996 Argia cuprea (Hagen, 1861) Argia oenea Hagen in Selys, 1865 Argia sp Brechmorhoga praecox (Hagen, 1861) Dythemis nigrescens Calvert, 1899 Dythemis sp Enallagma civile (Hagen, 1861) Erpetogomphus crotalinus (Hagen, 1854) Erythrodiplax umbrata (Linnaeus, 1758) Hetaerina americana (Fabricius, 1798) Hetaerina vulnerata Hagen in Selys, 1853 Ischnura denticollis (Burmeister, 1839) Libellula croceipennis (Selys, 1868) Libellula saturata Uhler, 1857 Libellula sp. Orthemis ferruginea (Fabricius, 1775) Paltothemis lineatipes Karsch, 1890 Pantala flavescens (Fabricius, 1798) Perithemis domitia (Drury, 1773) Progomphus borealis McLachlan in Selys, 1873 Pseudoleon superbus (Hagen, 1861) Sympetrum corruptum Newman, 1833 Sympetrum illotum (Hagen, 1861) Telebasis salva (Hagen, 1861)

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Wetland Science & Practice January 2022 47

PANTANAL WETLANDS

Veredas: Upper Basin Wetlands Feed the Pantanal Year-round Arnildo Pott1, Vali Joana Pott2, and Suzana Neves Moreira3 INTRODUCTION Veredas4 are a unique type of wetlands found in Brazil’s Cerrado on the headwaters of streams feeding rivers of Central Brazil, such as the tributaries of the Paraguay River in the Pantanal wetland (Figure 1). They usually occur in flat valleys, although they sometimes begin on slopes. Their size in length and width is variable. Their soils are hydromorphic organic that store water like a sponge and release it gradually throughout the year, despite the seasonality of region’s rainfall. These organic soils are permanently waterlogged, often enhanced by impermeable laterite bedrock which produces high iron contents in soil and water. While veredas form small creeks that maintain the rivers running to the Pantanal lowland, in dry years or under drainage, the regularity of outflow is reduced, thereby eliminating river overflow on the adjacent plains. Low rainfall in both highland and lowland favors wildfires, such as occurred in 2020 (Damasceno et al. 2021). The severe drought continues in 2021. Therefore, veredas are now under increasing concern for the urgent need for conservation. FLORA OF THE VEREDAS Veredas shelter a high plant richness, around 1,000 species of angiosperms (Moreira 2015). The species-richest families are Poaceae, Cyperaceae, Asteraceae, Fabaceae, Melastomataceae, Lamiaceae, Rubiaceae, Eriocaulaceae and Lentibulariaceae. Some of the most abundant species are filiform graminoids, such as tussock grasses and sedges. The main grasses are Andropogon virgatus, Anthaenantia lanata, Anthaenantiopsis trachystachya, Axonopus uninodis, Eriochrysis spp., Paspalum spp., Saccharum spp., and Setaria paucifolia. Some of the most frequent sedges are Rhynchospora spp. (e.g., R. emaciata). Other filiform or slender plants reaching the top of the grassland stands belong to various families and genera, e.g., Achyrocline, Aeschynomene, Buchnera, Cyrtopodium, Habenaria, Lessingianthus, Schwenckia, Sizyrinchium, Widgrenia 1 Visiting Professor, INBIO-Instituto de Biociências, UFMS-Universidade Federal de Mato Grosso do Sul, Campo Grande, MS, Brazil. Corresponding author e-mail address: arnildo.pott@gmail.com 2 Botanist, Herbarium CGMS, INBIO, Universidade Estadual de Mato Grosso do Sul, Campo Grande, MS, Brazil. 3 Universidade Estadual de Mato Grosso do Sul, Campo Grande, MS, Brazil. 4 Vereda means path in Portuguese because it was the way of access through the Brazilian woody savanna, the Cerrado, while also providing water and grass for horses and oxen.

48 Wetland Science & Practice January 2022

and Xyris. Two endemic species were recently described: Cyperus valiae, so far only found in veredas of the Serra da Bodoquena plateau (Pereira-Silva et al. 2018), and Passiflora pottiae (Cervi and Imig 2013). A few ferns colonize the decaying straw underneath grass tussocks, like Pityrogramma calomelanos, Cyclosorus spp., and Thelypteris serrata, later to be replaced by shrubs. The most common shrubs are Byrsonima umbellata, Heteropterys subcoriacea, Ilex affinis, Ludwigia nervosa and Miconia chamissois, and some scattered treelets such as Cecropia pachystachya, Myrsine spp., and Tapirira guianensis. Treelets become denser toward the streamlet, mainly where the channel deepens, forming a creek with downstream gallery forest. Trees also occur on the vereda head, where water emerges near the slope. The palm Mauritia flexuosa is often present (Figure 1) and although its presence is not mandatory to characterize a vereda (Figure 2), it is used as an indicator for its easy recognition. Unfortunately similar wet grasslands without the palm are overlooked and not recognized as veredas and not protected (Moreira et al. 2015). The occurrence of M. flexuosa southwards seems limited by frequent frost. Another indicator species is Miconia chamissois (Figure 3) which grows in variable densities over the grassland. Most often, there are three zones in the veredas, with varying floristic compositions (Oliveira et al. 2009). The vegetation varies from open marshy grassland, mainly in the external zone, to grassy-shrubby and swampy gallery forest inwards. The ground is quite firm near the edge, with short herbs. It becomes boggy inside, often like a water mattress, and water pours to the surface, forming puddles among grassy mounds and flows slowly, mainly in animal tracks. Intermingled small aquatic plants grow in those gaps, e.g., Drosera, Mayaca, and Utricularia. In some

Figure 1. Vereda wetland with the Mauritia flexuosa palm. (Note: All photos courtesy of the authors)

areas patches of Sphagnum spp. can be found. The third zone is the bottom (the lowest elevation), where a stream collects the outflow. Temporary ponds with aquatic plants may occur within the veredas (Moreira et al. 2011).

Figure 2. Vereda wetland without the Mauritia flexuosa palm. Note the filiform grasses.

Note that while the flora of the Pantanal wetlands (Pott and Pott 2020) is distinct from the veredas, some patches of vereda occur in the eastern edge of the Pantanal lowland, only along the few short permanent streams. The occurrence of Mauritia flexuosa along the Aquidauana River, however, are just palm stands because they lack the other species characteristic of the veredas (Moreira et al. 2017; Pott and Pott 2020). FAUNA OF THE VEREDAS Veredas function as water and food source, nursery and refuge for the Cerrado fauna. Flowers occur year-round for pollinators, e.g., Ferdinandusa brasiliensis and Sinningia elatior are visited by hummingbirds. Many species are bird-spread and provide fruits for fauna, e.g., Byrsonima umbellata, Cecropia pachystachya, Mauritia flexuosa, Miconia chamissois (Figure 3), Myrsine leuconeura, Piper fuligineum and Tapirira guianensis. Some of the common animals are anaconda, urutu (Bothrops alternatus), caiman, giant anteater, tapir, and the maned wolf that is depicted on the current top-value Brazilian bill (R$200). Many birds nest in the veredas, such as the blue-and-yellow macaw (Ara ararauna), on dead stems of M. flexuosa. OTHER VALUES OF THE VEREDAS Veredas function as a sponge, filter and storage, producing clean water year-round, feeding streams and rivers. For example, the main water supply of the capital of Mato Grosso do Sul comes from veredas of the Guariroba preservation area - we can drink veredas water straight from the tap! (No filtration required.) Besides being a vital water resource, the veredas are an income source for local people. Traditional handicraft is jewelry from Syngonanthus nitens inflorescence stalk. The Mauritia palm is very useful: 1) the young leaf fiber is woven into various objects, 2) the petioles are used a very light wood for furniture, and 3) the oily fruit pulp (rich in carotene) is used to produce a sort of marmalade, ice cream, and cosmetics. The orange-colored roots of the obligate hemiparasitic plant Escobedia grandiflora is traditionally used as a natural colorant for food and medicines (Pennel 1931).

Figure 3. Shrub Miconia chamissois, an indicator of vereda wetland.

THREATS Veredas has not been under much direct human pressure. Cattle access drinking water and, in the dry season, often there is grazing. Small impoundments are often built on the outlet stream for cattle. However, the most severe disturbances are changes by drainage. Despite being protected by law as permanent preservation areas, some were recently drained for rice on the headwaters of the Sucuriu River and for other crops (oats, corn, soybean) on the headwaters of Wetland Science & Practice January 2022 49

the Prata River (prata is silver in Portuguese, alluding to the crystalline water), affecting tourist resorts by turbidity (Pott et al. 2019). On karstic ground, with underground flow and sinkholes, marshes of the Bodoquena plateau often have sawgrass (Cladium mariscus subsp. jamaicense). Drainage causes water table drop and consequent increase of ruderal annual plants, shrubs and treelets, and invasion of exotic trees such as Leucaena leucocephala. Silting from erosion of surrounding deforested slopes can kill the vereda, choking the herbaceous and woody vegetation. Trees die in standing water dammed by road embankment, including M. flexuosa. Although a wetland palm, it does not survive when its pneumatophores are drowned in stagnant water or buried. Fire may also occur, causing severe damage when the organic soil is dry and burns. Natural or human interferences induce plant succession, primarily via changes in rainfall and human activities inside the veredas or in the surrounding lands. Lowering the water table leads to increased woody plant abundance. Moreover, lack of conservation of veredas on the upper basin hinders the hydrologic regime of the Pantanal wetland. CONCLUSION Besides the species richness and the vital role of this water resource, it is crucial to preserve the vereda wetlands with or without the Mauritia flexuosa palm on the upper basin of the Pantanal to maintain the hydrology of the Pantanal floodplain.

REFERENCES

Damasceno-Junior G.A., F.O. Roque, L.C. Garcia, C.B. Ribeiro, W.M. Tomas, E. Scremin-Dias, B.H.M. Ferreira, R. Libonati, J.A. Rodrigues, F.L.M. Santos, E.B. Souza, L.K. Reis, M.R. Oliveira, A.H.A. Souza, D.A. Manrique-Pineda, and I.M. Bortolotto, and A.Pott. 2021. Lessons to be learned from the wildfire catastrophe of 2020 in the Pantanal wetland. Wetland Science & Practice 38: 107-117. Cervi, A.C., and D. C. Imig. 2013. A new species of Passiflora (Passifloraceae) from Mato Grosso do Sul, Brazil. Phytotaxa 103(1). https://doi. org/10.11646/phytotaxa.103.1.3 Moreira, S.N. 2015. Flora, distribuição e estrutura da vegetação das áreas úmidas de uma região savânica brasileira: implicações para a conservação da biodiversidade: 1-131. Tese (Doutorado em Biologia Vegetal) – Universidade Federal de Minas Gerais, Belo Horizonte. Avaiable at: https://repositorio.ufmg.br/bitstream/1843/BUBDA35JZ6/1/tese_suzanamoreira_final.pdf. Accessed on: October 01, 2021. Moreira S. N., A. Pott, V.J. Pott, and G.A. Damasceno-Junior. 2011. Structure of pond vegetation of a vereda in the Brazilian Cerrado. Rodriguésia 62: 721-729. http://rodriguesia.jbrj.gov.br/rodrig62-4/01%20 -%20ID280.pdf Moreira S.N., P.V. Eisenlohr, A. Pott, V.J. Pott, and A.T. Oliveira-Filho. 2015. Similar vegetation structure in protected and non-protected wetlands in Central Brazil: conservation significance. Environmental Conservation 42: 1-7. https://doi.org/10.1017/s0376892915000107. Moreira S.N., V.J. Pott, and A. Pott. 2017. Are Muritia flexuosa L.f. palm swamps in the Pantanal true veredas? A floristic appraisal. Bol. Museu Emílio Goeldi, Ciências Naturais 12: 221-238. https://boletimcn.museugoeldi.br. Moreira S.N., V.J. Pott, A. Pott, R.H. Silva, and G.A. Damasceno-Junior. 2019. Flora and vegetation structure of a vereda in southwestern Cerrado. Oecologia Australis 23: 776-798. https://doi.org/10.4257/ oeco.2019.2304.06. Munhoz C.B.R., C.U.O. Eugenio, and R.C. Oliveira. 2011.Vereda: Guia de Campo. Brasília, Rede de Sementes do Cerrado, v.1, 224 pp. Oliveira G.C., G.M. Araújo, and A.A. Barbosa. 2009. Florística e zonação de espécies em veredas no Triângulo Mineiro, Brasil. Rodriguésia 60(4): 1077-85. https://revistas.ufrj.br. Pennell, F.W. 1931. Escobedia: A neotropical genus of the Scrophularlaceae. Proceedings of the Academy of Natural Sciences of Philadelphia 83: 411-426. Pereira-Silva L., S.M. Hoefler, and R. Trevisan. 2018. Cyperus longiculmis and C. valiae (Cyperaceae), two new species from Brazil. Systematic Botany 43(3): 741-746. https://doi.org/10.1600/036364418X697454. Pott A., V.J. Pott, G. Catian, and E. Scremin-Dias. 2019. Floristic elements as basis for conservation of wetlands and public policies: the case of veredas of the Prata River. Oecologia Australis 23(4): 744-763. https://doi.org/10.4257/oeco.2019.2304.04. Pott A., and V.J. Pott. 2020. What is the flora of the Pantanal wetland? Wetland Science & Practice 37(4): 261-266. Ribeiro J.F. and B.M.T. Walter. 1998. Fitofisionomias do bioma Cerrado: os biomas do Cerrado. In: S.M. Sano and S. P. Almeida (eds.) Cerrado: Ambiente e Flora. Pp. 89-166. Embrapa, Planaltina.

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PLANT STRESSORS

Tolerance and Recovery Capacity of Tropical Native Tree Species to Water and Saline Stress: An Experimental Approach for Wetland Rehabilitation Wilmer O. Rivera-De Jesús1 and Elsie Rivera-Ocasio2 INTRODUCTION The accelerated urban expansion in coastal areas in combination with the effects of climate change promotes excessive loss of green infrastructure, including urban wetlands (Faulkner 2004). Coastal estuarine wetland structure and composition is mainly determined by their microclimate, hydrological dynamics, tidal influence, and edaphic composition (Lambs et al. 2015). All of these factors determine the physiological success or response capacity of wetland plant species and are identified as abiotic determinants (Lambers et al. 2008; Chaipin III et al. 2002; Figure 1). Climate change models for the Caribbean region predict an accelerated sea level rise, frequent drought periods and changes in the intensity, frequency, and distribution of the precipitation events (Lambs et al. 2015; CardonaOlarte et al. 2013; Erwin 2009; PRCCC 2013). Abiotic determinants are and will continue to change in the region. These changes could result in a greater reduction in coastal wetlands and thus, the loss of important ecological services such as flood control, climate regulation, pollutant retention, pest control, carbon sequestration, among others (Mitsh and Gosselink 2000; Farber et al. 2006; Zedler and Kercher 2005). Studies from estuarine coastal wetlands in the Caribbean and Puerto Rico have recorded frequent and prolonged periods of drought and significant increases in soil and groundwater salinity as a result of the expansion of the marine intrusion and sea level rise (Rivera-De Jesús 2019; Lambs et al. 2015; Colón-Rivera et al. 2014; Bompy et al. 2014; Flower and Imbert 2006). Soil and groundwater salinity is mainly caused by the increase in concentration of ions, especially Na+ and Cl-, which promotes reductions in freshwater availability. This condition results in water and salt stress in plant species associated with these ecosystems. Rehabilitation practices in wetlands ecosystems must consider the present and future climate changes when developing a restoration or rehabilitation project since variations in abiotic conditions will influence the tolerance 1 Assistant Professor of Biology, Department of Biological Sciences & University High School, University of Puerto Rico, Río Piedras Campus, San Juan, Puerto Rico, 00926; corresponding author wilmer.rivera2@upr.edu 2 Associate Professor, Department of Biology, University of Puerto Rico at Bayamón, Carr. 174 núm. 170, Industrial Minillas, Bayamón, Puerto Rico,00959-1919.

and recovery potential of the species. The tolerance capacity of a species refers to the resistance that it can present, maintaining its physiological functioning within a range of environmental conditions (Lambers et al. 2008; Osmond et al. 1987). Rosenthal and Kotanen (1994) define tolerance in plant species as the ability to maintain their physiological functioning and the maintenance of their biological processes, such as growth, energy processing and reproduction, within a stressful environmental range. In this study we want to establish the range of tolerance and recovery capacity of three species that inhabit estuarine coastal wetlands in the Caribbean region and are frequently used in rehabilitation projects: Pterocarpus officinalis (Jacq.), Thespesia populnea (L.), and Amphitecna latifolia (Mill.) A. H. Gentry). These species are used due to their adaptations to flooded conditions, while more information is needed about the range of tolerance of these species to water and saline stress. To achieve these objectives, we evaluated structural and ecophysiological parameters, and species survival rates at different saline treatments under greenhouse conditions. The knowledge provided by this study will help to make effective decisions and promote best practices in the use of these species in reforestation and rehabilitation projects in estuarine-coastal wetlands. METHODS

Plant Species

The three studied species - Pterocarpus officinalis, Amphitecna latifolia and Thespesia populnea - are native species commonly used in rehabilitation and reforestation projects in estuarine coastal wetlands in the Caribbean and Puerto Rico (Figure 2). Pterocarpus officinalis, a member of the Fabaceae family (Little et al. 1988; Little and Wadsworth 1995), is a tropical tree that can reach up to 20m height. Members of this species are easily recognized by its basal roots that form narrow plates extending from the trunk and the presence of roots nodules for bacteria nitrogen fixation (Little et al. 1988; Little and Wadsworth 1995). This species can occupy both riparian and estuarine coastal wetlands, with the capacity to tolerate prolonged flooding and saline conditions (Eusse and Aide 1999). Its ability to tolerate levels of salinity has been attributed to its capacity to accumulate Na in the leaf rachis and to maintain high K/Na ratios in the leaf blades (Medina et al. 2007). Nevertheless, additional information is necessary to better understand its tolerance to changes in salinity conditions and to determine how these conditions may affect the ecophysiological fitness of this species.

Amphitecna latifolia is another tropical tree species (Family Bignoniaceae) that can reach 10m height

Wetland Science & Practice January 2022 51

with relatively large leaves and round seeds between 7cm to 10cm in diameter. While it can occupy fresh water flooded areas (Gilman and Watson 1993), there is limited information about its salt tolerance although it has been reported as a halophyte in Mexico and Florida, (Meerow et al. 2001; Flores-Olvera et al. 2016). Its specific range of tolerance to salinity and water stress are unknown. Thespesia populnea is a tropical tree (Family Malvaceae) that commonly occupy coastal areas, with a wide distribution that includes inland and riparian wetlands. It is an evergreen tree with a height between 6m to 10m. Individuals of this species can grow on flooded and saline-influenced lands (Feng-ying et al. 2011). Despite its frequency in coastal areas particularly in sand dunes, few studies have evaluated the ecophysiological performance of adult trees under saline conditions (Miah 2013). Nonetheless, a recent

study found that T. populnea seedlings can survive and maintain high rates of photosynthesis and stomatal conductance even at 90% seawater (De Sedas et al. 2019). Thus, more information is needed about the tolerance and ecophysiological mechanisms used by the species to cope with changing salinity conditions in both juveniles young and adult individuals. Greenhouse Experiments To determine the range of tolerance of these species to water and salt stress, we first collected seeds from adult individuals planted in field conditions, and then germinated them under greenhouse conditions. The individuals were left to grow until they reached an approximate age of 2 to 3 years - the juvenile stage. The experiments were performed using multiple flow hydroponic systems, where four individuals of each species were planted (Figure 3). Each system was supplied with a stable concentration of nutrients, but salt concentration was varied in the order of 5ppt,

Figure 1. Conceptual model that expose how the survival of a plant species is related to physiological filters associated to local abiotic determinants. Physiological filters are influenced by regional and global changes that may occur in a temporal scale. (Modified model from Chapin III et al. 2002 and Lambers et al. 2008)

Figure 2. Study species: A) Pterocarpus officinalis, B) Amphitecna latifolia, and C) Thespesia populnea. (Photos by Wilmer Rivera-A & C,and Herbario-Universidad de Panamá-B)

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Figure 3. Experimental multiple flow systems, salinity treatments and distribution of the individuals by species type.

Figure 4. Temporal variation in the number of leaves in all the study species as a function of experimental treatments: 0ppt (dark line), 5ppt (red line), 15ppt (blue line) and 25ppt (green line). Each point represents the total number of leaves counted per species for each period and per treatment. The red vertical line identifies the recovery period from week 20 to week 52.

Figure 5. Temporal variation in stem size (A, B, and C) in all the study species as a function of experimental treatments: 0ppt (dark line), 5ppt (red line), 15ppt (blue line) and 25ppt (green line). Each point represents the average stem size per species for each period and per treatment. The red vertical line identifies the recovery period from week 20 to week 52.

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15ppt and 25ppt (Figure 3). These salt concentrations were established based on data collected during field conditions in wetlands where these species are used for reforestation purposes (Rivera-De Jesús 2019). A control group (0ppt) was maintained for each of the species to establish comparisons with the experimental groups, in terms of structural and ecophysiological parameters and survival rates. The experiments were performed for 5 months, and at the end of the eighth, fifteen and twenty weeks, structural measurements were made in terms of number of developed leaves and stem height(cm)., We also counted the number of dead individuals during the experiment to establish survival rates for each species. In terms of ecophysiological parameters we evaluated the net carbon assimilation (A) in μmol CO2 m-2 s-1, the stomatal conductance (gs) in μmol m-2 s-1 and the long-term water use efficiency (WUE) through the isotopic signal of δ13C. The net carbon assimilation and the stomatal conductance were determined using a portable gas exchange system, CIRAS-3-Photosynthesis System. High values of net carbon assimilation (A) indicate a greater degree of stomatal opening at the foliar level, while high values of stomatal conductance (gs) indicate a high rate of net carbon assimilation by the plant. Long-term (WUE) can be determined by the concentration of the 13C isotope in leaf tissues. Medina and Francisco (1997); Lin and Sternberg (1992) and Farquhar et al. (1989) demonstrated that tolerant species, to avoid the effects of water and saline stress, increase and maintain a stable WUE, which is positively correlated with an increase in the concentration of carbon 13 isotope (δ13C) in leaf tissues. After five months of experiment, in the surviving plants, we removed the experimental salinity treatments with continuous flows of fresh water to evaluate the recuperation capacity of these species. The recovery period lasted until week 52 from the final week of the experiment (week 20). In the recovery period we evaluated the same structural parameters (number of leaves and stem height) and the same ecophysiological parameters (A, gs and WUE) to compare with the experimental period. Statistical Analysis To determine differences between salinity treatments and between experimental periods on structural changes and ecophysiological parameters we did an analysis of variance for each species. To establish the range of tolerance of the species to salinity and hydric stress, we compared the A, gs and WUE between experimental periods. We compared the mean differences in the number of leaves, stem height and survival rates for each species. To establish the recovery capacity of the species to water and saline stress we compared the same structural and ecophysiological parameters 54 Wetland Science & Practice January 2022

Figure 6. Relationship between stomatal conductance (A) and net carbon assimilation (gs) (n = 97, R2 = 0.55), p <0.001 according to the statistical regression model Mallows’s Cp. The points in the dispersion represent values of gs and A among the study species in all the evaluated periods in which the measurements were made.

between the final period of experiment and the recovery period. We used the Tukey-Kramer post-hoc test with a significance level of p < 0.05. Statistical analyzes and figures were developed using Info-Stat and Past 4v. RESULTS AND DISCUSSION Structural Variations: Leaves and Stem There were differences in the structural variables from individuals of each species in the different saline treatment and in the time in which these treatments were sustained. The temporal variation in the number of leaves resulted in considerable reductions when the salinity levels and the exposure time increased for all species (Figure 4). In control groups, all the species showed the highest number of leaves, which indicates that the saline condition impacts important plant structures such as leaf tissue (Figure 4-dark lines). Wright and Cornejo (1990) demonstrated that in the face of low water availability, many species present a high loss in their leaf tissue to avoid the effects of water limitation, the excessive loss of water by transpiration and the limitation in biomass production under these stressful conditions. In comparison with the other two species, A. latifolia showed a considerable reduction in the total number of leaves in all saline treatments: 5ppt- (91 to 75) ; 15ppt- (91 to 76) ; 15ppt- (75 to 3) (Figure 4). The greatest loss occurred at 25ppt, with an abrupt fall in the number of leaves from the eighth week of experiment to a substantial removal of the leaf tissue towards the final period of the experiment (week 20). Among all species A. latifolia showed

Figure 7. Temporal variation of stomatal conductance (gs) (graphs A, C and E) and net carbon assimilation (A) (graphs B, D and F) in all study species as a function of the experimental treatments: 0ppt (dark line), 5ppt (red line), 15ppt (blue line) and 25ppt (green line). Each point represents the average obtained in (gs) and (A) by species type and experimental salinity treatments. The red vertical line identifies the recovery period from week 20 to week 52.

the lowest tolerance to water and salt stress. Both P. officinalis and T. populnea maintained a more stable number of leaves in the treatments ranging from 0ppt to 5ppt and showed less pronounced losses compared to A. latifolia in the high salinity treatments at 15ppt and 25ppt (Figure 4B and 4C - blue and green lines). The reduction in number of leaves become more evident from week 15, particularly in P. officinalis compared to T. populnea. In Thespesia the presence of leaves was observed in all experimental treatments, including the extreme salinity treatment (i.e., 25ppt) and also during the recovery period (Figure 4C). All of the species showed continuous growth in stem size at all experimental treatments during the first weeks,

but there was a reduction at week 20 (Figure 5). A. latifolia showed an earlier reduction in stem size on week 15 in the 15ppt and 25ppt treatments (Figure 5A - blue and green lines). The hydric and salt stress were more pronounced in this species and produced a high impact in the stem size compared to the other species. During the recovery period, the effects that these saline treatments exerted in Amphitecna, in terms of number of leaves and stem height, prevented a recovery, suggesting that its recuperation capacity is limited. Ecophysiological Response to Water and Saline Stress Stomatal conductance (gs) and net carbon assimilation (A) were different on each species at the experimental treatWetland Science & Practice January 2022 55

have a greater ecophysiological tolerance and a greater recovery capacity to water and salt stress by maintaining their photosynthetic structures (leaves), a greater degree of stomatal opening (gs) and a high net carbon assimilation (A). It is important to highlight that during its recovery period, T. populnea showed carbon assimilation rates equals to the initial values when the experimental period was started (Figure 7E and 7F). In this species, values of (A) rised significantly, relative to the other species, in the order of 8 to 12 μmol CO2 m-2 s-1, during the recovery period.

Figure 8. WUE comparisons between the beginning of the experimental period (week 0), the end of the experimental period (week 20) and the recovery period (week 52), by means of the isotopic signal of δ13C, by species type and treatments. The (*) represent significant difference in (WUE-δ13C) between periods, according to the Tukey-Kramer test, p < 0.05.

ments. The values showed a direct proportional relationship between stomatal conductance and the net carbon assimilation in all the species (Figure 6; n = 97, R2 = 0.55, p <0.001). This significant relation indicates that high values in stomatal conductance are positively related to elevated net carbon assimilation, product of a greater stomatal opening. Through the experimental treatments we found reductions in both gs and A in all species as the saline and hydric stress increased, particularly in the experimental treatments of 15ppt and 25ppt (Figure 7). The values that we found in gs and A showed a greater tolerance and recovery capacity to water and saline stress in T. populnea, followed by P. officinalis and a greater vulnerability in A. latifolia. In the most tolerant species, high values of gs and A were observed, including at the highest salinity treatments of 15ppt and 25ppt (Figures 7C to 7F). These values were maintained throughout the entire experimental period and showed a recovery, mainly in the net carbon assimilation during the recuperation period towards week 52 (Figures 7C to 7F). This implies that T. populnea and P. officinalis 56 Wetland Science & Practice January 2022

In terms of water use efficiency (WUE), through the isotopic signal of δ13C, T. populnea presented the lower and more consistent WUE at all experimental treatments compared to the other species including the recovery phase (mean = -30 to -28; Figure 8C). All species showed an increase in WUE due to water and saline stress, but P. officinalis and A. latifolia showed a significant increase in WUE from the initial to the final period of the experimental weeks and even in the recovery phase, mainly in the extreme salinity treatments at 15ppt and 25ppt (Figure 8A, mean = -28.5 and Figure 8B, mean = -26). The increase in WUE reflected in P. officinalis and A. latifolia demonstrated the high sensitivity of these species to the increase in water and salt stress and to the persistence of these conditions. The increase in WUE explains the reductions in gs and A, which is related to a low tolerance and recovery capacity to water and saline conditions. In T. populnea, the low WUE, relative to the other species, reflected the highest tolerance to salinity and water stress of the species, which is directly related to high values in gs and A throughout all the treatments including the recovery period (Figure 8C). The WUE observed in Thespesia and its relationship with gs explains the greater net carbon assimilation (A) reflected in this species, including at the highest salinity treatments and during the recovery phase, and therefore explains the structural stability and survival rates reflected by the experiment. The consistency in WUE, gs, and A in T. populnea shows its greater ecophysiological tolerance to water and salt stress and its greater recovery capacity to these conditions. Survival and Salinity Range of Tolerance The mortality showed by all species tells us about the specific range of tolerance that they show to different salinity levels. Mortality in our species increased considerably from the week 15 of the experiment to between 50% to 70%, mainly in the saline treatments at 15ppt and 25ppt. A. latifolia and P. officinalis experienced the lowest survival rates (0%), particularly in the highest salinity treatment at 25ppt (Figure 9).A. latifolia showed the earliest mortality, at 8th week of experiment, also in the lowest saline treatment

at 5ppt.. The lowest survival rate demonstrated its high sensitivity to water and saline stress relative to the other species. Rivera-Ocasio and others (2007) found that species with a lower tolerance to water and salt stress experienced higher mortality associated with physiological problems due to the toxic effect of salinity. In our study T. populnea had the survival rates (Figure 9). At the extreme salinity treatments, the total mortality in this species did not exceed 50%, therefore, it is the only species that had live individuals at the end of all the experimental treatments (Figure 9). The highest survival rates reflected in this species can be explained by its ecophysiological properties. Such trends are consistent with experimental studies evaluating net carbon assimilation rates (A) and stomatal conductance (gs), where an increase in water deficit reduced both parameters significantly, leading to increased mortality (Angelopoulos et al. 1996). CONCLUSION Increased water and salt stress affects plant structure and ecophysiological response as well as survival rates of P. officinalis, T. populnea and A. latifolia. The limitation in freshwater availability due to an increase in salinity promotes differentiated effects in terms of A, gs and WUE. Compared to the other species, T. populnea showed better structural and ecophysiological responses. The survival rate observed in this species, even under extreme conditions, is the product of its ability to preserve its stomatal opening and maintain its net carbon assimilation, avoiding the effects of water and saline stress. The maintenance in the

Figure 9. Survival rates and salinity range of tolerance in all study species. Purple line represents Pterocarpus officinalis, orange line Amphitecna latifolia and green line Thespesia populnea.

net carbon assimilation (A) allows this species to constantly generate biomass for its survival and structural development (Miah 2013; Atkinson et al. 2000, 1999). Species tolerant to water deficit not only show a better stomatal opening under a low water availability, but also a high carbon assimilation, which is reflected in the production of biomass, structural development and survival of the plant within an extreme environmental condition (Maréchaux et al. 2015). In A. latifolia the stomatal closure experienced by this species limited the entry of CO2, affecting the rate of photosynthesis and forcing the plant to use the carbon that has already been synthesized into organic compounds to maintain its basic cellular functions (respiration) (Pezeshki et al. 1996). When a plant is in a physiological respiration, it implies that the net photosynthesis rate is limited, reflecting A values equal to 0 μmol CO2 m-2 s-1 or less, as was the case of A. latifolia. In terms of the recovery capacity to water and saline stress, both T. populnea and P. officinalis demonstrated the highest ecophysiological recovery as shown by their net carbon assimilation, maintaining their stomatal opening, and their survival, even during the recovery phase. However, we do not know for how long these more tolerant species could persist under these water and saline conditions. Considering the survival rates, we found that the range of tolerance to saline conditions varied between the species. T. populnea exhibited the highest range of tolerance to salinity from 0ppt to 25ppt (Figure 9). In the case of P. officinalis we established a tolerance range from 0ppt to 15ppt and for A. latifolia a lesser range from 0ppt to 5ppt (Figure 9). The optimal tolerance range for all the species generally occurs between 0ppt to 5ppt, extending up to 15ppt in the case of T. populnea (Figure 9). Addressing changes in environmental conditions experienced in wetland ecosystems associated with climate change and their impacts on biodiversity are challenges that any rehabilitation and reforestation effort must consider. This work can be used to identify species that exhibit a great environmental plasticity as well as those with a high vulnerability to environmental stressors. For conservation and management purposes vulnerable species could then be relocated to areas with optimal conditions for their development and allows the identification of habitat conditions that will likely promote the successful establishment of these species. Rehabilitation practices should consider the response capacity of species (including ecotypes) and focus on selecting tolerant individuals with a greater environmental plasticity. Wetlands experience changes in their abiotic conditions as a result of climate change and anthropogenic impacts. Consequently, our experimental approach and similar studies can be used to help improve rehabilitation practices in these dynamic ecosystems. Wetland Science & Practice January 2022 57

ACKNOWLEDGMENTS The author of this study appreciates the collaboration of the “Corredor del Yaguazo Inc.”, and its president Mr. Pedro Carrión for the technical and logistics support offered in the experimental work. To Mr. Larry Díaz, UPR-CATEC-EcoLab technician and the lab technicians of the Puerto RicoInternational Tropical Forestry Institute for the leaf samples processing for isotopic analysis. Financial support for this study was provided by the Department of Environmental Sciences at the UPR-Río Piedras, UPR-CATEC, Puma Energy Caribe, the Ford Foundation, and the University of Berkeley.

Feng Ying, Q., L. Bao Wen, and F. Ming. 2011. Salt tolerance of semimangrove plant Thespesia populnea seedlings. Forest Research Beijing 24(1): 51-55.

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Experiences in the Restoration of Tropical Freshwater Swamps

Patricia Moreno-Casasola1, Hugo López Rosas2, Judith Vázquez Benavides1, Oscar Sánchez Luna 1, Marco Rivera-Ocasio, E., T.M. Aide, and N. Ríos-López. 2007. The effects of salinity on the dynamics of a Pterocarpus officinalis forest stand in Puerto González Nochebuena1, Laura D. Aguirre1, and E. Sánchez Rico. Journal of Tropical Ecology 23(5): 559-568. García1 Rosenthal, J. P., and P.M. Kotanen. 1994. Terrestrial plant tolerance to herbivory. Trends in Ecology & Evolution 9(4): 145-148.

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A variety of tropical wetlands is found on the floodplains of coastal regions in Mexico, including mangroves, swamps, and marshes. The first (mangroves) are well known and have received much scientific and social attention, including research, inclusion in blue carbon programs, an appreciation of their role in fisheries, and restoration efforts. However, tropical coastal freshwater wetlands have been neglected, and members of society and government have little awareness of their benefits. Tropical coastal freshwater forested and herbaceous wetlands like the mangroves represent important carbon deposits (Sjögersten et al. 2021), store water and help reduce flood peaks (Campos et al. 2011), are important for fisheries, and produce fibers, wood, and other resources that are used by local people (González-Marín et al. 2012; VázquezGonzález et al. 2015), among other environmental services. They are frequently used for raising cattle, while swamps are logged, and grasses are introduced as cattle fodder, some of them exotic African grass species. Marshes feed cattle during the dry season, and aggressive grasses are planted to induce changes in plant composition. Tropical freshwater forested and herbaceous wetlands have also been drained for housing projects, thereby reducing the area of freshwater wetlands, mainly swamps. While wetlands provide diverse and invaluable ecosystem services, their extent worldwide is decreasing and the condition of many of the remaining wetlands is deteriorating worldwide. Approximately 62% of coastal wetlands have been lost globally (Ramsar 2015). For example, in Mexico alone (the focus of this study), an estimated 7 million ha of herbaceous and forested wetlands have been lost, 33% of which are in coastal municipalities (Landgrave and Moreno-Casasola 2012). Restoring tropical freshwater swamps is becoming increasing important given continued threats. In this paper we are putting together the results obtained by different members of our working group. We will present a picture Red de Ecología Funcional, Instituto de Ecología, A.C., Carretera Antigua a Coatepec No. 351, El Haya C. P. 91073, Xalapa, Veracruz, México; corresponding author: patricia.moreno@inecol.mx., 2 Academia de Desarrollo Regional Sustentable, El Colegio de Veracruz, Carrillo Puerto 26, Zona Centro, Xalapa, 91000, Veracruz, México. 1

Wetland Science & Practice January 2022 59

of the problems encountered and the possible alternatives during the restoration of the freshwater forested wetland of Ciénaga del Fuerte. STUDY AREA: CIÉNAGA DEL FUERTE PROTECTED NATURAL AREA (CFPNA) On the Gulf coast, in Veracruz, there once were many swamps (freshwater forested wetlands) - today only a few sites remain. One of them, Ciénaga del Fuerte Protected Natural Area (CFPNA), is in the county of Tecolutla (Figure 1). The climate in the region is wet and warm with three seasons: rainy from July to October, cold with northerly winds and rain from November to February, and dry from March to June (Infante Mata et al. 2012). The mean annual temperature is 24 °C, and the total annual precipitation is around 1450 mm (Figure 1). While Ciénaga del Fuerte Protected Natural Area is a swamp largely dominated by the money tree (Pachira aquatica), it also contains patches of cattail marshes dominated by Typha domingensis along with Cladium

jamaicense (Jamaica swamp grass), and Cyperus giganteus (Mexican papyrus), and broad-leaved marshes that include Pontederia sagittata (pickerelweed), Thalia geniculata (alligator flag), Sagittaria lancifolia (bulltongue arrowhead), and Leersia hexandra (southern cutgrass or swamp rice grass) (Figures 2 and 3). CFPNA is separated from the coast by a field of low parallel dunes used for cattle ranching. Different freshwater forested wetlands are distributed along a gradient with changes in the elevation (topography) and duration of flooding. Pachira aquatica dominates areas where inundation can last up to six months and water currents are present, while pond apple (Annona glabra) is found in depressions where flooding lasts two to four months. Swamps dominated by fig trees are the driest lowlands with only one to two months of flooding. Hydroperiod is one of the characteristics that define

Figure 2. Aerial image from November 2014 showing Ciénaga del Fuerte Protected Natural Area: mostly swamp (dark green area) with considerable marsh (brown and light green areas) on the northern and southern borders. (Source: Pleiades) The upper photograph shows the swamp and the sandy area towards the water. The lower photograph shows swamp patches imbedded in a marsh matrix. (Photos by Roberto Monroy)

Figure 1. Overview of the study site and its climate and hydrology: a) location of the study site (Ciénaga del Fuerte Protected Natural Area) on the coast of Veracruz, Mexico, on the Gulf of Mexico with an aerial image showing a strip of swamp and several smaller patches immersed in an herbaceous matrix (Source: Google Earth; letters indicate study sites by the river = R and on the border of forest fragments = F), b) precipitation and temperature data for the area, and c) water level fluctuation in the study area - the dashed line represents the soil level and the blue line is the water level. As shown in c) the water level starts rising in August and remains high until the dry season begins in February or March. Note that after the rains in the area stop, the area continues to receive water that slowly flows down the Sierra Madre Occidental mountain range. 60 Wetland Science & Practice January 2022

Figure 3. Major plants of coastal freshwater wetlands in Mexico. Left to right, top line – marsh plants: Thalia geniculata, Pontederia sagittata, Cyperus giganteus; lower line – swamp plants: Pachira aquatica, Annona glabra, and Ficus species. (Photos by Gerardo Sánchez Vigil)

Figure 4. Water level fluctuation in two types of tropical coastal wetlands. On the left is a swamp dominated by Pachira aquatica (data from February 2019 to August 2020). On the right side is a Leersia hexandra grass marsh (data from March 2020 to June 2021). The latter was flooded longer, had higher levels of flooding, and the roots remained saturated most of the time, whereas the swamp experienced significant drops in the water level during the dry season as in 2019.

wetland types. Mitsch and Gosselink (2000) refer to it as the wetland signature. Figure 4 shows fluctuations in water level graphs for two freshwater wetland types located in the natural protected area of CFNPA: Pachira aquatica swamp and Leersia hexandra grass-dominated marsh surrounding the swamp. Direct human impact in the study site includes the expansion of human settlements and productive rural activities (mainly crops and cattle ranching), along with the construction of the Poza Rica-Veracruz coastal highway. These activities have caused changes in the hydrological regime and in the structure and composition of the original vegetation of the protected area (Programa de Manejo 2002). Swamps trees are felled to increase the area for cattle ranching, and ditches are sometimes built to change watercourses and reduce flooding. The highway also acts as a barrier to water flow as there are insufficient water passes, particularly during tropical storms and hurricanes. Hurricanes have also significantly degraded the site (Robledo 2013). Tecolutla is one of the areas in Veracruz where more hurricanes and tropical storms reach land. Tropical depression Number 11 (October 1999) changed the location of the outlets where water flowed out of the swamp, and Hurricanes Dean (August 2007) and Lorenzo (September 2007) and tropical storm Barry (June 2013) also had significant impacts. Their strong winds knocked down numerous swamp trees, many covered by lianas, thus increasing the damage and connecting the clearings that had progressively resulted from diverse climatic and anthropogenic events in previous years. In these clearings,

a species-poor herbaceous community established – one dominated by the grass Leersia hexandra, and the herbaceous climbers Ipomoea tiliacea and Ipomoea indica. IS THE SWAMP REGENERATING? We worked in the several swamp forests in Veracruz to understand their floristic composition and variability (Infante-Mata et al. 2011), productivity (Infante-Mata et al. 2012) and carbon storage in the soil (Marín-Muñiz et al. 2014; Moreno-Casasola et al. 2017; Sjögersten et al. 2021). More recently, our work in this tropical freshwater swamp is focused on disturbed areas where the native grass Leersia hexandra dominates open areas, creating a grass matrix surrounding patches of Pachira swamp forest (Figure 5). Leersia hexandra thrives, forming cushions of dry matter that cover the soil, thereby creating a potential obstacle to seed dispersal (Vázquez-Benavides et al. 2020). While grasses (native and exotic) and trees are both commonly found on tropical floodplains, grasses expand very quickly and outgrow tree seedlings. We have seen this in the case of swamps affected by hurricanes, in felled areas transformed to pastures for cattle ranching, and where exotic grasses have been introduced as cattle fodder in wetlands. METHODS To understand if there was any possibility for natural regeneration of forest to occur in the Leersia matrix, we set up transects in the grass matrix in two zones: close to the river (R) and bordering the tree fragments (F) (Figures 5 and 6). We quantified the seed and seedling presence, survival, and the growth of seeds and seedlings found in Wetland Science & Practice January 2022 61

Figure 5. Study sites and field research design. The figure represents the layout of seed dispersal, germination treatments, and the establishment of Pachira aquatica seedlings in two zones (R = next to river; F = next to fragment) of the three study sites in each zone (R1, R2, R3, F1, F2, F3). Transect position is displayed as two red bars, and consisted of 3m x 1m monitoring units in which natural arriving seeds and established seedlings were counted. In addition, two seedlings and three seeds were placed in these same quadrats (on the transects) and monitored at different distances from the river (R) and the swamp fragment (F). The photograph shows the cushion formed by Leersia hexandra. (Images on the left from Google Earth; Photos by Judith Vázquez-Benavides)

situ and those variables for experimentally introduced seeds and seedlings in the field. We monitored their germination and seedling survival and development, as well as that of the transplanted seedlings. We set up monitoring units along transects in areas close to the river (R) where seeds arrive by currents and in the border of swamp fragments (F) where adult Pachira trees are producing seeds (Figure 5). 62 Wetland Science & Practice January 2022

RESULTS There was a negative relationship between the number of seeds and established seedlings and the distance to the river or fragment and the grass (Figure 6). West Indian marsh grass (Hymenachne amplexicaulis) is more abundant on the river border and does not form a grass cushion. It was present in the first meters of R1 and R2 transects, and

Figure 6. The number of naturally dispersed Pachira aquatica seeds, established seedlings (naturally established seedlings), and non-established seedlings (seedlings planted manually) found in the transects (black line) and the average thickness of the grass cushions (gray dashed line) at the sites next to a fragment (F1, F2, F3) and the areas next to the river (R1, R2, R3). See Figure 5. (Redrawn form Vázquez-Benavides et al. 2020.)

dominated R3 transect. Leersia hexandra has a different habit and accumulates dead plant material forming a cushion that acts as a barrier to seed dispersal. In R3, the grass cushion was non-existent because of the presence of the grass H. amplexicaulis, thus seeds can travel further inland. Thus, in zone R, the presence of the grass cushion was more variable between sites. In the F zone, this cushion was always present. In both zones, the thickness of the grass cushion was a function of distance, from the river and from the forest fragment. Seeds and seedlings were found at a greater distance from the river than in the sites next

to the fragments. In the F zone, more seeds and seedlings were found (241), especially in F3 (204), than in the R zone (103), and the seeds and seedlings did not travel more than 5 m, as reported by Vázquez-Benavides et al. (2020). When seeds were sown after clearing the grass cushion, germination success was high, and the transplanted seedlings had a greater survival rate and final height than the seedlings sprouting from the planted seeds. So, these stages are not limited in the area once the grass has been removed. Seedling survival rates were inversely related to grass cover, showing that seedlings overgrown by grass had Wetland Science & Practice January 2022 63

low survival rates (Vázquez-Benavides et al. 2020). Thus, L. hexandra cover had a negative effect on both types of seedlings. We can conclude that the native wetland grass L. hexandra dominates in open areas and presents an obstacle to Pachira aquatica seed dispersal by acting as a physical barrier that traps the seeds in its dry biomass, preventing them from reaching the soil and effectively arresting succession. Leersia hexandra also limits their germination and seedling. When seeds are sown on bare soil opening the grass cushion, germination success is high, so this stage is not limited. These seedlings established, grew, and had good survival rates, showing that in general the environmental conditions are suitable for seedling recruitment and survival (Figure 7; Vázquez-Benavides et al. 2020). The same happened with seedlings transplanted directly. However, seedling survival rates were inversely related to grass cover, and seedlings overgrown by grass had low survival rates CAN PRIOR TREATMENTS HELP RESTORATION? Planting seedlings is not always effective for ensuring their establishment and survival. Many times, it is necessary to pre-treat the area to reduce the presence of competing plants. In this section, we evaluate the survival and growth of P. aquatica seedlings after their planting with and without previous treatments.

METHODS To evaluate the effectiveness of different treatments to reduce and control herbs and climbers, and to promote the establishment of P. aquatica in open areas dominated by the grass Leersia spp., we planted Pachira saplings in quadrats subjected to different manipulations. We used 20 plots (3 × 3 m each). The vegetation of the corridors surrounding the plots was cut to ground level with a machete and hook, and maintenance was carried out every three months. The herbaceous plants from one plot did not enter another, nor were there any plants from outside of the experimental area. Four P. aquatica saplings were planted in each plot, with 20 saplings per treatment (Sánchez Luna et al. 2021). Saplings had an average height of 70.05 ± 9.36 cm at the time of planting. The treatments we used were: a) Control - the area was covered by the dominant native grasses, with no maintenance (cutting, weeding) or intervention of any kind; b) Cut - herbaceous vegetation in the plot was cut to ground level (with a machete and a hook) prior to planting the saplings and again two days after tree planting. The cut plant material was removed from the plot, but there was no maintenance afterwards; c) Vegetation Mulch - the surrounding vegetation was cut as in treatment b, and the plant material was chopped and piled on the plot, around the sapling, as mulch. After eight months, the cutting was repeated and climbers removed (mainly I. tiliacea and I. indica) from the trees, d) Plastic Cover - the surrounding vegetation was cut to ground level as in the previous

Figure 7. Germination percentage and seedling survival rate (percentage) of seedlings from seeds sown on bare soil and germinating along the transect, and seedlings transplanted directly. Result for transects at the sites next to a fragment (F1, F2, F3) and the areas next to the river (R1, R2, R3) can be seen. In all cases there was seed germination and seedling establishment. 64 Wetland Science & Practice January 2022

treatments, this plant material was removed, and a black plastic sheet was placed on the ground. After eight months, maintenance was applied once. In all treatments, sapling survival was recorded at the beginning of the experiment and every three months for two years. Tree height, crown cover, and basal diameter (at 5 cm from the ground) were measured right after planting and then two years later. To evaluate the effect of the treatments on the herbaceous vegetation cover in each plot, the number of species, height, and species cover of the herbaceous vegetation was recorded at 12 and 24 months (Sánchez Luna et al. 2021). RESULTS Our findings show that herbaceous vegetation can reduce seedling establishment and growth (Table 1). In the Cut and Vegetation Mulch treatments, the herbaceous vegetation took two to three months to recover (one trimester), giving the P. aquatica saplings a better chance to survive. In the Plastic Cover treatment, the herbaceous vegetation took even longer to recover. The final height of the herbaceous vegetation was 140 cm in Control, 200 cm in Cut, 90 cm in Vegetation Mulch, and 70 cm in Plastic Cover (Table 2). This means that, to increase the survival of the seedlings, it is necessary that they have sizes of more than 125 cm in height at the time of planting. If seedlings of less than 125 cm are planted, to increase their survival the herbs should be pruned continuously, or the growth of the herbs should be reduced with a padding or by covering with black plastic. While all the treatments had a positive effect on the survival and growth of the P. aquatica saplings, the Plastic Cover treatment produced the best results. When the costeffectiveness is considered, the Vegetation Mulch approach is the best for controlling the herbaceous layer because it gives P. aquatica a growth advantage at a lower cost and does not generate waste (plastic) that takes decades to degrade (Table 3). Even though the establishment of P.

aquatica plantings reduced the cover of the herbaceous vegetation, maintenance, especially the removal of climbing plants, is also essential for the successful establishment of the tree saplings (Sánchez Luna et al. 2021). ARE CLIMBERS OF THE GENUS IPOMOEA AFFECTING PLANTED JUVENILES OF THE TREE PACHIRA AQUATICA TREES? Globally, woody and herbaceous climber proliferation is becoming frequent in forest gaps and disturbed forests (Allen et al. 2007; Murphy et al. 2016). At its worst, this phenomenon can lead to the replacement of forest vegetation by a dense layer of climbers that prevents ecological succession, even when active restoration efforts are underway, which is happening in the Ciénaga del Fuerte restoration site. Thus, the removal climbers is considered necessary to improve tree recruitment, growth, and survival (César et al. 2016). These climbers grow using the trees as supports, and in doing so they both shade and damage the trees (Figure 8), which can be considered as a parasitic structural relationship (Schnitzer et al. 2000; Tang et al. 2012). Aguirre (2019) studied this interaction under restoration conditions by evaluating the regeneration capacity and biomass accumulation of I. tiliacea and I. indica under contrasting conditions of supports and light availability, as well as the effect of herbaceous vegetation removal as a control strategy. From that study, it became evident that the planted saplings provided a new source of supports and enhanced the biomass accumulation of climbers. Ipomoea tiliacea presents a great challenge because its initial biomass is high in the wetland, and it can take advantage of supports under shaded conditions. When the herbaceous layer is removed at the beginning of the dry season, it is possible to significantly reduce the biomass of both climbers by the end of the dry season, relative to areas where there was no

Figure 8. The climber Ipomoea tiliacea covers the young trees, bending them. In reforestation sites, local inhabitants have to carry out maintenance to ensure tree survival and growth. (Photos by Marco González Nochebuena) Wetland Science & Practice January 2022 65

Treatment

Initial N

Control Cut Vegetation Mulch Plastic Cover

20 20 20 20

Final N) 1 7 14 19

Survival (%) 5.0 ± 5.0 c 35.0 ± 12.7 bc 70.0 ± 9.3 ab 95.0 ± 5.0 a

Final height (cm) 122.0 143.8 ± 9.58 a 161.2 ± 6.02 a 163.6 ± 1.81 a

Table 1. Survival (mean ± 1 SE), final height and relative growth rate (RGR) of P. aquatica planted saplings in four treatments: 1) Control (planting saplings); 2) Cut (herbaceous vegetation cut and saplings planted); 3) Vegetation Mulch (herbaceous vegetation cut and used as plant mulch, saplings planted); 4) Plastic Cover (herbaceous vegetation cut, plastic cover over soil, saplings planted). Different letters indicate significant differences among treatments with the Holm-Sidak test (*P ≤ 0.05, ***P ≤ 0.001, ns = non-significant at P > 0.05). Survival is indicated as the mean of survival percentage for each treatment after 24 months. In the Control only one sapling survived in one of the five blocks therefore, the control was not included in the two-way ANOVA for final height. Transformed values were used for statistical comparisons. N = number of sapling per treatment.

Treatment Control

Initial plant cover (%)

Final plant cover (%)

100

Cut

Vegetation Mulch Plastic Cover

Species with the highest cover Leersia hexandra, Ipomoea tiliacea, Ipomoea indica Leersia hexandra, Leersia oryzoides, Ludwigia octovalvis Leersia hexandra, Pistia stratiotes, Leersia oryzoides, Pistia stratiotes, Leersia hexandra, Ipomoea tiliacea

200

Trimester regrowth of the herbaceous cover Never disappeared First

First

Second

Final height (cm) 140

Table 2. The time that the herbaceous layer needs to recover after each treatment.

Treatment

Activities)

Materials

Total (USD) $8.9

Control

Plant transportation, sapling planting

P. aquatica saplings

Cut

Cutting the herbaceous layer, plant transportation, sapling planting Cutting the herbaceous layer, preparing and applying plant mulch, plant transportation, sapling planting Cutting the herbaceous layer, fixing plastic cover, plant transportation, sapling planting and maintenance

P. aquatica saplings, materi- $15.9 als for plant cutting P. aquatica saplings, materi- $22.9 als for plant cutting

Vegetation Mulch Plastic Cover

P. aquatica saplings, materi- $44.2 als for plant cutting, plastic

Total/ha (USD) $1,993 $ 3,551 $5,106 $9,824

Table 3. Cost of activities and materials required for each treatment to control the herbaceous vegetation in Ciénaga del Fuerte, Veracruz, Mexico. The total cost is calculated for one hectare.

66 Wetland Science & Practice January 2022

herbaceous removal. However, if supports are available, I. tiliacea can grow very fast and its biomass accumulation can be very similar to its growth in non-herbaceous removal areas. The flood level gradient in the wetland was the factor that limited I. tiliacea growth. It seems that this species has a low flood tolerance, and its biomass in areas with high flood level areas was very low. Also, high flood levels limit re-sprouting in climbers, which delays both regeneration and growth. WHAT IS THE EFFECT OF RESTORATION ON THE JICOTEA TURTLE? Freshwater wetlands are the habitat for a high diversity of fauna such as reptiles, amphibians, and birds. In oviparous reptiles, oviposition is essential to complete their reproductive cycle. Nesting site selection is a characteristic of the female behavior that contributes to the survival and permanence of the species (Kolbe and Janzen 2002). The freshwater jicotea turtle (Trachemys venusta, Emydidae family) is a species endemic to southeastern Mexico and Central America. Due to its high nutritional, cultural, and ornamental value, it is one of the most overexploited species (Flores 2009). Its population reduction is also due to the loss and fragmentation of its habitat, and it is currently considered an at-risk species. Therefore, we wanted to see if the environment created by the dominance of Leersia hexandra was suitable for this species to nest. Six sites were selected to search for nests along transects parallel to the riverbank (Note: They are described in Vázquez-Benavides 2019). Data were collected during the dry season of 2017 and twice in 2018 (March and May in both years). Forty-four nests were found, of which 34 had been predated; only ten were active. The site with the highest number of nests was a reforestation site (started in 2013 with Pachira aquatica trees). Here the presence of L. hexandra grass was low because of tree shading. Hymenachne amplexicaule dominated, but this species does not form grass cushions that pose a problem for seed germination so tree re-establishment was unaffected in contrast to sites dominated by Leersia (Figure 9). Since the sites where L. hexandra grass dominates have conditions that make it difficult for turtles to enter and dig directly into the ground to lay their eggs, turtles selected areas with less grass cover and less creeping vegetation. Thus, planting saplings of P. aquatica favors the recovery of turtle populations by providing suitable nesting sites. COMMUNITY PARTICIPATION People in rural communities are very dependent on wetland resources and the environmental services they provide. They are frequently interested in restoration projects. Ten

years ago, we worked with local people to organize an ecotourism group to guide people through the swamp. The guides are local farmers who know the area very well and who used to fish in the river that crosses the freshwater forested wetlands, until the catch dwindled to almost nothing. Working as guides in their ecotourism project is a means of earning money and is increasing their knowledge of the ecosystems that surround them and how they function. This activity also expands their awareness of the importance of environmental services and conservation. They are therefore a natural group to support work on the restoration of their environment. They have actively participated in choosing sites, growing plants, maintaining the plantations, and keeping an eye on the whole process (Figure 10). Some of them are very observant and actively contribute to the decision-making process for everyday maintenance and restoration activities. Their help is truly invaluable. They take visitors to the restoration sites and explain the process as part of their guided tours. They are paid wages for their day’s work and this benefits them economically – a win-win for local residents and the environment. CONCLUSION AND RECOMMENDATIONS It is necessary to control Leersia hexandra and Ipomoea tiliacea to increase the survivorship of planted trees during the restoration of the Ciénaga del Fuerte swamp. Both are native wetland species but are problematic because they do not allow natural regeneration of floodplain forest, and therefore are an obstacle to restoration. The planted Pachira aquatica seedlings should be surrounded by a cleared area, allowing the plantings to grow without competition from these two species for a time long enough for saplings to become well established and eventually to outgrow the grass. So, the first step is to remove the grass cushion to give the saplings an advantage. Once established the saplings grow above the grass and are not greatly affected by it. Since the grass tolerates flooding better than the tree saplings do, the restoration site has to be checked periodically to ensure that saplings are growing sufficiently. The removal of the vines and the grass should be carried out during the weeks before the water level increases above the soil’s surface, i.e., from July to August, in order to ensure that the underground biomass of the grasses and vines remains in flooded soils to limit vegetative regeneration for as long as possible. Both will sprout when the water level decreases, while the grass can produce some sprouts under flooded conditions (Vázquez-Benavides 2019). It is important to recognize that cutting and plant removal should be avoided when the dry season begins since the low level of the water table will Wetland Science & Practice January 2022 67

Figure 9. Two sites by the river. On the left, a river border with Pachira aquatica trees and an herbaceous cover of Hymenachne amplexicaule are shown. On the right is a site dominated by Leersia hexandra, with its thick cushion of dry organic matter. (Photos by Judith Vázquez Benavides).

Figure 10. Community activities in the restoration process: lifting and removing the Leersia hexandra organic matter cushion (first four photographs), collecting young Pachira aquatica saplings from the seedling bank, growing plants in the nursery, planting saplings, visiting the sites during high flood levels, and helping set up the experiments. (Photos by Marco González Nochebuena)

favor the growth of both Leersia hexandra and Ipomoea tiliacea. Shading seems to reduce climber growth, so high sapling densities should be used, and plantings should be done in patches. Areas where Ipomoea has not invaded are preferred restoration sites. Experiments done with Annona glabra, another freshwater forested wetland species, show that planting saplings on mounds elevated 30 cm above the soil level increased survival, although the herbaceous layer also had to be removed periodically (Sánchez García 2020). These 68 Wetland Science & Practice January 2022

mounds reduce flooding time and the stress on the plants. Restoration projects should include a maintenance budget for several years until the Pachira aquatica trees have grown a trunk strong enough to not bend and thus survive. The findings reported herein are based on five years of work in Ciénega del Fuerte. More research is needed to understand the biology of the different species involved and their response to both abiotic factors (flooding) and competition. Long-term monitoring of restoration projects should be an important component

of any restoration plan. After a few years, the degree of success resulting from the elimination of highly competitive species (Leersia hexandra, Ipomoea tiliacea) and the possibility of recovering the forested wetland should be evaluated. Only through monitoring will we be able to determine project success and to improve our restoration techniques. ACKNOWLEDGMENTS We thank Roberto Monroy for his help with editing the figures. We thank B. Delfosse for revising the English. REFERENCES

Aguirre Franco, L.D. 2019. Regeneración y Crecimiento de Dos Plantas Trepadoras Herbáceas en Áreas Destinadas para la Restauración de Selva Inundable en la Planicie Costera del Centro del Golfo de México. (Master’s Thesis). Posgrado en Ciencias Biológicas, UNAM. Morelia, Mexico. Allen, B.P., R.R. Sharitz, and P.C. Goebel. 2007. Are lianas increasing in importance in temperate floodplain forests in the southeastern United States? Forest Ecology and Management 242(1): 17–23. Campos, A., M.E. Hernández, P. Moreno-Casasola, E. Cejudo Espinosa, A. Robledo, and D. Infante-Mata. 2011. Soil water retention and carbon pools in tropical forested wetlands and marshes of the Gulf of Mexico. Hydrological Sciences Journal 56(8): 1388-1406.

Moreno-Casasola, P., M.E. Hernández, and A. Campos C. 2017. Hydrology, soil carbon sequestration and water retention along a coastal wetland gradient in the Alvarado Lagoon System, Veracruz, Mexico. Journal of Coastal Research 77(10077): 104-115. Murphy, H.T., D.J. Metcalfe, T. Forest, and H. Murphy. 2016. The perfect storm: Weed invasion and intense storms in tropical forests. Austral Ecology 41: 864–874. Programa de Manejo. 2002. Programa de Manejo. Área Natural Protegida Ciénaga del Fuerte. Coordinación Estatal de Medio Ambiente, Xalapa, Mexico Ramsar. 2015. Nota Informativa 7: Estado de los Humedales del Mundo y de los Servicios que Prestan a las Personas. Convención Ramsar, Gland, Switzerland. Robledo, R.A. 2013. Análisis de los Servicios Mitigación de Impactos por Tormentas y Huracanes que proporcionan los Humedales de Ciénega del Fuerte para Tecolutla, Veracruz. (Master’s Thesis). Facultad de Ingeniería Química, Universidad Veracruzana. Xalapa, Mexico. Sánchez García, E.A. 2020. Germinación y Crecimiento de Annona glabra L. Bajo Condiciones Experimentales como Base para la Restauración de un Humedal Arbóreo en la Zona Costera del Centro de Veracruz. (Master’s Thesis). Instituto de Ecología A.C. Xalapa, Mexico. Sánchez Luna, O., T. Toledo Acevedo, H. López-Rosas, and P. MorenoCasasola. 2021. Effectiveness of restoration plantings with Pachira aquatica in swamps. Restoration Ecology e13472. http://doi.org/10.1111/ rec.13472.

César, R.G., K.D. Holl, V.J. Girão, F.N. Mello, E. Vidal, M.C. Alves, and P.H. Brancalion. 2016. Evaluating climber cutting as a strategy to restore degraded tropical forests. Biological Conservation 201: 309–313.

Schnitzer, S.A., J.W. Dalling, and W.P. Carson. 2000. The impact of lianas on tree regeneration in tropical forest canopy gaps: Evidence for an alternative pathway of gap-phase regeneration. Journal of Ecology 88(4): 655–666.

González-Marín, R.M., P. Moreno-Casasola, R. Orellana, and A. Castillo. 2012. Traditional wetland palm uses in construction and cooking in Veracruz, Gulf of Mexico. Indian Journal of Traditional Knowledge 11(3): 408-413.

Tang, Y., R. Kitching, and M. Cao. 2012. Lianas as structural parasites: A re-evaluation. Chinese Science Bulletin 57(4): 307-312.

Flores, L. 2009. Valoración y Uso de Tortugas Dulceacuícolas en la Cuenca Baja del Papaloapan, Veracruz. (Master’s Thesis). Instituto de Ecología, A.C. Xalapa, Mexico.

Infante-Mata, D., P. Moreno-Casasola, C. Madero-Vega, G. CastilloCampos, and B.G. Warner. 2011. Floristic composition and soil characteristics of tropical freshwater forested wetlands of Veracruz on the coastal plain of the Gulf of Mexico. Forest Ecology and Management 262 (8): 1514-1531. Infante-Mata, D., P. Moreno-Casasola, and C. Madero-Vega. 2012. Litterfall of tropical forested wetlands of Veracruz in the coastal floodplains of the Gulf of Mexico. Aquatic Botany 98(1): 1-11. Kolbe, J.J., and F.J. Janzen. 2002. Impact of nest-site selection on nest success and nest temperature in natural and disturbed habitats. Ecology 83(1): 269-281. Landgrave, R., and P. Moreno-Casasola. 2012. Evaluación cuantitativa de la pérdida de humedales en México. Investigación Ambiental 4 (1): 35-51.

Sjögersten, S., B. Batista de la Barruda, C. Brown, D. Boyd, H. LopezRosas, E. Hernández, M. Rincón, C. Vane, C. Moss-Hayes, J. HoyosSantillan, and P. Moreno-Casasola. 2021. Coastal wetland ecosystems deliver large carbon stocks in tropical Mexico. Geoderma 403: 115173. https://doi.org/10.1016/j.geoderma.2021.115173

Vázquez-Benavides, J. 2019. Estudio del Pasto Nativo Leersia hexandra Sw. y su Efecto en la Dispersión, Germinación y Establecimiento de Semillas y Plántulas de Pachira aquatica Aubl. y como Recurso para la Anidación de la Tortuga Iinta Trachemys venusta (Gray, 1855) en el ANP Ciénaga del Fuerte, Tecolutla, Veracruz. (Master’s Thesis). Instituto de Ecología A.C. Xalapa, Mexico. Vázquez-Benavides, J., P. Moreno-Casasola, and H. López Rosas. 2020. Effect of the grass Leersia hexandra on the dispersal, seed germination and establishment of Pachira aquatica seedlings. Freshwater Biology 65(10): 1702-1717. Vázquez-González, C., P. Moreno-Casasola, A. Juárez, N. RiveraGuzmán, R. Monroy, and I. Espejel. 2015. Trade-offs in fishery yield between wetland conservation and land conversion on the Gulf of Mexico. Ocean & Coastal Management 114: 194-203.

Marín-Muñiz, J.L., M.E. Hernández, and P. Moreno-Casasola. 2014. Comparing soil carbon sequestration in coastal freshwater wetlands with various geomorphic features and plant communities in Veracruz, Mexico. Plant and Soil 378(1): 189-203. Mitsch, W.J., and J.G. Gosselink. 2000. The value of wetlands: importance of scale and landscape setting. Ecological Economics 35(1): 25-33.

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URBAN/SUBURBAN WETLANDS

Assessment and Perceptions of the Environmental Quality of the Urban and Suburban Wetlands of Leticia (Amazonas, Colombia) María J. Arias-B 1, Jhon Ch. Donato- Rondón 2 and Santiago R. Duque 3*

INTRODUCTION The Amazon River Basin, the most extensive and main hotspot of diversity on the planet (Bertzky et al. 2013), hosts the largest terrestrial biodiversity of plants and animals. However, it is not only relevant for its ecological attributes, but also in social and cultural dynamics and, in particular, as a reference in the various responses to global change (Betts et al. 2008; Vargas et al. 2019). This had led several countries (including Colombia) to consider the Amazon Basin as a subject of law, in which the State is primarily responsible for its conservation and protection (Macías 2018; Trujillo 2013). On the other hand, the Amazon region exhibits all the ecological characteristics associated with global aquatic species (Albert et al. 2011). For example, phylogenetic and biogeographic patterns, in the case of Neotropical freshwater fishes, suggest that in this region most speciation occurred along geographic rather than ecological events (Albert and Crampton 2010). Likewise, based on vegetation types in the Colombian Amazon Basin, there is a high ecosystem richness (Rudas 2009). In the case of aquatic ecosystems, the region registers 16 out of the 89 types of wetlands (Ricaurte et al. 2019). It is worth emphasizing that Amazonian aquatic ecosystems have an evolutionary history that exceeds 23 million years (Hoorn et al. 2010). In recent times, since the arrival of the first humans some 12,000 years ago (Morcote et al. 2017), life and culture have been closely linked to these ecosystems (Santos et al. 2013). Wetlands, due to their geological, biogeographical, and ecological history (McInnes 2011; Horwitz and Finlayson 2011) are one of the most productive and diverse ecosystems in the world (Gopal and Junk 2000). This biological diversity and the ecosystem services they provide are used for diverse economic activities, Environmental Engineering, Departamento de Geociencias y Medio Ambiente, Universidad Nacional de Colombia Sede Medellin. 2 Departamento de Biología, Universidad Nacional de Colombia Sede Bogotá. jcdonator@unal.edu.co. ORCID-ID:0000-0002-3111-1067 3 Instituto Amazónico de Investigaciones (IMANI), Universidad Nacional de Colombia Sede Amazonia. srduquee@unal.edu.co. ORCIDID:0000-0003-2567-6042 * Correspondence author: Santiago R. Duque. srduquee@unal.edu.co 1

70 Wetland Science & Practice January 2022

such as fishing or aquaculture production, as well as to cover basic needs of hygiene and food (Carrillo et al. 2011; Duque et al. 2018). Consequently, wetlands are important for territorial planning, especially as a figure of environmental determinants, a norm of superior hierarchy and of mandatory compliance in land use plans (Jara 2017). (Note: This norm is stipulated specifically for the municipality of Leticia [Corpoamazonia 2014] which is the subject area for this article.) The wetlands of the Amazon in general and particularly in Leticia, have been significantly transformed by factors, such as pollution, deforestation of their watersheds and riparian vegetation, sedimentation and habitat transformation (Duque 1993; Herrera et al. 2008; Senhadji et al. 2017). According to regional environmental history, the municipality of Leticia (Amazonas, Colombia) was located in a complex network of wetlands (Hoorn et al. 2010), which due to population growth were disappearing and then used for urban development (Vergel 2008). It is therefore important to consider human populations in the environmental, cultural, and social context. Knowing people’s perception of the environment helps us understand the environment and human relationships (Calixto and Herrera 2010), as well as social participation in environmental issues (Sureda et al. 2009). Furthermore, it is essential that people understand the attributes of the most diverse biome in the world such as the Amazon forest and its associated ecosystems, in order to initiate or help support actions that lead to the conservation of its megadiversity (Ribas and Aleixo 2019). The objective of our study was to address the environmental characteristics of wetlands in the urban and suburban area of Leticia and provide insight into their use and current environmental problems. It is hoped that this information will contribute to strengthening territorial and environmental planning strategies and unleash actions to strengthen citizen participation and regional environmental education. STUDY AREA The study area of the characterized wetlands in the field and those consulted in Carrillo et al. (2011) is located between 4°1’0” S to 4°14’0” S and 70°1’0” W to 69°53’0” W, in the jurisdiction of the municipality of Leticia (Amazonas, Colombia, Figure 1). The study area of Carrillo et al. (2011) covers three micro-watersheds: Yahuarcaca, Tacana and Pichuna, while our investigation also included work in urban streams (or what are known locally as “caños”) and wetlands associated with them, such as the Calderón, Urumutú and Simón Bolívar streams that drain into the Tacana. According to the Caldas-Lang classification, the climate is warm-humid (Rangel and Luengas 1997). The aver-

age annual precipitation is 3400 mm, with a monomodal rainfall regime, where the highest rainfall occurs between November and May, and the lowest between June and October. The latter is not considered a dry season since the month with the least rainfall exceeds 100 mm (Galvis et al. 2006). The average annual temperature is 25.8°C (IDEAM 2012). The warmest months of the year occur from October to November and between February and March, when the average temperature rises to 28°C. Minimum temperatures occur in June, July, and August, with values close to 24°C (Carrillo et al. 2011), coinciding with the arú or friaje season, which refers to the arrival of a cold wind that blows through the entire eastern plain of South America from Antarctica (Duque 1993; Caraballo et al. 2014). The wetlands in the region have a seasonal hydrological behavior because their waters can vary between 10 and 12 m deep (Torres et al. 2013). In the case of the river and lagoon systems associated with the Amazon River and the Yahuarcaca stream, there are four periods: high water between April and May, falling water between June and August, low water mainly in September and rising water until March (Salcedo et al. 2012; Torres et al. 2013).

The study area includes two well-defined morphological units. The first corresponds to the Leticia-Tabatinga terrace and the other to the river floodplain, where the oscillation in the water level of the Amazon River reaches 12 m vertically and causes a horizontal expansion of the water sheet with its suspended load close to 10 km (Jaramillo et al. 2013). This behavior creates unique ecological and hydrological conditions in the wetlands of Leticia (Jaramillo et al. 2013; Arbeláez et al. 2008). In the southern region of the Colombian Amazon, seven main classes of natural vegetation cover are recognized: high forests (canopy above 25 m), medium forests (canopy between 10-25 m), low forests (canopy < 10 m), alluvial forests, shrublands, grasslands (dominated by non-graminoid herbaceous vegetation) and high savannas (dominated by graminoid herbaceous vegetation) (Rudas 2007). However, in the case of the municipality of Leticia, given the modifications of anthropic origin, that it has suffered especially in the urban area, only a few patches of vegetation remain, such as the site of the old zoo and botanical garden, the forest of the Yahuarcaca stream and its lakes, and the forest of the National University of Colombia

Figure 1. Map showing location of urban and suburban wetlands studied.

Wetland Science & Practice January 2022 71

Amazon Headquarters (van Vliet and Duque 2019). METHODS Field Stage Wetlands were sampled between May and June of 2021, which corresponds to the period when rainfall begins to decrease and, therefore, coincides with the transition between high water and falling water. Twenty-six wetlands were studied with 46 samples taken for field physicochemical parameters (littoral zone and open water) such as dissolved oxygen (DO), % oxygen saturation, total dissolved solids (TDS), pH (H+), temperature (°C) and depth (m) with a HANNA HI 98194 Multiparameter probe (Table 1). The selection of wetlands sites was made according to the ease of sampling and granted access. Data from Carrillo et al. (2011) were used for the characterization of water quality and were collected in the framework of the inter-administrative agreement between the Government of Amazonas and Corpoamazonia. Only the variables collected in the field by the project team were used, and specifically those corresponding to wetlands. In this case, 23 pieces of data from different points on the urban and suburban area of the municipality of Leticia were included. A 1:100.000 scale map of the wetlands characterized in the field and those from the work of Carrillo et al. (2011) was produced using ArcGIS software version 10.4 and Google Earth Pro (Figure 1). Laboratory Stage An analysis of the physicochemical variables of the water was performed with R Studio software using the Principal Component Analysis (PCA) method, which allowed the dimensionality (variables) to be reduced to a smaller number of transformed variables (principal components) that explain the variability of the data (Bro and Smilde 2014). Each principal component would be a linear combination of the original variables and would also be independent or uncorrelated with each other (Bro and Smilde 2014). Initially, the data were arranged in a matrix and then transformed with log (x+1). We then proceeded to run the analysis using the PCA function of the aforementioned program. The total dissolved solids (TDS) variable was discarded since it produced a high collinearity with conductivity. Perception Surveys In order to know the perception that the inhabitants of Leticia have about wetlands and their environmental problems, a synchronous survey was carried out through personal interviews of 100 people or by electronic means of people in government institutions, universities and high schools. 72 Wetland Science & Practice January 2022

These surveys asked about the definition that each inhabitant had of what a wetland was, the type of problems and economic activities with the greatest impact on wetlands in Leticia, as well as the benefits they obtained from wetlands and what actions are being taken for their conservation. Perception surveys of the inhabitants of each wetland were conducted during the field campaigns. To process the responses from the perception surveys, the coding method was used, which consists of reducing the variability of responses to a few types of answers that can be tabulated and analyzed (Rincón 2014). The categories of responses were established beforehand by making a list of these comments and their frequency according to words or phrases with the same meaning, and each grouping was assigned a code that allows its tabulation and subsequent quantitative analysis. For this purpose, a code was designed in Google Colaboratory that uses Python 3 as programming language, with which it was possible to count the frequency of words and categorize them according to the researcher’s criteria and the objectives of this study (Popping 2015). After coding the responses, a Vester Matrix was made to allow for the identification of the causes and effects of environmental problems (Bermúdez and Gómez 2001). This consists of a double-entry format where the identified problems are located, the level of causality between them is established, that is, the relationship of a problem with the others (Bermúdez and Gómez 2001) and allows the qualification of the attributes in the following way: Not a cause Indirect cause Very direct cause

0 1 2

Having established the value of causality in the matrix between all the problems, we proceeded to graph this (Figure 4): the abscissa axis is the one that represents the range of passive problems or consequence problems, and the ordinate axis represents the active problems or cause problems. The sum of the causality/effect that each problem has over the others is the value reflected in the graph. The active problems represent the problems that have an influence over the other ones, but not by others, which means they are cause problems. The passive problems are those that don’t have an important influence over the other problems, but are caused by the majority, which means they are consequence problems. The critical problems usually represent one problem which is a noticeable cause of others, and which is caused by the rest. The indifferent problems represent the problems that don’t have any effect of causality on the analyzed set and are not caused by any of

Figure 2. Box plots of conductivity, temperature, pH, and % oxygen saturation. The dark line represents the median of the data, while the shaded box is delimited by the lower quartile and the upper quartile. The small dots, as visualized in the graphs of conductivity and temperature, indicate outliers with values well beyond the normal.

the problems. Finally, the Vester matrix allows the construction of a problem tree (Figure 5) where problems are put into a hierarchy and prioritized by means of the logical decomposition of cause-effect relationship (Bermúdez and Gómez 2001). RESULTS The results from field 46 samples and Carrillo et al. (2011) are presented in Table 1, whereas the variability of the physicochemical data is presented in box plots (Figure 2). It is possible to observe atypical records and extreme cases for conductivity with values above 200 µS/cm, as in the case of the wetlands of Parque Santander, with values of 252 µS/cm and 259 µS/cm; and Calderón, with 289 µS/ cm, both in the urban area. These values contrast with the average conductivity of 48 µS/cm. The average temperature is 26.0°C, with atypical values of 29°C and 29.5°C (Figure 2) in Finca El Retiro and Lake Zapatero, respectively. The pH level is in the neutral to acidic range (Figure 2). The average oxygen saturation percentage was 30.6% and several wetlands located in the suburban zones were found to have very low oxygen concentrations, representing hypoxic and in extreme cases anoxic conditions, as is the case of the wetlands of Universidad Nacional Sede Amazonia, Parque Santader, Inravisión, Chirui lake and the streams of Simón Bolívar and Calderón. The information collected in the field through the

characterization sheets (Table 2) shows that the wetlands located in the suburban area have protection figures with respect to urban wetlands. It undoubtedly favors the protection or less impact on these systems. In any case, there are already some wetlands, especially cananguchales, that are being affected by gradually increasing the population density on their territories. Suburban wetlands are mostly associated with La Beatriz and Yahuarcaca streams and to a lesser extent with Pichuna, Tacana, “Agua Negra” and Urumutú streams. The urban wetlands are located in the micro-watershed of the Urumutú stream, the Calderón stream and the Simón Bolívar stream. Less pressure occurs on suburban wetlands, first due to lower population density, second because several of them are located in conservation areas, either in indigenous territories (“resguardos”) or in protection zones of non-governmental organizations (civil society) and third, the suburban sector is important for the leisure time of the Letician people. Figure 3 represents the Principal Component Analysis (PCA) in which axis 1, associated with conductivity, explains 33% of the variance of the data while axis 2 explains 26.1%. This axis is mainly associated with temperature. By this, we could see that in general the correlation between the variables is low, but those are the variables that can explain most of the variance of the data. The wetlands located to the left of component 1 have a higher percentage of oxygen saturation and correspond in majority to the suburban area of Leticia (orange dots), Wetland Science & Practice January 2022 73

Table 1. Physicochemical parameters of Carrillo et al. 2011 (*) and those obtained in the current study (**).

Site

Wetland

1* 2* 3* 4* 5* 6* 7* 8* 9* 10* 11* 12* 13* 14*

Q. km 11 Q. km 13 Box culvert Castañal exit of Casa Abuelo Intake site of PTAP San Sebastian pond Dump Q. Urumutú dump exit Channel Spillway dump Well source Q. Urumutú Cananguchal source Yahuarcaca “Azul” lagoon Box culvert slide km 7,5 Between Mundo Amazónico and Avaque Balneario km 8, before Balneario km 8 Balneario km 8, after Q. Arenosa (Agape) Km 11 bridge Cananguchal afeter Balcón Paisa Km 16 lake water in box coulvert Ahead of Villa Kiara Q. landfill source La Beatriz

15* 16* 17* 18* 19* 20* 21* 22* 23* 24** 25** 26** 27** 28** 29** 30** 31** 32** 33** 34**

La Julianita property (Hábitat Sur) km 16 Los Balcones El Retiro km 22 Villa Santi km 15 Bruselas Alcaldía property km 18

74 Wetland Science & Practice January 2022

% oxygen saturation 17.5 14.7 73.3 13.3 22.3 21.3 23.1 11.4 15.4 57.2 55.3 26.0 42.5 69.4

Temperature (°C) 25.5 24.5 24.5 26.4 25.6 27.0 25.6 24.9 24.6 27.2 28.4 26.5 27.0 24.5

74.9 64.2 77.1 80.1 36.3 71.2 72.2 37.1 63.9 54.4 0.9 39.2 36.4 108.3 45.4 108.5 24,.3 26.0 1.3 0.0

24.3 24.9 24.4 24.8 26.1 28.5 26.7 26.3 24,.9 25.8 25.8 26.9 26.9 29.0 27.7 26.0 26.1 26.4 25.4 25.0

TDS

7 8 49 26 39 34 46 108 13 26 52 27 21 30

5.5 6.4 7.5 6.6 6.3 6.3 7.4 6.4 7.3 4.3 7.2 8.4 7.6 6.8

27 27 27 20 14 13 5 5 8 4 10 16 4 3 4 4 15 8 4 4

6.6 6.6 6.2 6.5 6.8 7.7 5.7 8.0 5.5 5.2 5.8 5.9 5.6 5.2 5.1 5.5 5.8 5.5 4.8 4.8

Conductivity (µS/cm) 7.4 8.4 36.8 46.1 37.5 33.9 47.7 107.5 13.1 25.9 51.5 27.4 21.5 29.9

Depth (m) 0.42 0.83 0.39 0.80 1.69 0.95 0.18 0.48 1.18 0.15 0.82 0.50 0.40 0.47

26.6 26.8 27.2 20.1 14.4 12.8 5.4 5.4 7.7 7.0 19.0 31.0 8.0 5.0 7.0 8.0 24.0

1.44 0.53 0.61 0.45 0.15 0.56 1.33 1.03 0.23 0.90 0.90 1.00 1.00 1.50 1.50 1.50 0.00 0.00 2.00 2.00

8.0 8.0

Site 35** 36** 37** 38** 39** 40** 41** 42** 43** 44** 45** 46** 47** 48** 49** 50** 51** 52** 53** 54** 55** 56** 57** 58** 59** 60** 61** 62** 63** 64** 65** 66** 67** 68** 69** 70**

Wetland La Manigua reserve km 18,5 Mundo Amazónico km 7 Villa Daniela km 15 Wetland km 15 site 1 Wetland km 15 site 2 Cabildo TIWA km 6 San Sebastián cananguchal Chirui lake Yahuarcaca Zapatero lake Yahuarcaca Tacana river Bora community Salado grande of Agua Negra Calderón behind the stadium Calderón La Ceiba neighborhood Simón Bolívar on the road to the Manguaré neighborhood Caño Urumutú Wetland of the Simón Bolivar - Barrio nuevo Inravisión wetland Wetland well of Parque Santander Wetland of Tomás Cárdenas property Wetland of UN Amazonia campus northeast Wetland of UN Amazonia campus north

5.2 5.0 6.4 6.0 5.8 5.9 5.4 5.3 5.9 5.6 5.9 5.7 6.8 6.8 7.2 6.9 57 6.4 5.4 5.8 6.8 6.4 6.5 6.5 6.4

Conductivity (µS/cm) 19.0 8.0 29.0 32.0 25.0 16.0 15.0 8.0 18.0 18.0 33.0 13.0 127.0 122.0 126.0 126.0 4.0 94.0 9.0 5.0 289.0 47.0 70.0 200.0 78.0

Depth (m) 2.00 2.00 0.60 0,.60 3.00 3.00 2.00 0.50 1.50 1.50 0.20 0.20 5.50 5.50 7.00 7.00 1.50 0.27 0.27 0.27 0.20 0.77 0.77 0.77 0.00

22 20 42

6.8 6.6 6.6

44.0 40.0 84.0

0.00 0.00 0.50

25.4 25.7 26.4 26.3 28.0 27.6 25.3

29 18 126 129 18 14 44

6.1 6.4 7.0 7.0 7.1 6.8 6.3

59.0 36.0 252.0 259.0 36.0 27.0 92.0

0.00 0.00 0.00 0.00 0.00 0.00 0.21

25.5

6.8

87.0

0.35

% oxygen saturation 44.5 57.9 46.9 0.0 21.2 22.1 5.5 0.0 0.0 0.0 0.7 0.0 0.0 0.0 54.1 0.0 65.9 26.2 31.5 69.1 0.0 0.0 0.0 24.1 0.0

Temperature (°C) 26.6 26.4 24.9 25.8 26.9 26.8 24.9 25.2 25.0 25.1 25.2 25.2 26.6 26.4 29.5 26.6 25.4 27.2 25.5 25.4 26.1 26.7 25.5 26.3 25.2

48.9 54.5 0.0

TDS

8 4 15 16 16 10 7 4 9 9 17 13 62 61 63 63 2 47 5 2 145 23 35 99 39

25.9 26.1 25.2

0.0 0.0 0.0 0.0 54.0 27.6 0.0 0.0

Wetland Science & Practice January 2022 75

Figure 3. Principal Component Analysis (PCA) of physicochemical variables for the sampled wetlands (our sites plus those of Carrillo et al. 2011; see Table 1 for site names).

which means better water quality conditions. On the other hand, the urban wetlands (green dots) are the ones with highest conductivity, as is the case of Calderón stream (points 55 and 58) along with the wetland well of Parque Santander (point 66), which means an increased amount of minerals explained by domestic wastewater discharge. It can be also seen that depth variable does not explain much of the variance of the data, since its vector that represents the correlation with the other ones has a lower value (shorter). The urban wetlands that have an optimal environmental quality are located on the premises of the Institute of Radio and Television (Inravisión) and the National University of Colombia (Amazon Headquarters), which have a high degree of protection: the former for being in a private area and the latter for being categorized as environmental heritage of the University. On the other hand, human settlements were found on the margins of the streams (Calde76 Wetland Science & Practice January 2022

rón wetland) indicating that there is no compliance with minimum riparian corridor. A recent resolution of the entity responsible for the environment in the region (Corpoamazonia), resolved in 2020 that wetlands and their water rounds are environmental determinants. In other words, in the improvements and adjustments of what is called territorial planning in Colombia, wetlands must be incorporated (e.g., taken into account). The results obtained from the field analysis on the problems and economic activities coincided with the perception of the inhabitants - where urbanization (and all processes and activities that it entails) is the main problem affecting the wetlands of Leticia. The results also suggest that 91.8% of those surveyed consider that there is at least one problem in the wetlands of Leticia and more than 80% mentioned contamination by solid waste and the dumping of domestic wastewater. The perception survey results show that urbanization is

the most critical problem facing wetlands in Leticia (Figure 4). Land use changes related to urbanization pose the greatest threat to wetlands. Deforestation, filling, draining, and illegal settlements are “active problems” while contamination by solid waste and the introduction of invasive species are “passive problems” recognized by the survey participants. Finally, the indifferent problems are dumping of domestic wastewater, damming and hydraulic regulation, which represent activities that have no causal effect on the analyzed set. Also, agricultural activities (poultry, fish, and livestock) generate negative impacts (pollution and habitat transformation) which are more evident in the suburban area. The construction of the problem tree (Figure 5) allowed defining the main problem that is affecting the wetlands, corresponding to urbanization. When building a house, a road or any other artificial structure, people look to the necessity of cutting down trees, filling and draining the humid zones and not to adapt themselves to these conditions. Additionally, some social problems that occur elsewhere in the region or in the country (which we do not analyze in this article) have created some displacements to the municipality of Leticia generating illegal settlements in apparently “wasteland” areas, which later become more urban areas. Finally, as a consequence of all the processes involved in urbanization, added to an inadequate planning of the city, it can be observed that contamination by solid wastes and by domestic wastewater provide evidence of a lack of proper waste separation in Leticia. All of this ends up transforming the initial conditions of wetlands and reducing their environmental quality. The results of the characterization visits and perception surveys indicated that the main benefits obtained from wetlands are, in order of importance, food (such as fruits or seeds) and medicines from vegetation, followed by water regulation and water availability. Other uses include recreation either in the form of tourism or as local entertainment, cultural or sacred resources, and fishing. The wetlands used as examples for environmental education and scientific research are the wetlands of the National University of Colombia Amazon campus, Mundo Amazónico wetland and Hábitat Sur Natural Reserve. These in turn have protection mechanisms for their conservation. The wetland that stands out for having a cultural use is Zapatero Lake, which has a special and sacred protection for the Organization of Artisanal Fishers of the Yuhuarcaca Lakes (TIKA). Indigenous peoples when living and coexisting with nature have developed mechanisms of relationship with nature, which are necessary for their survival and behavior. Like environmental regulations, sacred themes are control mechanisms to protect nature.

DISCUSSION Since Humboldt’s time in the Amazon (19th Century) there has been talk of colored rivers (Sioli 1967), recognizing three types: white, black, and clear waters. In the southern region of the Colombian Amazon these same considerations have been taken into account, but showing some differences with Sioli’s classification, who worked more in the central Brazilian Amazon. In Colombia and in particular in the region of Leticia there are white waters type I and black waters type I, due to a geographical and geological difference that occurs with more northern sectors of the basin in Colombia where they are of type II. Thus, lakes Zapatero and Shirui, when being affected by the Amazon River, are type I whitewater wetlands with conductivities higher than 100 µS/cm while the rest of the wetlands have type I blackwater and their conductivity is always lower than this value (Duque 1997; Núñez-Avellaneda and Duque 2001). Therefore, the higher values found in some urban wetlands, such as Parque Santander and those associated to Calderón micro-basin, are due to a clear disturbance of these systems by human action. The examination of environmental variables in the urban and suburban wetlands of the municipality of Leticia reveals an axis of variation in water quality and ecological status in general. The gradient is explained by changes in conductivity and low oxygen content, in addition to habitat transformations (littoral zone and riparian vegetation), particularly in urban wetlands. It is important to consider that hypoxic conditions, explained by high rates of organic matter decomposition (Junk et al. 2012) and intense metabolic activity (Lampert and Sommer 2007; Salcedo et al. 2012), are typical of Amazonian wetlands and in broad terms of shallow tropical lakes. However, anthropogenic actions such as deforestation of the watershed and riparian vegetation (Trancoso et al. 2009), urbanization (Senhadji et al. 2017) and wastewater discharges (Cardona 2020), are influential factors of high impact that are transforming wetlands and explain the environmental axis in the ecological conditions found in the wetlands of Leticia. The results obtained coincide with those found by Carrillo et al. (2011) who concluded that the main source of wetland contamination comes from the dumping of wastewater by communities located on the banks of streams and the disposal of solid waste into water bodies. These aspects are the consequence of the urbanization of the wetlands of Leticia. This agrees with Paredes (2010) who mentions that solid waste is the obvious threat to urban wetlands. While urbanization is not a recent process in the Amazon (Heckenberger et al. 2008), urbanization processes have accelerated and have led to the transformation of wetlands Wetland Science & Practice January 2022 77

Figure 4. Results of Vester matrix for environmental problems and impacts of the wetlands in Leticia. P1: Solid waste contamination, P2: Dumping of domestic wastewater, P3: Dammed, P4: Urbanization, P5: Deforestation, P6: Filling, P7: Drainage, P8: Illegal settlements, P9: Species introduction, P10: Hydraulic regulation.

Figure 5. Problem tree of the wetlands in Leticia.

78 Wetland Science & Practice January 2022

Table 2. Location, sample date, property type, protection status, neighborhood or path, and micro-basin associated with the visited wetlands in the municipality of Leticia. *AICA: Area of International Importance for Bird Conservation. Wetland

Name

Longitude

Date (DD/ MM/YY)

Location Latitude

Zone

Property

Protection

Neighborhood/Path

Microbasin

La Julianita property km 16

-4,092060

-69,984260

4/5/2021

Suburban

Private

Natural reserve of civil society

Leticia-Tarapacá km 16

La Beatriz

Los Balcones property km 21

-4,057190

-69,995470

4/5/2021

Suburban

Private

Private property

Leticia-Tarapacá km 21

Tacana

El Retiro property km 22

-4,057550

-69,996640

4/5/2021

Suburban

Private

Private property

Leticia-Tarapacá km 22

Pichuna

Villa Santi property km 15

-4,093690

-69,983860

4/5/2021

Suburban

Private

None

Leticia-Tarapacá km 15

La Beatriz

Bruselas Alcaldía property km 18

-4,082010

-69,996050

7/5/2021

Suburban

Private

City hall property

Leticia-Tarapacá km 18

La Beatriz

La Manigua reserve km 18,5

-4,078760

-69,999580

7/5/2021

Suburban

Private

Forest reserve

Leticia-Tarapacá km 18.5

La Beatriz

Mundo Amazónico km 7

-4,144070

-69,929120

11/5/2021

Suburban

Private

Natural reserve of civil society

Leticia-Tarapacá km 7

Yahuarcaca

Villa Daniela km 15

-4,099903

-69,979869

11/5/2021

Suburban

Private

Private property

Leticia-Tarapacá km 15.2

La Beatriz

Wetland km 15 site 1

-4,097334

-69,979464

11/5/2021

Suburban

Private

None

Leticia-Tarapacá km 15.5

La Beatriz

H10

Wetland km 15 site 2 (“Ojo de agua”)

-4,097259

-69,978813

11/5/2021

Suburban

Private

Private property

Leticia-Tarapacá km 15.5

La Beatriz

H11

Indigenous Council TIWA km 6

-4,162020

-69,930034

11/5/2021

Suburban

Public

In process of constitution as an indigenous settlement

Indigenous council (km 6)

Yahuarcaca

H12

Mixed cananguchal San Sebastián

-4,176890

-69,948750

18/5/2021

Suburban

Private

None

San Sebastián, 50m from the old landfill

Urumutú

Public

AICA* and special protection area under the organization’s fishing agreements TIKA

Chirui lake of lagoon system of Yahuarcaca

Yahuarcaca and Amazonas river

Zapatero lake of lagoon system of Yahuarcaca

Yahuarcaca and Amazonas river

H13

Chirui lake Yahuarcaca

-4,175005

-69,958320

18/5/2021

Suburban

H14

Zapatero lake Yahuarcaca

-4,174060

-69,963780

18/5/2021

Suburban

Public

AICA* and special protection area under the organization’s fishing agreements TIKA

H15

Tacana river Bora community

-4,066070

-69,980347

23/5/2021

Suburban

Public

Indigenous territory

Bora community km 17

Tacana

H16

Salado Grande of Agua Negra

-4,038425

-69,950658

23/5/2021

Suburban

Public

Indigenous territory

Indigenous teeritory Ticuna Huitoto km 6-11

Agua Negra

H17

Calderón behind the stadium

-4,206667

-69,939861

28/5/2021

Urban

Public

None

José María Hernández

Calderón

H18

Calderón La Ceiba neighborhood

-4,201056

-69,941250

28/5/2021

Urban

Public

None

La Ceiba

Calderón

H19

Simón Bolívar on the road to the Manguaré neighborhood

-4,193750

-69,930444

28/5/2021

Urban

Public

None

Barrio Nuevo

Simón Bolívar

H20

Caño Urumutú

-4,190389

-69,928000

28/5/2021

Urban

Public

None

Tabatinga

Urumutú

H21

Wetland of Simón Bolivar - Barrio Nuevo

-4,194667

-69,929417

28/5/2021

Urban

Public

None

Barrio Nuevo

Simón Bolívar

H22

Inravisión wetland

-4,216654

-69,936481

1/6/2021

Urban

Public

None

Avenida Internacional

None

Wetland Science & Practice January 2022 79

Table 2 Continued. Location, sample date, property type, protection status, neighborhood or path, and micro-basin associated with the visited wetlands in the municipality of Leticia. *AICA: Area of International Importance for Bird Conservation. Wetland

Name

Longitude

Date (DD/ MM/YY)

Location Latitude

Zone

Property

Protection

Neighborhood/Path

Microbasin

H23

Wetland well of Parque Santander

-4,212647

-69,942997

1/6/2021

Urban

Public

None

Centro

None

H24

Wetland of Tomás Cárdenas property

-4,214870

-69,933270

1/6/2021

Urban

Private

None

Barrio Costa Rica

None

Public

Environmental heritage of Universidad Nacional de Colombia Sede Amazonia

Km 2 antigua vía Tarapacá

Urumutú

Public

Environmental heritage of Universidad Nacional de Colombia Sede Amazonia

Km 2 antigua vía Tarapacá

Urumutú

H25

H26

Wetland of UN Amazonia campus northeast

Wetland of UN Amazonia campus north

-4,192999

-4,191590

-69,937629

-69,939163

4/6/2021

in Leticia. The wetland well of Parque Santander located in the center of Leticia, provides an extreme example. Here high levels of conductivity and hypoxic conditions have led to eutrophication of the water supply which has been replaced by pumping water sources from the municipal aqueduct. This transformation of urban environments not only has effects on natural conditions, but also impacts environmental uses or benefits, as it modifies the supply of ecosystem services (Horwitz and Finlayson 2011). Finally, while there are several policies and regulations to protect and conserve wetlands in Colombia, these have not been successful. Therefore, it is necessary to encourage, support and finance participatory research of citizen science projects in order to make the inhabitants appropriate the value of the cultural and environmental heritage of these wetlands (Contreras 2002). Policies should not only take into account the ecological particularities but also the cultural, political and economic context of the region. CONCLUSION Regarding environmental quality, differences were found between urban and suburban wetlands in the municipality of Leticia. These differences generate a gradient in conductivity, pH, temperature, and oxygen concentration. Despite the problems facing wetlands in Leticia, particularly urban wetlands, 94.8% of respondents considered wetlands important for their well-being and 97.9% emphasized the value of wetland conservation and restoration. The findings of this research provided information on the causes of the decrease in the environmental quality of wetlands in the municipality of Leticia and will hopefully aid ongoing efforts to improve territorial planning and environmental management. ACKNOWLEDGEMENTS The authors thank to National University of Colombia 80 Wetland Science & Practice January 2022

Urban

(Bogotá and Amazonia campus) for the financial support for the project “Environmental and social appropriation of the urban and suburban wetlands of Leticia” (HERMES No. 47320). REFERENCES

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Junk, W.J., M.T.F. Piedade, J. Schöngart, and F. Wittmann. 2012. A classification of major natural habitats of Amazonian white-water river floodplains (várzeas). Wetlands Ecology and Management 20(6): 461–475. https://doi.org/10.1007/s11273-012-9268-0 Lampert, W. and U. Sommer. 2007. Limnoecology: the Ecology of Lakes and Streams. Oxford University Press. 324 pp. McInnes, R.J. 2011. Managing wetlands for multifunctional benefits. In: B.A. Le Page (ed). Wetlands: Integrating Muldisciplinary Concepts. Springer New York. pp 205-222. Macías, L.F. 2018. ¿Qué significa que la amazonia sea un sujeto de derecho? Revista Colombiana Amazónica 1(11): 103-120. Morcote -R, G., D. Mahecha, and C. Franky. 2017. Recorrido en el tiempo: 12000 años de ocupación de la Amazonia. In: Universidad Nacional de Colombia (ed.). Universidad y territorio. Bogotá: v. 5, t.1, p. 66-93. Paredes, D. 2010. Determinación de amenazas en humedales urbanos: Estudio de tres humedales de Valdivia, Chile. (Trabajo de pregado, Ingeniero en Conservación de Recursos Naturales). Universidad Austral de Chile, Valdivia, Chile. 35 p. Popping, R. 2015. Analyzing Open-ended Questions by Means of Text Analysis Procedures. Bulletin de Méthodologie Sociologique BMS 128: 23-39. http://dx.doi.org/10.1177/0759106315597389 Rangel, E. and B. Luengas. 1997. Clima y aguas del eje Apaporis Tabatinga. Pp. 49-68. En: IGAC – SINCHI – Universidad Nacional de Colombia. Zonificación ambiental para el plan modelo colombo-brasilero (eje Apaporis-Tabatinga: PAT). Editorial Linotipia Bolivar. Bogotá, Colombia. 410 pp. Ribas, C.C. and A. Aleixo. 2019. Diversity and evolution of amazonian birds: Implications for conservation and biogeography. Anais Da Academia Brasileira de Ciencias 91 (3): 1-9. https://doi.org/10.1590/00013765201920190218

Gopal, B. and W.J. Junk. 2000. Biodiversity in wetlands: an introduction. In B. Gopal, W. J. Junk, and J. A. Davis (Eds.), Biodiversity in wetlands: assessment, function and conservation. Vol. 1 (pp. 1-10). Leiden: Backhuys Publishers.

Ricaurte, L.F., J.E. Patiño, D.F.R. Zambrano, J.C. Arias, O. Acevedo, C. Aponte, and W.J. Junk. 2019. A classification system for Colombian wetlands: an essential step forward in open environmental policy-making. Wetlands 39(5): 971–990. https://doi.org/10.1007/s13157-019-01149-8

Heckenberger, M.J., J.C. Russell, C. Fausto, J.R. Toney, M.J. Schmidt, E. Pereira, B. Franchetto, and A. Kuikuro. 2008. Pre-Columbian Urbanism, Anthropogenic Landscapes, and the Future of the Amazon. Science 321: 1214 – 1217.

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Herrera, M.A., M.V. Sepúlveda, and N.J. Aguirre. 2008. Análisis sobre la aplicabilidad de las herramientas de gestión ambiental para el manejo de los humedales naturales interiores de Colombia. Gestión y ambiente 11(2): 7-20. Hoorn, C., F.P. Wesselingh, J. Hovikoski, and J. Guerrero. 2010. The development of the amazonian mega-wetland (Miocene; Brazil, Colombia, Peru, Bolivia). Amazonia, Landscape and Species Evolution: A Look into the Past 123: 123-142. Horwitz, P. and C.M. Finlayson. 2011. Wetlands as settings for human health: incorporating ecosystem services and health impact assessment into water resource management. BioScience 61(9): 678-688. IDEAM. 2012. Climatología aeronáutica. Aeródromo Alfredo Vásquez Cobo Sklt–Leticia, Colombia. ideam.gov.co/ documents/290086/75945771/SKLT/0c696ee4-8601-4039-aaa89a6e83a7a442 Jara, C.F. 2017. Los determinantes ambientales y su efecto en la planificación del territorio. Universidad Santo Tomás. Bogotá. 37 pp. Jaramillo, A.J., L.N.P. Sánchez, and J.O. Rangel. 2013. Geomorfología y estratigrafía de las formaciones cuaternarias en la región del trapecio amazónico colombiano: Geomorphology and stratigraphy of the Quaternary formations in the Colombian Amazon rain forest región. Caldasia 35(2): 429-464.

Rudas, A. 2009. Unidades ecogeográficas y su relación con la diversidad vegetal de la amazonia colombiana. Ph.D. thesis, Universidad Nacional de Colombia. Bogotá, Colombia. Rudas, A. 2007. La diversidad de la vegetación: estado actual del conocimiento. In S.L. Ruiz (Ed.). Diversidad Biológica y Cultural del Sur de la Amazonia Colombiana. Corpoamazonia, Instituto Humboldt, SINCHI, UAESPNN, Bogotá. pp. 98-102. Salcedo, M.J., S.R. Duque, L. Palma, A. Torres, D. Montenegro, N. Bahamón, L. Lagos, L.F. Alvarado, M. Gómez, and A.P. Alba. 2012. Ecología del fitoplancton y dinámica hidrológica del sistema lagunar de Yahuarcaca, Amazonas, Colombia: análisis integrado de 16 años de estudio. Mundo amazónico 3: 9-41. Santos A., E. Cassú, M. Pérez and S.R. Duque 2013. Memoria ambiental de los tikuna en los lagos de Yahuarcaca (Amazonia Colombiana). Colombia Amazónica (Instituto Sinchi) 6: 41-67. Senhadji, K., M.A. Ruiz, and J.P. Rodríguez. 2017. Estado ecológico de algunos humedales colombianos en los últimos 15 años: Una evaluación prospectiva. Colombia Forestal 20(2): 191-200. Sioli, H. 1967. Studies in Amazonian waters. Atas do Simposio a Biota Amazónica 3: 9-50. Sureda, J., M. Gili, R. Comas, L. Tudela, and E. Duce. 2009. Ciudadanía y Medio Ambiente en las Islas Baleares: el ecobarómetro como instrumento de análisis. M+A, Revista Electrónica de Medioambiente 7: 23–40. https://doi.org/10.5209/MARE.15915 Wetland Science & Practice January 2022 81

WETLAND CONSERVATION

Effective Conservation and Good Governance at the Ramsar Site Bahía Lomas, Tierra del Fuego, Chile Carmen Espoz1, Ricardo Matus1,2, Daniela Haro1, Diego Luna3 and Heraldo V. Norambuena1

INTRODUCTION Bahía Lomas, in South America, is the most southerly Ramsar site, located at the eastern end of the Magellan Strait, on the northern coast of the island of Tierra del Fuego, Chile. This marine wetland covers approximately 58,946 ha (Figure 1) and belongs to the State of Chile, while the surrounding area is privately owned. Bahía Lomas is the most critical wintering area of Red Knot (Calidris canutus rufa) and is the second most important place for Hudsonian Godwit (Limosa haemastica; Morrison and Ross 1989, Morrison et al. 2004, Niles et al. 2008). This wetland also has significant numbers of Magellanic Oystercatcher (Haematopus leucopodus), White-rumped Sandpiper (C. fuscicollis), Two-banded Plover (Charadrius falklandicus), Least Seedsnipe (Thinocorus rumicivorus), and Magellanic Plover (Pluvianellus socialis; Espoz et al. 2016). Red Knot has undergone a massive population decline over the past two decades (Harrington and Flowers 1996, Morrison and Harrington 1992, Morrison and Ross 1989, Morrison et al.

2004, Niles et al. 2008). The abundance of Red Knot has contracted to the point that virtually the entire population is now confined to Tierra del Fuego where numbers decreased from 53,232 in 1986 (Morrison and Ross 1989) to 14,800 in 2008 (Niles et al. 2008). Regarding flora, grassland ecosystems dominate the wetland, with a predominance of Poa and Festuca grasses. In some sectors towards the coastal edge of the intertidal plain, tussock, scrub, and vegetative components of annual and biannual grasses dominate (Espoz et al. 2016). Characteristic species are Sarcocornia magellanica and Atriplex vulgatissima (Figure 2). After 20 years of research and monitoring, this article aims to present the conservation and governance process that has allowed Bahía Lomas to have effective conservation and represents a management model for the conservation of shorebirds and wetlands in the southern hemisphere. BAHÍA LOMAS Bahía Lomas is the southernmost Ramsar site in Chile, located near the eastern end of the Straits of Magellan on the north shore of the main island of Tierra del Fuego. Geographically, the bay is situated between Punta Catalina on the east (52°32’05’’S-68°49’4’’W) and Punta Anegada on the west (52°27’3’’S-69°26’14’’W; Figure 1). According to the Ramsar classification, Bahía Lomas is a coastal marine wetland (Frazier 1996) that represents the Chilean coast’s most extensive tidal variation range. In South America only the existing plain in Bahía San Sebastián in Tierra del Fuego, Argentina is comparable. In Bahía Lomas, the low tide zone exceeds 7 kilometers daily, measured from the highest tide line in the direction of the sea. Consequently, this bay contains a wide area of continuous and channeled muddy plains (Morrison and Ross 1989),

Figure 1. Geographic location of the Bahía Lomas marine wetland in Tierra del Fuego, Chile. Key to colors: green = preservation zone, brown = area of conservation and particular uses, yellow = buffer zone, and red = areas of sustainable management of natural resources. 1 Centro Bahía Lomas, Facultad de Ciencias, Universidad Santo Tomás, Chile. Corresponding authors E-mail: cespoz@santotomas.cl - buteonis@gmail.com 2 Centro de Rehabilitación de Aves Leñadura, Punta Arenas, Chile. 3 WHSRN Executive Office-Manomet, Plymouth, MA, USA.

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Figure 2. Sarcocornia magellanica and Side River in Bahía Lomas marine wetland, Tierra del Fuego, Chile. (Photo by Antonio Larrea)

after which large stretches of sand predominate. The linear distance between each margin of the bay is approximately 69 km. In the austral summer season (December to March), Bahía Lomas is characterized by low temperatures (between 6° and 12° C), winds exceeding 80-90 km/h, and low rainfall, with common abrupt climatic changes. In austral winter (June-August), the climatic conditions change towards temperatures below -1 °C and mild winds that do not exceed, most of the time, 60-70 km/h. CONSERVATION ADVANCES Over time, research in Bahía Lomas has been a critical factor in producing the scientific and technical knowledge necessary to make decisions that secure the effective management and conservation of the wetland. The first conservation efforts of the Bahía Lomas marine wetland date back to 2001 with the beginning of periodic visits to the area - the first censuses of shorebirds carried out by Chilean researchers (Figure 3) in collaboration with Guy Morrison and Ken Ross (see Morrison and Ross 1989), and campaigns to capture and ring shorebirds led by Lawrence Niles and funded by the New Jersey Division of Endangered and Nongame Species from the United States. Both aerial and terrestrial surveys are currently supported by the National Wildlife Research Center of Canada and “Empresa Nacional del Petróleo” (ENAP). In 2003, an ecological monitoring program led by a team from Universidad Santo Tomás was started. The aims are to evaluate abundance and distribution of macroinvertebrates in the mudflats, the trophic ecology of Calidris canutus rufa, and the physical, chemical, and biological characteristics of the sediments and water of Bahía Lomas (Espoz et al. 2008, 2011). This monitoring program is in

effect as of today. Since sheep farming is the main land use activity in the area, it was necessary to have sheep farmers join conservation efforts as allies. This relationship has had to be constantly reinforced over time, as new owners arrive at the “estancias” near the wetland. In the year 2004, with the support of Ministry of the Environment (formerly CONAMA), this wetland received recognition as Ramsar Site – a wetland of international importance (Vilina et al. 2004). Later in 2009, the wetland was designated as a Site of Hemispheric Importance for Shorebirds (WHSRN site) for the abundance of Red Knot and Hudsonian Godwit. Moreover, in April 2020, the State of Chile declared Bahía Lomas a Nature Sanctuary, the first in the Magallanes and Chilean Antarctic Region. This last categorization required the development of a management plan and the participation of an operating committee for the conservation of Bahía Lomas. While this wetland has had a management plan in place since 2011 (see Espoz et al. 2011), it was updated and expanded in 2021. The conservation targets of the latest management plan are: 1) Red Knot, 2) Hudsonian Godwit, 3) Magellanic Plover, 4) austral shorebirds, 5) tidal flat, 6) coastal edge vegetation, 7) Side River mouth, 8) archaeological sites, and 9) cetaceans. THREATS TO THE WETLAND During the last participatory process of updating the management plan of Bahía Lomas, workshops and interviews were held with key actors in the conservation and study of the area. Ten threats to the wetland were identified: 1. Risks of spills due to oil platforms (active and inactive) in Bahía Lomas and its area of influence (Figure 4a and d). 2. The impact generated by the transit of vessels in the Strait of Magellan on Bahía Lomas and its conservation objects (Figure 4c). 3. Risks of spills from oil wells, pipelines, storage facilities, and compressors located in Bahía Lomas and its area of influence. 4. Risk of accidents involving trucks that transport hydrocarbons or dangerous substances at the crossing of the Side River with the international route CH-259.

Figure 3. A mixed flock of Red Knot (Calidris canutus rufa) and Hudsonian Godwit (Limosa haemastica) registered during the aerial census in January 2021, Bahía Lomas marine wetland, Tierra del Fuego, Chile. (Photo by Heraldo V. Norambuena)

5. Tourism not oriented towards the conservation of Bahía Lomas.

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6. Livestock practices that do not consider the declaration of Bahía Lomas as a Nature Sanctuary (Figure 4b). 7. Discharge of untreated sewage, hydrocarbons, and polluting substances to the Side River. 8. Exploration and expansion of mining activity on land and in the maritime zone of Bahía Lomas. 9. Presence of invasive alien species. 10. Development of investment projects using wind turbines (or wind farms) on the route of migratory shorebirds. Following the methodology of open standards for conservation, it was proposed that many of these threats be monitored, controlled, and managed to maintain and ideally improve the conservation status of Bahía Lomas in the next ten years. GOVERNANCE The administration of the Sanctuary is the responsibility of the Bahía Lomas Conservation and Management Center Corporation (“Centro Bahía Lomas”) under the supervision and custody of the Ministry of the Environment. The Centro Bahía Lomas is part of the Faculty of Sciences of the Universidad Santo Tomás, Chile, and its objective is research, education, and social development around the conservation of the Bahía Lomas Ramsar site (Tierra

del Fuego, Chile). It is based in the city of Punta Arenas, Magallanes and Chilean Antarctic Region. In addition to the Centro Bahía Lomas, the governance figure of the area includes an operating committee of 11 members that include: Centro Bahía Lomas, Ministry of the Environment, Provincial Government, Municipality of Primavera, Chilean Navy, “Empresa Nacional del Petróleo” (ENAP), WHSRN Executive Office-Manomet, “Centro de Rehabilitación de Aves Leñadura”, “Museo de Historia Natural de Río Seco”, Council of National Monuments, and representative of owners of territories near the Bahía Lomas Nature Sanctuary. The Bahía Lomas Nature Sanctuary (SNBL) will be under the supervision and custody of the Ministry of the Environment of Chile. The operating committee conducts semiannual meetings where the Centro Bahía Lomas presents progress on the management and monitoring plans. FUTURE CHALLENGES In the management plan period 2021-2030, it has been proposed several goals for the conservation of Bahía Lomas, among which the following stand out: 1. By 2025, the operating committee will implement 100% of the actions of the Work Plan with ENAP, including monitoring and safety programs of HC’s exploitation, operation, storage, transport activities, and other polluting substances to be followed in the SNBL and its area of influence.

Figure 4. Activities in the region of the Nature Sanctuary of Bahía Lomas: A) oil platforms, B) extensive sheep farming, C) transit of vessels in the Strait of Magellan, and D) oil platforms and shorebirds. (Photos by Antonio Larrea) 84 Wetland Science & Practice January 2022

By 2025, a General Strategic Communication Plan will be developed to strengthen the capacities and avoid accidents of ENAP staff and companies related to the exploitation, operation, storage, and transport of oil or other polluting substances in the Bahía Lomas and its area of influence.

3. By 2024, the Rescue Plan for birds affected by the oil spill in the Bahía Lomas Nature Sanctuary and its area of influence will be implemented. 4. 5.

By 2025, prepare an Action Plan to develop sustainable tourism associated with Bahía Lomas. By 2025, implement a Side River Water Quality Monitoring Program in the Sanctuary’s area of influence.

6. All these actions must ensure the population stability of the conservation targets of Bahía Lomas, maintaining population abundances like those registered in the last decade. In addition to the progress related to the management plan, we hope to hire a professional to oversee livestock issues. Also, in 2022 we should have a biological station established in Bahía Lomas, which will include facilities for researchers, and an environmental interpretation center. ACKNOWLEDGMENTS Special thanks to Dr. Rob Clay and Manomet, Inc. for funding support to carry out this work. We thank Olivia Blank for the fieldwork support and Antonio Larrea for his help with photographs. A special thanks to Stuart Mackenzie and Bird Studies Canada for their essential and continued support. We thank “Empresa Nacional del Petróleo” (ENAP) for logistical support. Special thanks to Germaynee VelaRuiz, Gabriela Garrido, Fabio Labra, and the operative committee of the Bahía Lomas Nature Sanctuary.

REFERENCES

Espoz C., R. Matus, and D. Luna-Quevedo. 2016. Bahía Lomas, Ramsar Site (Chile). In: C.M. Finlayson et al. (eds.), The Wetland Book. pp 1-7.

Espoz C., A. Ponce, R. Matus, O. Blank, N. Rozbaczylo, H. Sitters, S. Rodríguez, A. Dey, and L.J. Niles. 2008. Trophic ecology of the Red Knot Calidris canutus rufa at Bahía Lomas, Tierra del Fuego, Chile. Wader Study Group Bull. 15: 69–76. Espoz C., F. Labra, R. Matus, A. Ponce, I. Barría, B. Saavedra, A. Figueroa, and M. Rondanelli. 2011. Plan de manejo para el sitio RAMSAR Bahía Lomas. Santiago: Ministerio del Medio Ambiente/Universidad Santo Tomás/Wildlife Conservation Society. 131 pp. Frazier, S. 1996. An overview of the world’s Ramsar sites. Wetlands International Publ.29. 58 p.

Harrington, B. and C. Flowers. 1996. The Flight of the Red Knot. W.W. Norton and Company, New York. Morrison, R.I.G., and R. K. Ross. 1989. Atlas of Nearctic shorebirds on the coast of South America. Canadian Wildlife Service Special Publication, Ottawa. Morrison, R.I.G. and B.A. Harrington. 1992. The migration system of the red knot Calidris canutus rufa in the New World. Wader Study Group Bull. 64: 71–84. Morrison, R.I.G., Y. Aubry, R.W. Butler, G.W. Beyersbergen, G.M. Donaldson, C.L. Gratto-Trevor, P.W. Hicklin, V.H. Johnston, and R.K. Ross. 2001. Declines in North American shorebird populations. Wader Study Group Bull. 94: 34-38. Morrison, R.I.G., R.K. Ross and L.J. Niles. 2004. Declines in wintering populations of Red Knots in southern South America. Condor 106: 60–70. Niles, L.J., H.P. Sitters, A.D. Dey, P.W. Atkinson, A.J. Baker, K.A. Bennett, R.C. Carmona, K.E. Clark, N.A. Clark, C. Espoz, P.M. González, B.A. Harrington, D.E. Hernández, K.S. Kalasz, R.G. Lathrop, R.N. Matus, C.D.T. Minton, R.I.G. Morrison, M.K. Peck, W. Pitts, R.A. Robinson and I.L. Serrano. 2008. Status of the Red Knot (Calidris canutus rufa) in the Western Hemisphere. Studies in Avian Biology No. 36. Los Angeles, Cooper Ornithological Society. Vilina, Y., J. Gibbons, and N. Núñez. 2004. Propuesta para la inclusión de Bahía Lomas, Isla Tierra del Fuego, Chile, como Sitio Ramsar. 29 pp.

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WETLAND OUTREACH

Water and Wetlands Outreach Project in the Sierra Gorda Biosphere Reserve, Mexico Tatiana Lobato-de Magalhães1, Marinus L. Otte2, and Leticia Rocha Mier3 BACKGROUND The development of sustainable water security strategies that preserve freshwater ecosystem functions and services to reduce water-related risks to humans and conserve biodiversity and abundance of biota is urgent. Nevertheless, wetland losses and degradation rates are alarmingly high, particularly in inland wetlands that face prolonged drought (Davidson 2014; Davidson et al. 2019). Through a Fulbright Specialist Project (FSP-P006854) with a multidisciplinary and multi-partner team, we worked in the Sierra Gorda Biosphere Reserve in the State of Querétaro, in collaboration with representatives of the government (Queretaro State Environmental Secretariat and Water Commission and others), rural communities (Cuatro Palos and El Éden in Pinal de Amoles, Tepozán in Arroyo Seco, and El Pocito and Carrizal de los Durán in Jalpan de Serra), Civil Societies and Non-Government Organizations, as well as with students, scientists, and practitioners of academic institutions: North Dakota State University (NDSU), Fargo, ND, USA and the Universidad Autónoma de Querétaro (UAQ), QRO, Mexico, the host institution. The 4-week consultation in September 2021 by the Fulbright Specialist in water and wetlands, Dr. Marinus L. Otte, was key to support and improve UAQ’s ongoing efforts, particularly to develop strategies for sustainable management of water and wetlands, by improving rainwater harvesting and wastewater treatment and by creation of artificial ponds for improvements of ecology and biodiversity. Enabling capacity-building in wetland sciences is a crucial objective of our project, aligning with the Fulbright Program mission, and key for safeguarding water of good quality in a sustainable manner. The project focused on promoting and improving water and wetland management and conservation in the State of Querétaro through a three-pronged approach combining (1) aquatic science education, (2) wetland awareness, and (3) engagement of local stakeholders.

FSP Project Director, Natural Sciences Faculty, Universidad Autónoma de Querétaro, Mexico; corresponding author: tatiana.lobato@uaq.mx

THE SIERRA GORDA BIOSPHERE RESERVE The Sierra Gorda Biosphere Reserve, Querétaro State, Mexico, is situated in a predominantly semiarid environment with karstic geology at elevations from 350 to 3,160 m a.s.l. The main town in the area is Jalpan de Serra (21°13’05.2” N, 99°28’26.6” W, 760 m a.s.l.). The intermittent availability of water and the porous substrates complicate sustainable management of water (Mireles 2006). This region harbors many types of vegetation as high elevation pine-oak forests, lowland tropical forests, moist montane forests, and is home to a great diversity of cacti. The Sierra Gorda Biosphere Reserve is a very species-rich region, consisting of butterflies (800 species), birds (342), reptiles and amphibians (131), fungi (127), mammals (110), and fish (27), among others. It harbors the last remaining population of the Ara militaris (military macaw) in Mexico (Grupo Ecológico Sierra Gorda 2021). Surface water and wetlands

Fulbright Specialist, North Dakota State University, USA. Education and Communities Support Department, Universidad Autónoma de Querétaro, Mexico.

2 3

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Figure 1. A) Santa María River with high sediment load due to the rainy season. B) Altered wetland (excavated) in Landa de Matamoros, Sierra Gorda Biosphere Reserve, Mexico. (Photos by Marinus Otte)

Figure 4. Teaching in the mountains: The Project Coordinator and Fulbright Specialist showing participants how to create new wetlands in high-altitude, steeply sloped lands. (Photo by José Gómez)

Figure 2. A concrete cistern, use for rainwater harvesting from the roofs of homes in El Éden, Sierra Gorda Biosphere Reserve, Mexico. This one was leaking a bit around the faucet, but the owners did not want to repair it yet, because the wasps living by their home used that spot for drinking. (Photo by Marinus Otte)

Figure 5. Sharing course materials with people from the Cuatro Palos community. (Photo by Marinus Otte)

Figure 3. The Fulbright Specialist Dr. Marinus Otte showing instructional materials. (Photo by Tatiana Lobato de Magalhães)

in this region are scarce and increasingly threatened by land-use transformation, urban and agricultural expansion, and invasive species (Figure 1). Groundwater extraction, pollution, and unsustainable agricultural practices such as flood and drip irrigation are major concerns. The problem is further magnified because the Sierra Gorda Biosphere Reserve is an economically marginal area from which inhabitants migrate to either larger cities or the United States (Arroyo-Quiroz and Perez-Gil 2007). Scientific knowledge about how and where to restore and construct wetlands is essential to enhance sustainable management of water, and to secure ecological services and functions, such as suitable habitat for wildlife. This will also safeguard sustainable access to adequate quantities of Wetland Science & Practice January 2022 87

water of acceptable quality for human well-being and sustainable socio-economic development. In the Sierra Gorda Biosphere Reserve, four rural communities have about 100 rainwater harvesting systems as their principal water source (Cuatro Palos, El Éden, Tepozán, and El Pocito). They were constructed by the Autonomous University of Queretaro (UAQ) in collaboration with Cáritas de Querétaro I.A.P. and Peace Corps (Figure 2). In Cuatro Palos and Tepozán the rainwater harvesting complements the water they received by the pipe systems of the State Water Commission. However, these systems are interrupted very frequently. In Carrizal de los Durán the springs are the main source of water and they therefore do not have rainwater harvesting systems. Other communities of Landa de Matamoros municipality face severe water treatment issues and had specific questions about better and ecologically sound water management. Through the Linking Relations Department (Dirección de Vinculación) and the Education and Communities Support Department (Coordinación de Educación y Apoyo Comunitário, CEACOM), the UAQ already had a long-standing relationship with rural communities in the Sierra Gorda, a huge advantage that enabled us to achieve good results within the short period of the field visits. TECHNICAL APPROACH

within the Reserve. As part of the courses, local people, Dr. Otte, and UAQ faculty exchanged experiences relating to water and wetlands, focusing on the specific needs of each locality (e.g., water pollution and the potential use of constructed wetlands for water treatment, spring management and protection, habitat for wildlife, and education). All courses were open to undergraduate students from various rural campuses of UAQ located in the Sierra Gorda (Concá, Jalpan de Sierra, and Pinal de Amoles). Due to a lack of infrastructure (e.g., energy and suitable space) and the need to be in outside areas due to the COVID-19 pandemic, we created nine bilingual pamphlets (in both Spanish and English; Figures 3 and 4) covering several topics on wetlands: What is a wetland? Why are wetlands important? What do wetland plants look like? Wetlands and mosquitos. Threats to wetlands. Constructed wetlands. How to construct wetlands? Holistic management of springs. Wetlands for water management in small dwellings. We shared the pamphlets with the participants during the interactive presentations and donated all pamphlet sets to the communities along with other instructional materials (e.g., boards, pencil, markers, and paper; Figure 5). Finally, we invited participants to make drawings or written messages about the importance of water and wetlands. That activity was important to engage participants

Eight-Day Expedition to the Sierra Gorda Biosphere Reserve We carried out an eight-day expedition across the Sierra Gorda Biosphere Reserve (September 18-25, 2021), visiting four municipalities, interacting with at least 300 participants of communities, university, civil societies, and government. During this expedition we provided short courses and workshops on water and wetlands to five rural communities, visited several wetlands, water reserves, treatment systems, and water collection systems, organized two meetings with local representatives from government, university and civil society, visited two rural campuses of the UAQ, collected material for diagnostics (notes, surveys), took at least 4,000 photographs for a book, and filmed more than 14 hours of audiovisual material for a documentary, including 38 recorded interviews. In addition, one undergraduate and two graduate students (from Anthropology, Watershed Management, and Biological Sciences programs) supported the activities during the expedition, which, in turn, provided them with opportunities to connect directly with Dr. Otte and UAQ faculty members and learn from their experience and expertise. Training and Instructional Materials We organized one-day courses for at least five communities 88 Wetland Science & Practice January 2022

Figure 6. A woman from the El Pocito community expressing her feelings about water and wetlands through drawings “For me water is life.” (Photo by Emiliano Plata)

and learn about their perspectives, particularly for those who were perhaps too shy to speak up during the presentations or couldn’t read or write (Figure 6). Work-groups and Visits to Key Sites Dr. Otte, UAQ scientists, and local representatives visited some of the natural wetlands in the region to evaluate their condition as well as to identify species of interest to potentially use in restoration and in artificial wetlands. We also visited wastewater treatment systems (Figure 7) and contaminated streams. We further developed training and activities which enabled local communities to learn basic concepts of water management and wetland ecology, such as water quality, harvesting water techniques, recognition of aquatic plant species, importance and types of wetlands (i.e., the role of wetlands in water security and maintenance of biodiversity) in what is a very sensitive region, the Sierra Gorda Biosphere Reserve. Engagement of Local Stakeholders Drs. Otte and Lobato were interviewed by TV UAQ on 9 September 2021, which led a follow-up meeting with a colleague from the Querétaro Water Commission. We further organized two workshops about water and wetland management for government representatives in order to engage stakeholders in the development of water security practices and strategies. Dr. Otte shared the experiences and challenges of the Sierra Gorda communities and lessons-learned worldwide to inspire decision-makers to include practices that involve artificial wetlands and protect the remaining natural ones in future projects in this region. The first workshop was in the Sierra Gorda Biosphere Reserve (Campus Jalpan, UAQ) with the second in the city of Querétaro

Figure 7. Visiting the Jalpan de Sierra wastewater treatment system and talking about opportunities to use constructed wetlands to clean up the water in the town. (Photo by Marinus Otte)

(UAQ principal Campus and livestream transmission). The workshops consisted of three sections: Introduction/ lecture about wetlands, Experiences from Sierra Gorda, and Conclusions. They included several activities to promote exchange among the participants (cognitive maps of water security, pros and cons of using wetlands to solve environmental problems; Figure 8). Finally, we asked them to form pairs to discuss wetland and water opportunities which the institutions they represented could potentially collaborate on in the short, medium or long term. Many excellent opportunities were identified, particularly regarding education and awareness. In addition, the Director of the Protected Areas National Commission CONANP (Guadalupe Sánchez Weimann) agreed to convene an ‘umbrella’ association of stakeholders in water and wetland management in the region, and to organize a conference for all stakeholders in 2022. Documentary (Storytelling about Perceptions of Safeguarding Water) While working with the Sierra Gorda Biosphere Reserve communities, we recorded about 14 hours of audiovisual materials of the project activities, the landscape and life in the Sierra Gorda Biosphere Reserve, and 38 interviews with local people (storytelling style) (Figure 9). People shared with us their stories about water and wetlands in this fragile region, changes in water management between their childhood and present (e.g., springs and wetlands that disappeared, new methodologies to get water resources, and pollution in rivers), and key messages to safeguard water for the future. We are planning to premier the documentary in Spanish with English subtitles in December 2021.

Figure 8. A cognitive map on water security created by the Querétaro State stakeholders during the workshop at the Universidad Autónoma de Querétaro. (Photo by Marinus Otte) Wetland Science & Practice January 2022 89

documentary arising from this project will be in both Spanish and English.

Figure 9. Interviewing a woman for the documentary. She lives in Cuatro Palos, the highest part of the watershed, and talked to us about the challenges she faced with water in the past and the rainwater harvesting systems that people currently use in this region. (Photo by Marinus Otte)

Scientific Lectures Dr. Otte presented seminars based on his experience with projects in the USA, Asia, Europe and other regions, and his experience as Editor-in-Chief of WETLANDS, as follows: (1) “Sustainable solutions for environmental problems with water-loving plants” for the Botanical Society of Mexico audience (more than 2K views) as part of the monthly webinar program of the Botanical Society of Mexico on August 25, 2021, (2) “Wetlands as solutions for environmental problems” for faculty and students of the Natural Sciences, Biological Sciences and Watershed Management programs at UAQ (118 participants) on September 7, 2021, (3) “Students vs Editors — or how to get your paper published” for graduate students of the Natural Sciences, UAQ’ (30 participants) on September 14, 2021, and (4) “Wetlands” for undergraduate students of the rural campus of UAQ in Concá, Arroyo Seco (30 participants) on September 20, 2021. Language Exchange Language can be a challenge for international outreach projects, particularly when they involve local people. In this context, we planned courses molded in small parts with continuous translations for answers and questions, introductions, and technical comments. All of our instructional materials were presented in both languages, Spanish and English. We believe that we can promote further opportunities better by improving bilingual capabilities for stakeholders, not only via the direct participants, but also via the children who will benefit from the bilingual materials we offered to the local schools. In addition, the book and the 90 Wetland Science & Practice January 2022

Outreach in Pandemic Time Running a wetland education, society linkage, and awareness project during the pandemic was a challenge for us in terms of logistics and planning. We held many small workshops across the Sierra Gorda Biosphere Reserve and all activities were at outdoors locations, instead of largescale indoor gatherings. Only a small number of participants were invited for all activities, to ensure groups remained small, while social-distancing and wearing masks. Challenging though this may have been, on the other hand this meant we were able to engage more directly with individuals, which we feel made the experience more productive for all. OPPORTUNITIES FOR WATER AND WETLANDS MANAGEMENT IN THE SIERRA GORDA We identified several opportunities for this region in terms of water and wetlands management, including: Biodiversity Improvement and Restoration of ‘Bordos’ “Bordo” is a local name for a pond (usually used by cattle) that may be entirely artificial or constructed in existing natural wetlands. Most of the bordos in the State of Querétaro were temporary wetlands that were altered. Regardless, the bordos are important landscape elements that have huge potential for wetland restoration or to create new wetland zones to improve sustainable management of water and biodiversity. Treatment Wetlands Urban towns across the Sierra Gorda Biosphere Reserve would benefit greatly from incorporation of wetlands in wastewater treatment systems (at least for tertiary treatment) both in existing and future projects. Particularly in rural communities we suggest implementation of small wetlands for individual households for treatment of grey water and subsequent re-use for irrigation of flower and vegetable gardens at the property. This is crucial for sustainable water management at higher elevations of the Reserve (20003000 m) where water is scarce. Conservation of Natural Springs People in the region are well aware of the importance of conservation of natural springs, but knowledge about how to protect and conserve them is lacking. In a region with karstic geology, it is not so easy to assess what the watershed of the springs is and exactly where the water comes from. In addition, changes in land uses, such as deforesta-

tion and road construction, resulting in erosion and pollution, compromise essential water resources. Promoting Well-being, Creating Awareness, and Citizen Science Projects Wetlands are crucial for promotion of health and well-being anywhere in the world, but especially in this karstic region. Perhaps our most important role during the visits was to promote more awareness about sustainable water management and wetland conservation and protection. As in many regions, such efforts are complicated, because the majority of wetlands in the region were destroyed or severally altered in the past. The majority of the present population did not know what the term “wetland” means, nor did they have any memory of what used to be there (the problem of “shifting baselines”). Perhaps most surprising is that most of the people we encountered in the city of Jalpan, the largest city in the Sierra Gorda and its economic center, did not know that the reservoir near the town, La Presa de Jalpan, is also a Ramsar site. It is an internationally important stopover and resting place for migratory birds. We observed lack of knowledge about the importance of wetlands not only in the rural communities but also among students and academics. Consequently there are many opportunities to raise awareness of wetlands and initiate dialogue about water and the environment in general. CONCLUSIONS This project aimed to advance knowledge of water and wetlands management, strengthen institutional linkages between academia, local communities, government and other stakeholders, and to support ongoing efforts to design innovative plans to harvest water, analyze water quality, and treat water using constructed wetlands. Several hundreds of people participated, and interest from rural as well as urban communities and stakeholders was very high. We plan to write a book to propose practices and techniques that people from this region could use to improve water management and wetland conservation and protection, to be published in 2022 by the UAQ. This project was key to establish a “wetland dialogue” between people in the region. We achieved some outstanding results from the contact with the diverse groups. Most notable is the realization of the need for an ‘umbrella’ as-

sociation of stakeholders in water and wetland management in the region. Finally, this Fulbright Specialist project was not a onceoff exercise but was the starting point for continued collaboration on wetland outreach, teaching, and research among both institutions: Universidad Autónoma de Querétaro and North Dakota State University. ACKNOWLEDGMENTS We thank the Fulbright Specialist Program; Dr. Mahinda Martínez, Dr. José Gómez, Dr. Mónica Queijeiro (Natural Sciences Faculty), MSc. Sofia Rivas, MSc. Paulina Becerril, Psicóloga Janete Hoyo (Dirección de Vinculación), Dr. Teresa Gasca (Rector) from the Universidad Autónoma de Querétaro for their motivation and support with logistics and fundraising; Jackal Tanelorn for his support through the US-Mexico Commission (COMEXUS); the UAQ students Lizeth Harzbecher, Miguel Sarmiento, Emiliano Plato for their support to collect data during the expedition in the Sierra Gorda Biosphere Reserve, particularly Emiliano for the photography database with ~4,000 registers; and Christian Rodriguez (UAQ Audiovisual coordination) for his genuine dedication on collecting audiovisual data for our documentary. This project was funded by the Developing Knowledge Fund (FONDEC-UAQ-2021), Fulbright Specialist Program (FSP-P006854), and Council on Science and Technology of the Querétaro State (CONCYTEQ-20101436). REFERENCES Arroyo-Quiroz, I., and R. Perez-Gil. 2007. Human-Wildlife Interactions in the Sierra Gorda Biosphere Reserve, Mexico: Annual Report Y2. Rufford Foundation-Faunam, AC Ciudad de México, Mexico. Davidson, N.C. 2014. How much wetland has the world lost? Longterm and recent trends in global wetland area. Marine and Freshwater Research 65(10): 934-941. Davidson, N.C., A.A. Van Dam, C.M. Finlayson, and R.J. McInnes. 2019. Worth of wetlands: revised global monetary values of coastal and inland wetland ecosystem services. Marine and Freshwater Research 70(8): 1189-1194. Grupo Ecológico Sierra Gorda. 2021. Available in https://sierragorda.net/ reserva-de-la-biosfera-sierra-gorda/ Visited: Sep 2021. Mireles, M.V. 2006. El mundo de la Sierra Gorda. Arqueología mexicana 13(77): 28-37.

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TREATMENT WETLANDS XXXXXX

The Pan-American Treatment Wetland Network, HUPANAM: A Brief Sketch of the Treatment Wetland Research Groups in the Latin-American and Caribbean Region M.A. Rodriguez-Dominguez1, 2, 3 *, P.H. Sezerino1, 4, and C.A. Arias 2, 3

INTRODUCTION Treatment wetlands (TW) technology is a nature-based solution where natural processes occurring in natural wetlands are emulated and optimized through engineered designs to improve water quality (Rodriguez-Dominguez et al. 2020). The technology is also known as constructed wetlands, phytoremediation systems, reed beds, soil infiltration beds, engineered wetlands root systems, biofilters, among others (Carvalho et al. 2017). The technology has been used around the world successfully for more than 60 years, treating polluted waters from different origins and across the world (Langergraber et al. 2019). TW have proved to be effective for treating rain-water run-off (Malaviya and Singh 2012; Istenič et al. 2011, 2012), and wastewater from domestic (Konnerup et al. 2009; Vymazal 2011; Wallace 2006; Brix and Arias 2005) , slaughterhouses (Gutiérrez-Sarabia et al. 2004; Soroko 2007), industrial (Vymazal 2014; Hadad et al. 2010; Maine et al. 2019) and urban sources (Pozo-Morales et al. 2017; Stefanakis 2019). TW have mostly been used for domestic wastewater treatment (DWWT) due its relative low operative and maintenance cost (Starkl et al. 2010; Uggetti et al. 2012; Mannino et al. 2008). Additionally, TW have shown ancillary benefits such as sustaining biodiversity, climate change mitigation, CO2 sequestration, flooding attenuation, enhancing public spaces, and contributing to the habitat conservation (Sanchez Celis et al. 2018). According to (Kadlec and Wallace 2008), natural wetlands have been used as wastewater discharge sites for at least 100 years, however, the history of TW is shorter, as it started in 1952 when the Max Planck Institute in West Germany published the results of the experiments regarding the use of TW for wastewater treatment. Thenceforth the Pan-American Treatment Wetland Network (HUPANAM) Department of Biology, Aarhus University, Ole Worms Allé 1, building 1135, 8000 Aarhus C, Denmark 3 Aarhus University Centre for Water Technology WATEC Aarhus University, Ny Munkegade 120, building 1521 DK-8000 Aarhus C, Denmark 4 Department of Sanitary and Environmental Engineering, Federal University of Santa Catarina, Campus Universitário, Trindade, Florianópolis, SC, CEP:88.010-970, Brazil 1 2

* Corresponding author: mard@bio.au.dk

92 Wetland Science & Practice January 2022

technology has spread around the globe, being used on all the continents, including experiments in Egypt, Morocco, Tanzania, Kenya, India, Vietnam, Taiwan, Nepal, Brazil, Colombia, Chile, Argentina, Nicaragua, Guatemala, Ecuador and Mexico (Naja and Volesky 2019). TW technology has also evolved. In 1952 the systems were characterized for not having filling media - today named Free Surface Systems (Kadlec and Wallace 2008). They evolved to the Subsurface Flow Systems, which use filling media like sand, gravel or coke to enhance treatment by increasing biofilm growth and therefore reducing the facility’s footprint. During the last decades, to enhance TW systems, intensification, such as the use of external energy sources, the input of air, or the use of advanced filling media, has allowed TWs to reach higher pollutant removal rates or to eliminate specific compounds from polluted waters (Fonder and Headley 2013). In the Latin-America and Caribbean (LAC) region, while TWs have been studied during the last two decades, only since 2010 have TW researchers, decision makers and practitioners of the region started to meet. The aim was to meet every two years in different participant countries to share and promote TW technology, to disseminate local results and to update the state of the art. These meetings led to the creation of the Pan-American Conference of Wetland Systems for Wastewater Treatment (CONFPANAM), now in its 5th edition. As the conferences thrived and the number of participants increased, a formal Pan-American Treatment Wetland Network (HUPANAM) was created and now includes members of most of the Pan-American countries. This article aims to briefly describe the evolution of the TW research in the region and provide updated information on the most active TW research groups in the region and the results of their work. HISTORY OF THE PAN-AMERICAN CONFERENCE OF WETLAND SYSTEMS FOR WASTEWATER TREATMENT The CONFPANAM (Table 1) is an initiative that was led in the beginning by Carlos A. Arias, Diego Paredes, Armando Rivas and other researchers in Latin America, aiming to become the first conference to bring to Spanish-speaking researchers, a platform and forum to exchange the results of the local experiments about TW (Arias 2021).

Table 1. Resume of all the editions of the Pan-American Conference of Wetland Systems for Wastewater Treatment Edition

Place

Date

Main host

Reference

Main topics

Pereira, Risaralda, Colombia

From: 26/02/12 To: 01/03/12

UTP

(CNPML 2011)

TW-design TW-removals TW-plants Natural Wetlands Cost Wetlands TW in small communities

Morelia, Michoacan, Mexico

From: 8/06/14 To: 12/06/14

IMTA

(Rivas-Hernández TW-design and Paredes-Cuervo TW-removals 2014) TW-plants Natural Wetlands Cost Wetlands TW in small communities TW for education TW-benefits

III

Santa Fe, Santa Fe, Argentina

From: 16/05/16 To: 19/05/16

UNL

(Hernán and Maine TW-design 2016) TW-removals TW-plants Natural Wetlands Cost Wetlands TW in small communities TW for education TW-benefits Emergent pollutants

Lima, Lima, Peru

From: 15/05/18 To: 18/05/18

UNALM

(Sanchez-Celis et al. TW-design 2018) TW-removals TW-plants Natural Wetlands Cost Wetlands TW in small communities TW for education TW-benefits Emergent pollutants TW and municipal swages TW for industrial ww TW for agriculture ww TW for landfills ww TW-innovation

Florianopolis, Santa Catarina, Brazil

From: 15/05/18 To: 18/05/18

UFSC

(Sezerino and Pelis- TW-design sari 2021) TW-removals TW-plants Natural Wetlands Cost Wetlands TW in small communities TW for education TW-benefits Emergent pollutants TW and municipal swages TW for industrial ww TW for agriculture ww TW for landfills ww TW-innovation Enhanced TW

UTP – Universidad tecnológica de Pereira (Technological University of Pereira) IMTA – Instituto Mexicano de Tecnología del Agua (Mexican Institute of Water Technology) UNL – Universidad Nacional del Litoral (National University of the Littoral) UNALM – Universidad Nacional Agraria La Molina (National Agrarian University La Molina) UFSC – Universidad Federal Santa de Catarina (Federal University of Santa Catarina)

Wetland Science & Practice January 2022 93

Table 2. Local TW researching groups recognized by the HUPANAM. Country

Name

Head of the group

Reference

Argentina

Argentina Wetland Research

Maria Alejandra Maine

(Argentina Wetland Research 2019)

Brazil

Wetlands Brasil.

Pablo Heleno Sezerino

(Wetlands Brasil 2011)

Chile

Red Humedales Construidos Chile

Gladys Vidal

(GIBA 2018)

Paraguay

Wetlands Paraguay

Tomás Rodrigo López Arias

(Wetlands Paraguay 2019)

Peru

Grupo de Investigación en Agua y Saneamiento Sostenible

Rosa Miglio Toledo

(UNALM 2021)

All the conferences have had participation of experts from different countries out of the LAC region, like United States, Germany, Australia, Austria, Denmark, Spain, France, and Canada resulting in a forum to share the latest developments around the world and the advances in the region allowing knowledge and tech transfer to reach the LAC region. HISTORY OF PAN-AMERICAN TREATMENT WETLAND NETWORK (HUPANAM) The HUPANAM network started in 2010 as a small informal network mainly formed by scientists from the LAC region that previously attended international wetland conferences like WETPOL and International Water Association (IWA), and similar expert groups interested in transferring TW know-how to the LAC region, especially the most recent information and technological advances. At the third CONFPANAM in Santa Fe, Argentina, attending scientists created a first steering committee of the network, with Dr. Armando Rivas from the Mexican Institute of Water Technology (IMTA) chosen as the President. Later, during the conference in Lima, Peru, the network selected the name HUPANAM, expanded the steering committee, and established for the first time the figure of “executive member”, selecting Dr. Alejandra Maine as the

President and having executive members from Argentina, Brazil, Catalonia, Chile, Colombia, Costa Rica, Paraguay, Mexico, and Peru. Most recently, during the recent fifth edition of the conference, the committee was renovated again, selecting Dr. Pablo Sezerino as President. TW RESEARCH GROUPS IN THE LAC REGION

Since the creation of HUPANAM, local researching groups have been created in different countries across the region to help spread and share the advances in the TW research. The groups recognized by the HUPANAM are shown in the Table 2. TW RESEARCH OUTPUTS IN THE LAC REGION According to Rodriguez-Dominguez et al. (2020), the number of published articles in the LAC region regarding TW have not changed much during the last decade (Figure 1). This situation points to the need for increasing the interest and the involved actors in the region in order to increase implementation and impact of the TW technology. During this last decade, more than 520 experiences were reported in 190 publications, reporting nutrient removal rates of chemical oxygen demand (COD), total nitrogen (TN), and total phosphorus (TP) of 65-83%, 55-72%, and 30-84%, respectively. The countries with the

Figure 1. Publications on constructed wetlands (CW): a) CW publications comparison among LAC and selected countries between 2009 and 2018, and b) global tendency of scientific production related to CW from 2009 to 2018. Modified from (RodriguezDominguez et al. 2020) 94 Wetland Science & Practice January 2022

treatment, the inclusion of TWs in public policies in LAC countries can improve sanitary conditions, increase the life quality, and protect the environment (Denny 1997) as well as meeting the United Nations goals (United Nations 2015). Additionally, the activities taken by the network should reach universities and technical schools to ensure the proper training of technicians and help promote the utilization of this technology across Latin America. REFERENCES

Argentina Wetland Research. 2019. Argentina Wetland Research Facebook. https://www.facebook.com/argentinawetlandresearch/. Accessed September 13, 2021. Arias, C.A. 2021. La historia del preambulo de la Conferencia Panamericana de Sistemas de Humedales. Personal communication, September 15, 2021. Ávila, C., C. Pelissari, P.H. Sezerino, M. Sgroi, P. Roccaro, and J. García. 2017. Enhancement of total nitrogen removal through effluent recirculation and fate of PPCPs in a hybrid constructed wetland system treating urban wastewater. Science of the Total Environment 584–585: 414–425. https://doi.org/10.1016/j.scitotenv.2017.01.024

Figure 2. Number of documents related with TW found per country during the last decade. (Modified from Rodriguez-Dominguez et al. 2020)

highest number of publications in the region were Brazil, Mexico, Argentina, and Colombia (Rodriguez-Dominguez et al. 2020; Figure 2). The most used technology in the region is the horizontal subsurface flow wetlands (HSSFW), followed by free water surface (FWS) one, and vertical flow (VF) and the intensified wetlands technology. The largest reported system in the region was a HSSFW of 6,000 m2 located in Rionegro, Colombia (Silva-Bedoya et al. 2016). The largest FWS system was reported in Argentina with a surface of 2000 m2, located in Santa Fe, Argentina (Maine et al. 2006). The largest VF system was established in the city of Palhoça, Santa Catarina, Brazil with a surface of 3144 m2 receiving domestic wastewater (Ávila et al. 2017). A total of 114 different plant species have been used in TW in the LAC region in the different experiments, with the most commonly used plants being Typha domingensis, Eichhornia crassipes, and Typha latifolia, while some systems use non-wetland species like Cocos nucifera (coconut),

Carica papaya (papaya), or Aloe vera.

CONCLUSION TW technology is gaining relevance in the region, increasing its impact and interest around all the countries, and improving water quality. Due the potential of the TW technology to bring sustainable and cheaper wastewater

Brix, H., and C.A. Arias. 2005. The use of vertical flow constructed wetlands for on-site treatment of domestic wastewater: New Danish guidelines. Ecological Engineering 25(5): 491–500. https://doi.org/10.1016/j. ecoleng.2005.07.009 Carvalho, P.N., C.A. Arias, and H. Brix. 2017. Constructed wetlands for water treatment: New developments. Water 9 (6): 397. https://doi. org/10.3390/w9060397 CNPML. 2011. Conferencia Panamericana en sistemas de humedales para el manejo, tratamiento y mejoramiento de la calidad del agua. http://www.cnpml.org/conferencia-panamericana-en-sistemas-de-humedales-para-el-manejo-tratamiento-y-mejoramiento-de-la-calidad-delagua/. Accessed September 10, 2021. Denny, P. 1997. Implementation of constructed wetlands in developing countries. Water Science and Technology 35(5): 27–34. https://doi. org/10.1016/S0273-1223(97)00049-8 Fonder, N., and T. Headley. 2013. The taxonomy of treatment wetlands: A proposed classification and nomenclature system. Ecological Engineering 51: 203–211. https://doi.org/10.1016/j.ecoleng.2012.12.011 GIBA. 2018. Red de Humedales Construidos Chile. http://www.eula.cl/ giba/red-humedales-construidos-chile/. Accessed September 17, 2021. Gutiérrez-Sarabia, A., G. Fernández-Villagómez, P. Martínez-Pereda, N. Rinderknecht-Seijas, and H.M. Poggi-Varaldo. 2004. Slaughterhouse Wastewater Treatment In a Full-scale System With Constructed Wetlands. Water Environment Research 76(4): 334–343. https://doi. org/10.2175/106143004X141924 Hadad, H.R., M.M. Mufarrege, M. Pinciroli, G.A. Di Luca, and M.A. Maine. 2010. Morphological response of typha domingensis to an industrial effluent containing heavy metals in a constructed wetland. Archives of Environmental Contamination and Toxicology 58(3): 666–675. https:// doi.org/10.1007/s00244-009-9454-0 Hadad H.R., and M.A. Maine. 2016. Memorias de la III Conferencia Panamericana de Sistemas de Humedales para el Tratamiento y Mejoramiento de la calidad del Agua. Segunda Conferencia Panamericana en Sistemas de Humedales Morelia, Michoacán, México.

Wetland Science & Practice January 2022 95

Figure 3. Examples of horizontal subsurface flow wetlands (HSSFW) for water treatment. On left, a 7,500 m2 full-scale HSSFW located in Jalisco, Mexico, planted with Reed (Phragmites australis), designed for treating up to 900 m3 of industrial wastewater, constructed by Soluciones Ecológicas del Desierto S.A. de C.V. (Photo reproduced with permission of Soluciones Ecológicas del Desierto S.A. de C.V.) To the right, a full-scale HSSFW installed in Ponte de Lima, Portugal, and planted with Canna flaccida, Canna indica, Zantedeschia aethiopica, Agapanthus africanus, and Watsonia borbonica. (Photo by Cristina Calheiros)

Figure 4. Examples of vertical subsurface flow (VF) wetlands. To the left, a VF wetland installed at the Center for Biologic and Aquaculture Research of Cuemanco, Autonomous Metropolitan University, Xochimilco, Mexico City, Mexico. (Photo courtesy of the designer Victor Manuel Luna Pabello.) To the right, a lab-scale vertical flow wetland (VF) installed at the Military University of Nueva Granada, Bogota, Colombia. (Photo by Tatiana Rodríguez Chaparro)

Figure 5. Two full-scale free water surface (FWS) wetlands, both located in Santa Fe, Santa Fe, Argentina. (Photos by Hernan Hadad) 96 Wetland Science & Practice January 2022

Istenič, D., C.A. Arias, V. Matamoros, J. Vollertsen, and H. Brix. 2011. Elimination and accumulation of polycyclic aromatic hydrocarbons in urban stormwater wet detention ponds. Water Science and Technology 64(4): 818–825. https://doi.org/10.2166/wst.2011.525 Kadlec, R., and S. Wallace. 2008. Treatment Wetlands 2nd Ed. CRC Press, Boca Raton, FL. Konnerup, D., T. Koottatep, and H. Brix. 2009. Treatment of domestic wastewater in tropical, subsurface flow constructed wetlands planted with Canna and Heliconia. Ecological Engineering 35(2): 248–257. https://doi.org/10.1016/j.ecoleng.2008.04.018 Langergraber, G., G. Dotro, J. Nivala, A. Rizzo, and O.R. Stein. 2019. Wetland Technology Practical Information on the Design and Application of Treatment Wetlands. IWA Publishing Alliance house. https:// www.iwapublishing.com/books/9781789060164/wetland-technologypractical-information-design-and-application-treatment Maine, M.A., G.C. Sanchez, H.R. Hadad, S. E. Caffaratti, M.C. Pedro, M.M. Mufarrege, and G.A. Di Luca. 2019. Hybrid constructed wetlands for the treatment of wastewater from a fertilizer manufacturing plant: Microcosms and field scale experiments. Science of the Total Environment 650: 297–302. https://doi.org/10.1016/j.scitotenv.2018.09.044 Maine, M.A., N. Suñe, H. Hadad, G. Sánchez, and C. Bonetto. 2006. Nutrient and metal removal in a constructed wetland for wastewater treatment from a metallurgic industry. Ecological Engineering 26(4): 341–347. https://doi.org/10.1016/j.ecoleng.2005.12.004 Malaviya, P., and A. Singh. 2012. Constructed wetlands for management of urban stormwater runoff. Critical Reviews in Environmental Science and Technology 42(20): 2153–2214. https://doi.org/10.1080/10643389.2 011.574107 Mannino, I., D. Franco, E. Piccioni, L. Favero, E. Mattiuzzo, and G. Zanetto. 2008. A cost-effectiveness analysis of seminatural wetlands and activated sludge wastewater-treatment systems. Environmental Management 41(1): 118–129. https://doi.org/10.1007/s00267-007-9001-6 Naja, G.M., and B. Volesky. 2019. Constructed wetlands for water treatment. In Murray Moo-Young (editor). Comprehensive Biotechnology. Third edition. pp. 307–322. https://doi.org/10.1016/B978-0-444-640468.00362-1 Pozo-Morales, L., C. Moron, D. Garvi, and J. Lebrato. 2017. Ecologically designed sanitary Sewer based on constructed wetlands technology - Case study in Managua (Nicaragua). Journal of Green Engineering 7(3): 421–450. https://doi.org/10.13052/jge1904-4720.735 Rivas-Hernández, A., and D. Paredes-Cuervo. 2014. Sistemas de humedales para el manejo, tratamiento y mejoramiento de la calidad del agua. Segunda Conferencia Panamericana En Sistemas de Humedales Morelia, Michoacán, México. pp. 1–189. https://www.imta.gob.mx/biblioteca/ libros_html/sistemas-de-humedales/files/assets/common/downloads/ publication.pdf. Accessed September 17, 2021. Rodriguez-Dominguez, M.A., D. Konnerup, H. Brix, and C.A. Arias. 2020. Constructed wetlands in Latin America and the Caribbean: A review of experiences during the last decade. Water 12(6): 1744. https:// doi.org/10.3390/w12061744

Sezerino, P.H., and C. Pelissari, 2021. Caderno da Conferência Panamericana de Sistemas Wetlands para Tratamento e Melhoramento da Qualidade da Água e do5o Simpósio Brasileiro sobre Aplicação de Wetlands Construídos no Tratamento de Águas Residuárias. Caderno Da Conferência Panamericana de Sistemas Wetlands Para Tratamento e Melhoramento Da Qualidade Da Água e Do5o Simpósio Brasileiro Sobre Aplicação de Wetlands Construídos No Tratamento de Águas Residuárias, 1–282. Silva-Bedoya, L.M., M.S. Sánchez-Pinzón, G.E. Cadavid-Restrepo, and C.X. Moreno-Herrera. 2016. Bacterial community analysis of an industrial wastewater treatment plant in Colombia with screening for lipiddegrading microorganisms. Microbiological Research 192: 313–325. https://doi.org/10.1016/j.micres.2016.08.006 Soroko, M. 2007. Treatment of wastewater from small slaughterhouse in hybrid constructed wetlands systems. Ecohydrology and Hydrobiology 7(3–4): 339–343. https://doi.org/10.1016/S1642-3593(07)70117-9 Starkl, M., L. Essl, J.L. Martinez, and E. Lopez. 2010. Water quality improvements through constructed wetlands: a case study in Mexico. Proceedings of the 3rd IASTED African Conference on Water Resource Management, AfricaWRM 2010. 10.2316/P.2010.686-078. https://doi. org/10.2316/p.2010.686-078 Stefanakis, A.I. 2019. The role of constructed wetlands as green infrastructure for sustainable urban water management. Sustainability 11(24): 6981. https://doi.org/10.3390/su11246981 Uggetti, E., I. Ferrer, C. Arias, H. Brix, and J. García. 2012. Carbon footprint of sludge treatment reed beds. Ecological Engineering 44: 298–302. https://doi.org/10.1016/j.ecoleng.2012.04.020 UNALM. 2021. Grupo de investigacion en aqua y saneamiento sostenible. https://web.lamolina.edu.pe/investigacion/index.php/grupos/ grupo-de-investigacion-en-agua-y-saneamiento-sostenible/. Accessed September 13, 2021. United Nations. 2015. Sustainable Development Goals. https://sdgs. un.org/goals. Accessed September 13, 2021. Vymazal, J. 2011. Constructed wetlands for wastewater treatment: five decades of experience. Environmental Science and Technology 45(1): 61–69. https://doi.org/10.1021/es101403q Vymazal, J. 2014. Constructed wetlands for treatment of industrial wastewaters: a review. Ecological Engineering 73: 724–751. https://doi. org/10.1016/j.ecoleng.2014.09.034 Wallace, S. 2006. Feasibility, Design Criteria, and O&M Requirements for Small Scale Constructed Wetland Wastewater Treatment Systems. IWA Report WERP, Volume 5. https://doi.org/10.2166/9781780403991 Wetlands Brasil. 2011. Wetlands Brasil. https://gesad.ufsc.br/apresenta caowetlandsbrasil/?fbclid=IwAR2b-6VUXLVKYx6qCElJUlurZR0Or3lO0ZSXwPPw5VDwt6FJbNA3C1CZUNg. Accessed September 17, 2021. Wetlands Paraguay. 2019. Wetlands Paraguay. https://www.facebook. com/478803742890432/posts/590226105081528/. Accessed September 21, 2021.

Sanchez-Celis, G., R.M. Miglio-Toledo, R. Vela-Cardich, and E.A. Cadillo de la Torre. 2018. Memorias IV Conferencia Panamericana de Sistemas de Humedales para el tratamiento y el mejoramiento de la calidad del agua. Universidad Agraria La Molina. http://www.lamolina. edu.xn--pe-d6t.

Wetland Science & Practice January 2022 97

WETLAND OF DISTINCTION

Cuatrociénegas Wetlands (Coahuila de Zaragoza, Mexico) Julie E. Nieset1, Wetland Ecologist, Illinois Natural History Survey, Prairie Research Institute at University of Illinois Urbana-Champaign The Cuatrociénegas wetland complex has been described as a “sea in the desert” (Espinosa et al. 2005). The wetland complex consists of around 500 water bodies approximately 840 km2 in size within the arid Chihuahuan Desert in the mountainous Northern Mexican Highlands (Figures 1 and 2), truly an oasis in the dry landscape. Mountains surround Cuatrociénegas which is essentially a bowl in the terrain that does not drain to the sea. Springs feed this area and are connected via systems of channels where the water from a deep aquifer is being continuously circulated from magmatic heat. Thus, even though this region receives less than 200 mm of precipitation per year the streams and rivers as well as ponds and lakes here are groundwater-fed (Figure 3). Scientists have discerned that the water at Cuatrociénegas exhibits similar marine osmolarity and properties as Precambrian oceans such as unbalanced nutrient stoichiometry that allow microbial mats and stromatolites to flourish (Souza et al. 2018; Figure 4). During rains the mountains that surround Cuatrociénegas release gypsum into the basin. Gypsum is a common mineral but is a rarity in the form of sand. Gypsum rich sand dunes and outcrops here form one of only three such dune fields found in all the Americas. The result of these specialized habitats and isolation is high biological diversity and unique species that are found nowhere else in the world (Souza et al. 2012). Álvarez and Ojeda (2019) note that there are at least 38 endemic species in this wetland complex. Eight endemic fish species (Minckley 1984) have been described. A survey of crustacean diversity revealed 45 species representing 4 classes, 18 families, and 32 genera and seven endemic species - three copepods, two amphipods, one isopod, and one shrimp (Álvarez and Villalobos 2019). There are 7 amphibian and 46 reptile species, of which 2 amphibian and 9 reptiles are endemic to Cuatrociénagas (García-Vázquez et al. 2019). Ochoterena and others (2020) report 297 species in 60 families of vascular plants growing on gypsum outcrops, of which 15 are endemic.

Author contact: jenieset@illinois.edu

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Figure 1. The basin of the Cuatrociénegas wetland complex (green symbol) is surrounded by mountains. It is located in northern Mexico, about 265 km west of Laredo, Texas. (Source: Imagery taken from the Wetlands of Distinction webpage utilizing Map data @2022 INEGI Imagery @ 2022 TerraMetrics)

Figure 2. The Cuatrociénegas wetland complex is a unique landscape containing numerous endemic species of plants and animals. (Photo by Tatiana Lobato-de Magalhães)

Figure 3. Springs feed water bodies such as this pond that are connected through channel systems to a deep aquifer. (Photo by Tatiana Lobato-de Magalhães)

REFERENCES

Álvarez, F., and M. Ojeda. 2019. The fauna of the Cuatro Ciénegas Basin, a unique assemblage of species, habitats, and evolutionary histories. Animal Diversity and Biogeography of the Cuatro Ciénegas Basin. pp 1-10. https://link.springer.com/chapter/10.1007/978-3-030-11262-2_1 Álvarez, F., and J. Villalobos. 2019. Crustaceans from the Cuatro Ciénegas Basin: diversity, origin, and endemism. Animal Diversity and Biogeography of the Cuatro Ciénegas Basin. pp. 77-90. https://link.springer. com/chapter/10.1007/978-3-030-11262-2_6 Espinosa, L.A. Escalante, L. Eguirte, and V. Soauza. 2005. El mar en el Desierto y su importancia para la conservación. CONABIO. Biodiversitas 58: 7-11.

Figure 4. Unique geological conditions that favor diverse microbial communities are found at Cuatrociénegas. (Photo by Tatiana Lobato-de Magalhães)

Cuatrociénagas is internationally recognized by UNESCO (as Cuatrociénegas Biosphere Reserve) as well as a wetland of international importance under the RAMSAR treaty (as Área de Protección de Flora y Fauna Cuatrociénegas). In 2006 U.S. and Mexican officials signed a Joint Declaration of Sister Park Partnerships to encourage collaboration between parks. As a result, the White Sands National Park and Cuatrociénagas were paired due to both being located in the Chihuahuan Desert and having the unique gypsum dune fields in an arid environment. Human activities threaten the Cuatrociénegas wetland complex in numerous ways. Industrial mining and extraction of gypsum and minerals from the dune fields destroy habitat. Agricultural activities create irrigation channels that pull aquifer water from the area, while overgrazing and burning for livestock harms vegetation. The SWS Wetlands of Distinction Program seeks to raise public awareness of wetlands and their benefits to human health and well-being and to the environment. While most of the designated wetlands are in the United States, recently the Program was expanded to include wetlands from other countries. The Cuatrociénegas wetland complex was the first Latin American wetland to join the program and currently is the only wetland from the region on the list. Consequently we are seeking nominations of other Latin America wetlands. Contact information, details about the nomination process, and a searchable map of current Wetlands of Distinction can be found on our website https:// www.sws.org/wetlands-of-distinction/. We look forward to adding more Latin American wetlands to the Program in the future.

García-Vázquez, U., M. Trujano-Ortega, A. Contreras-Arquieta, O. Ávalos-Hernández Omar, O. Escobedo-Correa, and P. Corcuera. 2019. Diversity of amphibians and reptiles in the Cuatro Ciénegas Basin. Animal Diversity and Biogeography of the Cuatro Ciénegas Basin. pp. 175188. https://link.springer.com/chapter/10.1007/978-3-030-11262-2_13 Minckley, W. L. 1984. Cuatro Cienegas fishes: research review and a local test of diversity versus habitat size. Journal of the Arizona-Nevada Academy of Science 19(1): 13–21. http://www.jstor.org/stable/40024286. Ochoterena, H., H. Flores-Olvera, C. Gómez-Hinostrosa, and M. Moore. 2020. Gypsum and plant species: a marvel of Cuatro Ciénegas and the Chihuahuan Desert. Plant Diversity and Ecology in the Chihuahuan Desert: 129-165. https://link.springer.com/chapter/10.1007/978-3-030-44963-6_9 Souza, V., J. Siefert, A. Escalante, J. Elser, and L. Eguiarte. 2012. The Cuatro Ciénegas Basin in Coahuila, Mexica: an astrobiological Precambrian park. Astrobiology 12(7): 641-647. Accessed November 15, 2021. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3426885/ Souza, V., A. Moreno-Letilier, M. Travisano, L. Alcaraz, G. Olmedo, and L. Eguiarte. 2018a. The lost world of Cuarto Ciénegas, a relictual bacterial niche in a desert oasis. eLIFE 2018; 7:e38278.

Wetland Science & Practice January 2022 99

NOTES XXXXXXFROM THE FIELD

or this issue, we received a quite a large number of images from numerous wetlanders and others with an interest in Latin American wetlands. From that group the following pictures were selected to show the beauty and variety of the region’s wetlands and some of their characteristic wildlife. Wetlands range from mangroves, salt marshes, and lagoon types along the coast to swamps and tropical rainforests further inland to high-elevation wetlands that include saline wetlands in deserts, fens fed from melting snow and springs in the mountains, and riparian wetlands along streams. Latin America and the Caribbean support about 60% of the world’s biodiversity including thousands of endemic species. Ten percent of the world’s biodiversity can be found in the Amazon Basin alone. Information on the status and threats to the region’s natural resources can be found in a 2016 United Nations Environment Programme report (https://www.cbd. int/gbo/gbo4/outlook-grulac-en.pdf ). Countries have joined together with the World Resources Institute and others to support natural resource conservation across the region – the 20x20 Initiative (https://initiative20x20.org/restoring-latinamericas-landscapes). Conservation efforts coming from an international treaty – the Ramsar Convention – have so far designated 212 sites as “wetlands of international importance” (a listing of these Ramsar sites with links to their profiles can be found at https://rsis.ramsar.org/ris-search?page=5&solrsort=ramsarid%20asc&pagetab=1&f%5B0%5D=regionCou ntry_en_ss%3ALatin%20America%20and%20the%20Caribbean).

Wetlands of Latin America Puerto Rico (See also cover image)

Mangrove swamp, San Juan Bay Estuary, Bosque Estatal de Piñones, Puerto Rico. (Photo by Doel Vázquez)

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Urban development encroaching on estuarine wetland, Ciénga Las Cucharillas Natural Reserve, Cataño, Puerto Rico. (Photo courtesy of Wilmer Rivera and Pedro Carrión)

Puerto Rico

Dwarf mangrove (Rizophora mangle) swamp, Ceiba, Puerto Rico. (Photo by Ricardo Colón)

Riverine wetlands, Añasco, Puerto Rico. (Photo by Luis VillanuevaCubero)

Freshwater forested wetland dominated by Pterocarpus officinalis, Humacao Natural Reserve, Humacao, Puerto Rico. (Photo by Doel Delgado)

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Mexico

High-elevation wetland in El Salado Lagoon (Laguna el Salado) during wet season, State of Puebla, Mexico. (Photo by Gerardo Sanchez Vigil; courtesy of PARES AC - Paisajes y Personas Resilientes, Asociación Civil)

High-elevation wetland in Laguna Totolcingo during the dry season, Puebla, Mexico. (Photo by Gerardo Sanchez Vigil)

Willows (Salix) in riparian forest, Panales, Tolimán, Querétaro, Mexico. (Photo by Olga Lidia Gómez-Nucamendi)

Temporary wetland characterized by Nymphoides fallax (yellow flower) and Sagittaria demersa (white flower) at 2,600 m a.s.l., Querétaro, Mexico. (Photo by Tatiana Lobato de Magalhães)

Highland freshwater wetland dominated by Nymphoides fallax, at 2,600 m a.s.l. Querétaro, Mexico. 2021. (Photo by Tatiana Lobato de Magalhães)

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Mexico

Streamside riparian wetland along Extorax Stream, El Salado, Peñamiller, Querétaro, Mexico. (Photo by Mahinda Martínez)

Nymphaea mexicana in a channel at Pátzcuaro Lake, Michoacán, Mexico. (Photo by Tatiana Lobato de Magalhães)

Nymphaea odorata bed in Zirahuén Lake, Michoacán, Mexico, with Schoenoplectus californicus marsh in back. (Photo by Tatiana Lobato de Magalhães) Lakeside wetland at sunrise in El Castillo, Xalapa Veracruz, Mexico. (Photo by Gerardo Sanchez Vigil) Wetland Science & Practice January 2022 103

Peru

The wetlands of Ite, Peru are coastal (estuarine) wetlands characterized by a variety of halophytes and they are an important site for waterfowl and waterbirds. (Photo by Jhonson Vizcarra)

High-altitude freshwater wetlands in Peru’s Andes; dominant vegetation is a rush (Distichia muscoides) that forms cushion-like mounds: a) Bofedales de Chilota and b) Bofedales de Pasto Grande. (Photos by Jhonson K. Vizcarra)

Two high Andean desert wetlands in Peru: a) Laguna Camaña and b) Laguna Toro Bravo. (Photos by Jhonson K. Vizcarra)

104 Wetland Science & Practice January 2022

Peru

Los Pantanos de Villa Ramsar site, a coastal saltmarsh in Lima, Peru. (Photo by Hector Aponte)

Emergent wetland in Lake Titicaca, a Ramsar site in Puno, Peru. (Photo by Hector Aponte)

Totora wetland in Puno Bay, part of Lake Titicaca, a wetland of international importance in Puno, Peru. (Photo by Hector Aponte)

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Chile

Wetland in Bio Bio, Chile. (Photo by Heraldo Norambuena)

Wetland along Rio Lluta, Arica, Chile. (Photo by Heraldo Norambuena)

Wetland in Rio Cruces, a Ramsar wetland of international importance, Chile. (Photo by Heraldo Norambuena)

Wetland along Rio Loa, Antofagasta, Chile. (Photo by Heraldo Norambuena)

106 Wetland Science & Practice January 2022

Argentina

A community of Schoenoplectus californicus along the shores of Pellegrini Lake, near Cinco Saltos, Río Negro Province, Argentina. (Photo by Adriel Jocou)

Shoreline wetland along Maturana lagoon, Senillosa, Neuquén Province, Argentina, with Schoenoplectus californicus in shallow water. (Photo by Adriel Jocou)

Water hyacinth (Eichornia azurea) in Lagunas y Esteros del Iberá, an Ramsar wetland of international importance in Argentina. (Photo by Sylvinia Casco)

“Mar Chiquita” lake, a Ramsar site, national park, hypersaline lake, is the home of one of the largest flamingo populations in the world (Phoenicopterus chilensis), Miramar, Córdoba, Argentina. (Photo by Hernán Hadad) Wetland Science & Practice January 2022 107

Latin American Wetland Wildlife

Argia anceps (Cerulean dancer) in copulation in Zannichellia, Las Moras, Moctezuma Stream, Cadereyta de Montes, Querétaro, Mexico. (Photo by Olga Lidia Gómez-Nucamendi)

Hetaerina vulnerata (Canyon rubyspot) in copulation, Maconí spring, Cadereyta de Montes, Querétaro, Mexico. (Photo by Olga Lidia GómezNucamendi)

Mexico has 24 species of tree frogs; this one was observed in El Oasis Spring, Peñamiller, Querétaro, Mexico. (Photo by Olga Lidia GómezNucamendi) 108 Wetland Science & Practice January 2022

Phoenicopterus chilensis (Chilean flamingo) in Mar Chiquita, a Ramsar site and national park, Miramar, Córdoba, Argentina. (Photo by Hernán Hadad)

Caiman yacaré (Alligator, Yacaré) in Jaaukanigas, another Ramsar wetland of international importance, in Villa Ocampo, Santa Fe, Argentina. (Photo by Hernán Hadad)

Busarellus nigricollis (Black-collared hawk, Aguilucho pampa) from Jaaukanigas, Argentina. (Photo by Hernán Hadad)

Campephilus leucopogon (Cream-backed woodpecker, Carpintero lomo blanco) from Jaaukanigas, Argentina. (Photo by Hernán Hadad)

Tigrisoma lineatum (Rufescent tiger-heron, Hocó colorado) observed in Middle Parana River floodplain, Santa Fe, Argentina. (Photos by Hernán Hadad)

Wetland Science & Practice January 2022 109

Calidris melanotos (Pectoral sandpiper, Playerito pectoral) observed in Middle Parana River floodplain, Santa Fe, Argentina. (Photos by Hernán Hadad)

Jacana jacana (Wattled jacana, Gallito de agua) observed in Middle Parana River floodplain, Santa Fe, Argentina. (Photos by Hernán Hadad)

Phrynops hilarii (Water turtle, Tortuga de agua) observed in Middle Parana River floodplain, Santa Fe, Argentina. (Photos by Hernán Hadad)

Egretta thula (Snowy egret) observed in Peru. (Photo by Andrea Olivera Jara)

110 Wetland Science & Practice January 2022

Leucophaeus pipixcan (Franklin’s gulls foreground), Phoenicopterus chilensis (Chilean flamingo adult with young birds; middle), and Podiceps major (Great grebe near marsh) from the Ite wetland (Humedales de Ite) at the mouth of the Locumba River, Peru. (Photo by Jhonson Vizcarra) Ardea alba (Great egret) preening in wooded wetland in Veracruz, Mexico. (Photo by Gerardo Sanchez Vigil)

Hydrochoerus hydrochaeris (Capybara, Carpincho), with young observed in Esteros del Iberá, a Ramsar wetland of international importance in Colonia Carlos Pellegrini, Corrientes, Argentina. (Photo by Hernán Hadad)

Pantara onca (Jaguar) from the banks of the Paraguay River. (From the April 2021 issue of WSP; photo by Geraldo Alves Damasceno Junior)

Wetland Science & Practice January 2022 111

Lama vicugna (Vicuña) is a wild South American camelid that can be found around high-altitude wetlands searching for drinking water and plants to feed on; here along Laguna Camaña (a lagoon in the middle of the high Andean desert), Peru. (Photo by Jhonson K. Vizcarra)

And we’ll wrap this section up with a sunset over Argentina’s Paraná River by Sylvina Casco.

112 Wetland Science & Practice January 2022

WETLANDS XXXXXXX IN THE NEWS

isted below are some links to some random news articles that may be of interest. Links from past issues can be accessed on the SWS website news page. The Association of State Wetland Managers' website contains a section entitled “Wetland News Digest.” This section includes links to newspaper articles that should be of interest. Members are encouraged to send links to articles about wetlands in their local area. Please send the links to WSP Editor at ralphtiner83@gmail.com and reference “Wetlands in the News” in the subject box. Thanks for your cooperation.

Inside the effort to save a tiny, endangered turtle in western Mass. Endangered Sea Turtles Found Washed up and Stranded on the Oregon Coast

Manatees are starving in Florida. Wildlife agencies are scrambling to save them Haryana plans wetland tag for Najafgarh lake

As seas rise, NJ’s wetlands disappear

Survey reveals submerged ancient ruins

‘We do need to back away from the coast,’ climate scientist warns

Newly-confirmed Arctic record 100-degree heat in Siberia setting off climate change "alarm bells," U.N. says

HIGHER GROUND: RETREAT - This RI neighborhood will soon be underwater forever. Can it be saved?

Wetland Research and Education Center / Atelier Z+

Big Cypress National Preserve: Six ways to experience the Everglades

Report: Feds signal plans to downlist Whooping Crane

How the coastal mid-Atlantic is haunted by sea-level rise

After years of debate, there's a new plan for these hulking warehouses near Louisiana wetlands

Rising sea levels threaten the lives and livelihood of those on a fragile U.S. coast In Iraq's famed marshlands, climate change is upending a way of life Alligator Alley: Take a hike in Florida panther country The Everglades: America's Wetland Cromwell residents urge wetlands agency to reject warehouse proposal

Mumbai's Panje wetland brims with life again; villagers net huge fishes, crabs Life in marine driftwood: The case of driftwood specialist talitrids One farmer's seaweed discovery could help slow methane emissions — and change the world

ENCOUNTER THE EVERGLADES IN THE PALM BEACHES

Massive wetland restoration project in the works for Upper Klamath Lake

The Big Cypress Swamp - Friends of the Everglades

Denman conservation charity buys 80 acres of forest and wetlands

Potential cost of saving U.S. coastal town from rising sea levels enormous: study

NSW to add two wetlands to national parks under 33,000ha land purchase

Audubon, Currituck Sound Coalition announce conservation plan for Currituck marshes

When Louisiana's trees vanish, so do its wetlands. Here's how some are trying bring them back.

‘State of the Slough’ report shows how Everglades fared during summer months

Trees Are Biggest Methane “Vents” in Wetland Areas – Significant Emissions Even When They’re Dry

Tropical forests recover after deforestation

Non-flooded riparian Amazon trees are a regionally significant methane source

Trees are biggest methane 'vents' in wetland areas – even when they're dry

The Secret Islands of the Everglades - PBS Terra (YouTube)

Victoria's 'extinct' purple-spotted gudgeon finds a home after farmers build wetland during lockdown

Abandoned cranberry bogs in Yarmouth slated for wetland restoration by state

Dragonflies disappearing as wetlands are lost

As Wetland Habitats Disappear, Dragonflies and Damselflies Are Threatened With Extinction

Haiderpur wetland in Uttar Pradesh recognised as Ramsar Site Britain’s wetlands could play vital role in fight against climate change As drought deepens, a city looks to restore dry riverbed into flowing river Dragonflies threatened as wetlands around the world disappear - IUCN Red List Wetland “Fen” In NW Iowa Revives, Home To Some Of State’s Rarest Plant Life Concept for wetland wins design award

Western Kentucky Wetland Monitoring Study - The Nature Conservancy New megadams threaten world’s biggest fish - National Geographic Florida has lost 44% of its wetlands since 1845. What is the environmental impact? Birds, frogs and sunset walks: how a wetlands project transformed the NSW town of Goulburn Tasmania's saltmarshes gain attention as councils look to preserve wetlands, filter runoff water

Wetland proposed for Micheldever to create 'nitrate credits'

Estuaries, Salt Marshes & Mangroves - MarineBio

From great chaos to great restoration: ‘Qinling Mountains defense battle’ reflects China's efforts to protect its ecological environment

Loxahatchee Wildlife Refuge: Window to the Everglades

Your Everglades moment of zen: The secret life of alligators

Hauz Khas, Sanjay lakes on list of 10 water bodies that may soon be notified as wetlands Wetland Science & Practice January 2022 113

WETLAND BOOKSHELF

here are no new books to add to this listing. Please help us add new books. If your agency, organization, or institution has a website where wetland information can be accessed, please send the information to the Editor of Wetland Science & Practice at ralphtiner83@gmail.com. Your cooperation is appreciated.

BOOKS • History of Wetland Science: A Perspective from Wetland Leaders

• Black Swan Lake – Life of a Wetland

• An Introduction to the Aquatic Insects of North

• Coastal Wetlands of the World: Geology, Ecology,

America (5th Edition)

Distributionand Applications

• Wading Right In: Discovering the Nature of Wetlands

• Florida’s Wetlands

• Sedges of Maine

• Mid-Atlantic Freshwater Wetlands: Science,

• Sedges and Rushes of Minnesota • Wetland & Stream Rapid Assessments: Development, Validation, and Application • Eager: The Surprising Secret Life of Beavers and Why They Matter • Wetland Indicators – A Guide to Wetland Formation, Identification, Delineation, Classification, and Mapping • Wetland Soils: Genesis, Hydrology, Landscapes, and Classification • Creating and Restoring Wetlands: From Theory to Practice • Salt Marsh Secrets. Who uncovered them and how?

Management,Policy, and Practice • The Atchafalaya River Basin: History and Ecology of an American Wetland • Tidal Wetlands Primer: An Introduction to their Ecology, Natural History, Status and Conservation • Wetland Landscape Characterization: Practical Tools, Methods, and Approaches for Landscape Ecology • Wetland Techniques (3 volumes) • Wildflowers and Other Plants of Iowa Wetlands • Wetland Restoration: A Handbook for New Zealand Freshwater Systems

• Remote Sensing of Wetlands: Applications and Advances.

• Wetland Ecosystems

• Wetlands (5th Edition).

• Constructed Wetlands and Sustainable Development

114 Wetland Science & Practice January 2022

WETLANDS JOURNAL

What's New in the SWS Journal - WETLANDS? The following articles appear in Volume 42, issue 1 of WETLANDS, Journal of the Society for Weland Scientists Zoning Seagrass Protection in Lap An Lagoon, Vietnam Using a Novel Integrated Framework for Sustainable Coastal Management Importance of Wetlands in Maintaining the Richness of Morocco’s Vascular Flora Vegetation Composition and Carbon Dioxide Fluxes on Rewetted Milled Peatlands — Comparison with Undisturbed Bogs Mercury distribution in an Upper St. Lawrence River wetland dominated by cattail (Typha angustifolia) Helminths of the Eurasian Coot Fulica atra at Lake Ladoga Coast (Northwestern Russia) Exploring the Influence of Multidimensional Tourist Satisfaction on Preferences for Wetland Ecotourism: a Case Study in Zhalong National Nature Reserve, China Locating Drainage Tiles at a Wetland Restoration Site within the Oak Openings Region of Ohio, United States Using UAV and Land Based Geophysical Techniques Influence of Selected Environmental Parameters on Rove Beetle (Coleoptera: Staphylinidae) Communities in Central European Floodplain Forests Changes of Community Structure and Functional Feeding Groups of Benthic Macrofauna After Mangrove Afforestation in a Subtropical Intertidal Zone, China The Peculiar Hydrology of West-Central Florida’s Sandhill Wetlands, Ponds, and Lakes—Part 1: Physical and Chemical Evidence of Connectivity to a Regional Water-Supply Aquifer Ecological Compromise: Can Alternative Beaver Management Maintain Aquatic Macroinvertebrate Biodiversity? Machine Learning Modeling Techniques for Forecasting the Trophic Level in a Restored South Mediterranean Lagoon Using Chlorophyll-a Pollution or Protection - What Early Survey Data Shows on Rapid Waterbird Utilisation of a Newly Established Sewage Treatment Plant in Urban Ghana, West Africa Effect of Water Level Fluctuation and Nitrate Concentration on Soil-Surface CO2 and CH4 Emissions from Riparian Freshwater Marsh Wetland Review of Greenhouse Gas Emissions from Rewetted Agricultural Soils Evolution of the Urban Wastewater Bio-treatment and Reuse System of East Kolkata Wetlands, India: an Appraisal Redefining Waters of the US: a Case Study from the Edge of the Okefenokee Swamp Influence of Macrophyte Complexity and Environmental Variables on Macroinvertebrate Assemblages Across a Subtropical Wetland System Sedimental Journey: Soil Fertility of Fluvial Islands Increases with Proximity to An Amazonian White-Water River A Spatially Distributed Groundwater Metric for Describing Hydrologic Changes in a Regional Population of Wetlands North of Tampa Bay, Florida, from 1990 to 2015 Understanding Bofedales as Cultural Landscapes in the Central Andes The Influence of Covid-19 on Perceived Health Effects of Wetland Parks in China Development of a Novel Fuzzy Logic-Based Wetland Health Assessment Approach for the Management of Freshwater Wetland Ecosystems Global Future Distributions of Mangrove Crabs in Response to Climate Change Water-stained leaves: formation and application as a field indicator of wetland hydrology

Wetland Science & Practice January 2022 115

WSP SUBMISSION GUIDELINES

About Wetland Science & Practice (WSP)

etland Science and Practice (WSP) is the SWS quarterly publication aimed at providing information on select SWS activities (technical committee summaries, chapter workshop overview/abstracts, and SWS-funded student activities), articles on ongoing or recently completed wetland research, restoration, or management projects, freelance articles on the general ecology and natural history of wetlands, and highlights of current events. The July issue is typically dedicated to publishing the proceedings of our annual conference. WSP also serves as an outlet for commentaries, perspectives and opinions on important developments in wetland science, theory, management and policy. Both invited and unsolicited manuscripts are reviewed by the WSP editor for suitability for publication. When deemed necessary or upon request, some articles are subject to scientific peer review. Student papers are welcomed. Please see publication guidelines herein. Electronic access to Wetland Science and Practice is included in your SWS membership. All issues published, except the current issue, are available via the internet to the general public. The current issue is only available to SWS members; it will be available to the public four months after its publication when the next issue is released (e.g., the January 2022 issue will be an open access issue in April 2022). WSP is an excellent choice to convey the results of your projects or interest in wetlands to others. Also note that as of January 2021, WSP will publish advertisements, contact info@sws. org for details. HOW YOU CAN HELP If you read something you like in WSP, or that you think someone else would find interesting, be sure to share. Share links to your Facebook, Twitter, Instagram and LinkedIn accounts. Make sure that all your SWS colleagues are checking out our recent issues, and help spread the word about SWS to non-members! Questions? Contact editor Ralph Tiner, PWS Emeritus (ralphtiner83@gmail.com).

116 Wetland Science & Practice January 2022

WSP Manuscript – General Guidelines LENGTH: Approximately 5,000 words; can be longer if necessary. STYLE: See existing articles from 2014 to more recent years available online at: https://members.sws.org/wetland-science-and-practice TEXT: Word document, 12 font, Times New Roman, singlespaced; keep tables and figures separate, although captions can be included in text. For reference citations in text use this format: (Smith 2016; Jones and Whithead 2014; Peterson et al. 2010). FIGURES: Please include full-color images of subject wetland(s). Image size should be a minimum of 1MB for this e-publication. High resolution images at 150 DPI are preferred. Figures should be original (not published elsewhere) or in the public domain. If published elsewhere, permission must be granted (author’s responsibility) from that publisher. Reference Citation Examples • Claus, S., S. Imgraben, K. Brennan, A. Carthey, B. Daly, R. Blakey, E. Turak, and N. Saintilan. 2011. Assessing the ex-tent and condition of wetlands in NSW: Supporting report A – Conceptual framework, Monitoring, evaluation and re-porting program, Technical report series, Office of Environ-ment and Heritage, Sydney, Australia. OEH 2011/0727. • Clements, F.E. 1916. Plant Succession: An Analysis of the Development of Vegetation. Carnegie Institution of Wash-ington. Washington D.C. Publication 242. • Clewell, A.F., C. Raymond, C.L. Coultas, W.M. Dennis, and J.P. Kelly. 2009. Spatially narrow wet prairies. Castanea 74: 146-159. • Colburn, E.A. 2004. Vernal Pools: Natural History and Conservation. McDonald & Woodward Publishing Company, Blacksburg, VA. • Cole, C.A. and R.P. Brooks. 2000. Patterns of wetland hydrology in the Ridge and Valley Province, Pennsylvania, USA. Wetlands 20: 438-447.

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Wetland Science & Practice January 2022 249 117 Wetland Science & Practice July 2021